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An analysis of the vegetation of Sphagnum-dominatedkettle-hole bogs in relation to environmental gradients DALE H. VITT Department of Botnny, The University ofAlberta, Edmonton, Albertn AND NANCY G. SLACK Department of Biology, Rirssell Snge College, Troy, New York Received March 22, 1974 VITT, D. H., and N. G. SLACK. 1975. An analysis of the vegetation of ~~lzcr~~nrm-dominated kettle-hole bogs in relation to environmental gradients. Can. J. Bot. 53: 332-359. Eight Sphngnirtndominated kettle-hole bogs in northern Michigan were analyzed to elucidate vegetation patterns of both vascular plants and Sphagnirm species in relation to measured bog gradients. Methods of both direct and indirect gradient analysis, including ordination and cluster analysis, were used. Community types as delineated in the ordination are discussed, including speciesdistributionfor Sphagnum and vascular plants. Segregation of community types followed gradients of pH, light, and calcium and magnesium ion concentrations. Two types of kettle-hole bogs were distinguished surrounding acid and alkaline bog lakes respectively, each with its own continuum of community types. VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation to environmental gradients. Can. J. Bot. 53: 332-359. Les auteurs ont ttudie huit tourbieres B sphaignes situees dans des creux de culot de glace fossile au nord du Michigan, dans le but d'tlucider les patrons de vegetation en relation avec les gradients mesurCs, aussl bien chez les plantes vasculaires que chez les especes de Sphagnum. Des methodes directes et indirectes d'analyse des gradients ont Cte utilisies, y compris des mtthodes d'ordination et de regroupement. Les auteurs discutent les types de communautCs rtvtlks par I'ordination, y compris la distribution des especes de Sphngnrrm et de plantes vasculaires. La stgregation des types de communautts suit les gradients du pH, de la lumiere et de la concentration des ions calcium et magntsium. Deux types de tourbieres dans les creux de culot de glace fossile ont ete distinguts, entourant respectivement les lacs de tourbiere acides et alcalins, et posstdant chacun son propre continuum de types de communautes. [Traduit par le journal] Introduction In eastern North America, bogs and fens, including those dominated by Sphagnum, have been studied by many ecologists. However, most of these vegetational analyses include vascular plants, but not Sphagnum and other bryophyte species. Successional patterns in many peatland types are often controlled by Sphagnum, which in addition to absorbing large amounts of water in areas of impeded drainage, also is able to cause changes in pH. A more acid habitat is created by exchange of hydrogen ions by Sphag- num for cations such as Ca2+, Mg2+, K+, and Na'. Thus Sphagnum species are able to influence the direction of bog and fen succession by creat- ing habitats suitable for other Sphagna and vascular plant species. The present analysis therefore includes vascular plants, Sphagnum, other mosses, hepatics, and lichens present in the bog and fen vegetation as well as physical measurements of important gradients. Eight bogs were studied in northern Michigan, situated between 45" and 47" N lat.; three in the Upper Peninsula and five in the northern portion of the Lower Peninsula. Although the analysis presented here used data only from these north- ern Michigan bogs, several bogs in Ontario, northern New York, and northeastern France were found to have similar patterns, especially of Sphagnum and Carex species. We think the results presented here may be generally applic- able, subject to plant geographical differences, to kettle-hole or basin-type bogs in glaciated regions, although not to patterned fens, blanket bogs, or raised bogs. Bog communities in the Michigan area have been studied on Isle Royale by Cooper (1913), and in northern Lower Michigan by Gates (1926, 1942) and by others, including Bevis (1960). These papers, as well as an important one by Dansereau and Segadas-Vianna (1952) on bogs of the Laurentian Shield area, use Can. J. Bot. Downloaded from www.nrcresearchpress.com by NORTH CAROLINA STATE on 12/14/14 For personal use only.

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Page 1: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

An analysis of the vegetation of Sphagnum-dominated kettle-hole bogs in relation to environmental gradients

DALE H VITT Department of Botnny The University ofAlberta Edmonton Albertn

A N D

NANCY G SLACK Department of Biology Rirssell Snge College Troy New York

Received March 22 1974

VITT D H and N G SLACK 1975 An analysis of the vegetation of ~~lzcr~~nrm-dominated kettle-hole bogs in relation to environmental gradients Can J Bot 53 332-359

Eight Sphngnirtndominated kettle-hole bogs in northern Michigan were analyzed to elucidate vegetation patterns of both vascular plants and Sphagnirm species in relation to measured bog gradients Methods of both direct and indirect gradient analysis including ordination and cluster analysis were used Community types as delineated in the ordination are discussed including speciesdistributionfor Sphagnum and vascular plants Segregation of community types followed gradients of pH light and calcium and magnesium ion concentrations Two types of kettle-hole bogs were distinguished surrounding acid and alkaline bog lakes respectively each with its own continuum of community types

VITT D H et N G SLACK 1975 An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation to environmental gradients Can J Bot 53 332-359

Les auteurs ont ttudie huit tourbieres B sphaignes situees dans des creux de culot de glace fossile au nord du Michigan dans le but dtlucider les patrons de vegetation en relation avec les gradients mesurCs aussl bien chez les plantes vasculaires que chez les especes de Sphagnum Des methodes directes et indirectes danalyse des gradients ont Cte utilisies y compris des mtthodes dordination et de regroupement Les auteurs discutent les types de communautCs rtvtlks par Iordination y compris la distribution des especes de Sphngnrrm et de plantes vasculaires La stgregation des types de communautts suit les gradients du pH de la lumiere et de la concentration des ions calcium et magntsium Deux types de tourbieres dans les creux de culot de glace fossile ont ete distinguts entourant respectivement les lacs de tourbiere acides et alcalins et posstdant chacun son propre continuum de types de communautes

[Traduit par le journal]

Introduction

In eastern North America bogs and fens including those dominated by Sphagnum have been studied by many ecologists However most of these vegetational analyses include vascular plants but not Sphagnum and other bryophyte species Successional patterns in many peatland types are often controlled by Sphagnum which in addition to absorbing large amounts of water in areas of impeded drainage also is able to cause changes in pH A more acid habitat is created by exchange of hydrogen ions by Sphag- num for cations such as Ca2+ Mg2+ K+ and Na Thus Sphagnum species are able to influence the direction of bog and fen succession by creat- ing habitats suitable for other Sphagna and vascular plant species The present analysis therefore includes vascular plants Sphagnum other mosses hepatics and lichens present in the bog and fen vegetation as well as physical measurements of important gradients

Eight bogs were studied in northern Michigan situated between 45 and 47 N lat three in the Upper Peninsula and five in the northern portion of the Lower Peninsula Although the analysis presented here used data only from these north- ern Michigan bogs several bogs in Ontario northern New York and northeastern France were found to have similar patterns especially of Sphagnum and Carex species We think the results presented here may be generally applic- able subject to plant geographical differences to kettle-hole or basin-type bogs in glaciated regions although not to patterned fens blanket bogs or raised bogs

Bog communities in the Michigan area have been studied on Isle Royale by Cooper (1913) and in northern Lower Michigan by Gates (1926 1942) and by others including Bevis (1960) These papers as well as an important one by Dansereau and Segadas-Vianna (1952) on bogs of the Laurentian Shield area use

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VITT AND SLACK BOG

descriptive or phytosociological methods largely of the Ziirich-Montpellier school and describe a variety of bog and fen communities based primarily on vascular plants Significant papers by Aldrich (1943) Catenhuisen (1950) Conway (1949) Heinselman (1963 1965 1970) Janssen (1967) Moss (1953) Nichols (1919) Osvald (1933) Sjors (1959 1963) and Transeau (1905- 1906) have added to the knowledge of peatlands in North America In Europe and Asia where peatlands have been studied intensively regional studies by Eurola (1962) Katz (1926) Kulczynski (1949) Malmer (1962) Osvald (1925 1949) Paasio (1 933) Rose (1953) Ruuhijarvi (1 960) Sjors (1948 1950) and Tolonen (1967) have greatly increased understanding of peatlands

The term bog lake in a restricted sense has been used by some authors to include only those bodies of water enclosed by ombrotrophic peat- land As compared with bogs fens would include peatlands with minerotrophic conditions Since in the area we studied truly ombrotrophic con- ditions do not appear to be present the kettle- hole lakes are strictly speaking fen lakes His- torically the peatland-surrounded basin lakes of northern Michigan have been called kettle-hole bogs or lakes We have used the term kettle-hole bog using bog in the sense of Sphagnum- dominated habitats including potentially ombro- trophic and minerotrophic conditions The bogs studied here are limited to those areas that have a Sphagnum mat at least partially surrounding a lake formed in a kettle-hole or similar depression The objectives of this study are (I) to analyze the composition structure and pattern of the vegeta- tion surrounding the open water of kettle-hole lakes (2) to determine the influence of certain physical parameters on the bog vegetation and (3) to determine plant gradient patterns and correlations of species in Sphagnunz-dominated vegetation Both gradient analysis and com- munity approaches to vegetation have been used

Any type of vegetation may be viewed as a group of communities each characterized by particular species populations vegetation struc- ture and in most cases environmental factors Such communities may or may not be separated be ecotonal areas Phytosociological methods reinforce this view and have been widely used in studies of bog vegetation on both sides of the Atlantic

Alternately one can view a type of vegetation as a continuum in which species populations and

[EGETATION ANALYSIS 333

community structure change in relation to en- vironmental gradients Although each species has its own physiological requirements differing to some extent from those of any other species groups of species with similar responses to environmental gradients may be recognized as community types Methods of vegetational analysis using this continuum viewpoint by some authors called gradient analysis have been developed by Ramensky (1930) and in North America by Whittaker (1948 1956 and later) and by Curtis and associates (195 1 and follow- ing) These techniques including ordination have been widely used in mountain vegetation and in the forests of Wisconsin and elsewhere For example ordination methods have recently been used to analyze corticolous lichen com- munities (Jesberger and Sheard 1973) marsh vegetation (Auclair et al 1973 Walker and Coupland 1968) and peatlands (Jeglum 1971 Dirschl and Coupland 1972) but they have not been used in this type of Sphagnum bog vegeta- tion We have used several of these methods including both indirect and direct gradient analysis (Whittaker 1967) In the former vegeta- tion samples are compared on the basis of species composition and arranged in an ordination along axes which may represent environmental gradi- ents In direct gradient analysis samples are arranged along actual environmental gradients Environmental gradients that may be important in bogs include light water level cation con- centration and pH all of which may in part be influenced by the Sphagnunz itself

Study Area The kettle-hole bogs sampled in this study are

located between 4520 and 4625 N latitude and 8515 and 84201 W longitude in the Great Lakes - St Lawrence forest region (Rowe 1972) The elevation of the area is about 200-220 m above sea level and does not vary to any con- siderable degree

The upland vegetation of the area is at present a mosaic of beech-maple forests and aspen woods with some areas dominated by oak white pine and other trees The lowland tree vegetation includes white cedar swamps and black spruce forests with a few areas dominated by larch and hemlock The region was extensively glaciated during Wisconsin times and the mineral soil is of glacial origin The bedrock is covered by glacial till and rock outcrops are rare Those that

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CAN J BOT VOL 53 1975

P R E C I P I T A T I O N I c m I

2 0

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FIG 1 Climograph showing mean monthly precipitation and temperature at Mackinaw City in northern Michigan (solid line) and at Willis in southeastern Michigan (broken line) Months numbered from 1 to 12 beginning with January

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are present are of limestone Gates (1912) has discussed the vegetation of the Douglas Lake region in the Lower Peninsula and Voss (1956) has given a history of the floristics of Emmet and Cheboygan Counties About 20-30 of the study region is covered by peatland vegetation most of this in the form of Tl~uja occidentalis - Picea mariana - Larix laricina swamps (Gates 1942)

Climate The climate of northern Michigan can be

considered cool temperate with mean precipita- tion of 840 mm and the mean monthly tempera- ture ranging from -7degC in February to 20degC in July and August Snowfall usually begins in October or November and may continue into April or early May averaging 1575 mm per year The absolute minimum temperature is - 39degC in February and the absolute maximum tempera- ture is 40degC in July (Gates 1942) The mean

monthly precipitation and temperature for 1970- 1971 are given in Fig 1 Temperatures peak in July and August while rainfall is highest in September In most years the maximum precipi- tation occurs in late summer whereas in southern Michigan the wettest season is spring and early summer (Fig 1) The relatively low annual precipitation and extremes of winter cold and summer heat are markedly different from the cool moist maritime climates of eg New- foundland and British Columbia

Methods Vegetation

A general reconnaissance of bog vegetation and bog distribution in the northern Michigan region was con- ducted in the summer of 1970 Based on this preliminary survey eight kettle-hole bogs were selected for study Two criteria were used in selecting the bogs for sampling (I) the presence of a Sphagnum mat and (2) the presence of open water forming a lake or pool in the center of the bog Five of the bogs are located in the Lower Peninsula

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 1

County range and township locality and stand numbers for the kettle-hole lakes in northern Michigan that were analyzed in this study

Kettle-hole lake County Range and township Stand No - -

Brown Lake Chippewa R5W T45N S6 SW-114 NE-114 8 9 10 Little Lake Mackinac R8W T44N S36 NE-114 E-118 6 26 27 Little Lake 16 (Fig 17) Cheboygan RIE T37N S16 SW-114 16 41 42 43 44 45 46 47 Livingston (Fig 15) Cheboygan R3W T36N S2 SW-114 NW-114 3 4 20 21 22 Mud Lake (Fig 18) Cheboygan R2W T37N S7 SE-114 S-118 5 7 23 24 25 28 29 30 Ryerse Lake Mackinac R7W T43N S9 SE-114 SE-114 11 12 13 31 32 33 34 35 Stutsmanville (Fig 16)

(or Hard Bottom Bog) Emmet R6W T36N S23 NE-114 SE-114 1 2 17 IS 19 Weber Lake Cheboygan R3W T34N S31 NW-114 SE-114 14 15 36 37 38 3940

FIG 2 Detail of five-county region in northern Michigan showing location of the eight kettle-hole lakes sampled Open circle is location of Mackinaw City from which precipitation and temperature were taken

in Emmet and Cheboygan Counties and three are located in the Upper Peninsula in Chippewa and Mackinac Counties The geographical coordinates for the bogs studied are given in Table 1 and the locations are shown in Fig 2

Quantitative data were gathered along transects oriented both parallel to the lake shore and perpendicular to it The parallel transects consisted of 10 quadrats while the perpendicular transects ran from open water to bog forest Within each transect I-mZ quadrats were chosen randomly from each 10-m interval Ocular esti- mates of percentage cover and frequency were recorded for each species Samples of bog water from natural depressions were taken along each transect and analyzed in the laboratory (within 6 h of sampling) with a Corning Model No 5 pH meter They were frozen and later analyzed on an atomic absorption spectrophotometer at theUniversity of Alberta for Ca2 + Na + K + and MgZ+

The data were compiled to give importance-value

indices (IV) based on a maximum of 200 by adding the relative frequency to the relative mean percentage cover Both frequency and percentage cover were calculated separately for the vascular plants and separately for Sphagnum plus other associated bryophytes and lichens

Authority names and nomenclature for the vascular plants follow Fernald (1950) Crum et 01 (1973) for the Sphagnum and mosses Schuster (1953) for the hepatics and Hale and Culberson (1970) for the lichens Voucher specimens are deposited in the herbaria of the University of Alberta (ALTA) and of Russell Sage College Troy New York

Ordination and Cluster Analysis Indirect gradient-analysis as well as direct gradient-

analysis methods were used in evaluating the data from the 300 quadrats Importance-value indices for all species of Sphagnum mosses hepatics lichens and vascular plants were used to calculate coefficients of

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336 CAN J BOT VOL 53 1975

FIG 3 Two-dimensional ordination of the 47 stands sampled Lines delimit community types S Sphagnum teres - Carex lasiocarpa - Calliergonella ctispidata community type A S clispidatlim - S papillos~rtn - Carex liniosa - C patiperctila community type C S capillacelim - Carex exilis - Vaccinitinz oxycoccos community type M S magellaniclin~ - S rectirurini- Ledtim groenlandictim - Picea mariarla community type F Zlex oerticillata- Netnopanthlis mlicronata community type T Tiirija occidentalis - S ~varnstorji community type H Tstiga canadensis - S centrale community type

similarity between stands using the formula C = [2w ( a + b ) ] x 100 where C is the coefficient of similarity rv is the sum of the lowest common importance values of the two stands and a and b are the sums of the impor- tance values of the stands A and B Species were not adjusted in relation to their maximum values of frequency and cover as in Bray and Curtis (1957) A two-dimen- sional ordination of communities was constructed from the dissimilarity values (1 - C ) and yielded a patterning for interpretation of the sampled stands (Bray and Curtis 1957 Beals 1960) A computer program was developed and all calculations were completed on the MTS and

Calcomp computer facilities at the University of Alberta The Bray-Curtis ordination was considered reliable for showing the relationships of the vegetation (see Gauch and Whittaker 1972 for a discussion)

Using the ordination the stands were grouped by visual inspection of axis 1 and 2 to reflect the spatial association of the ordination and from this grouping the mean importance values shown in Table 3 were obtained

Community types as delineated in the ordination (Fig 3 ) were named indicating the dominant vascular plant and dominant Sphagntint species as well as their most

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VITT AND SLACK BOG VEGETATION ANALYSIS 337

salient feature or distribution within the bog Bog com- munities often occur as overlapping vegetation zones and well-separated species groupings may or may not be present The associations of points of the ordination are not meant to imply that bog communities are found as distinct entities however they are presented to more easily discuss the vegetation

T o obtain another point of view of community struc- ture and to assess the validity of the community types derived from our ordination the same data as were used in the ordination were analyzed by a cluster analysis (Fig 4) Importance values of all species were used in a computer program using minimum variance clustering (agglomerative least squares (Pritchard and Anderson 1971) computer program obtained from A J B Ander- son)

Results and Discussion An analysis of the data indicates that two

distinct types of kettle-hole lakes exist in northern Michigan In the vegetation surrounding each of these two types of lakes there is a gradient of plant species from open water to bog forest zones Four Sphagnum-mat community types are described below as occurring around kettle-hole lakes in northern Michigan The outer marginal moat zone often termed a lagg or marginal fen is treated as a variable unit in which several alternative community types can be recognized The occurrence of two bog forest community types is discussed in relation to the Sphagnum- mat communities The two-dimensional ordina- tion is shown in Fig 3 illustrating the pattern of distribution of the community types The ordina- tion was constructed solely from dissimilarity values between sampled stands nevertheless the stand groupings indicate environmental gradi- ents which are considered important in bog vegetation The cluster analysis distinguishes community types similar to those of the ordina- tion while some of the differences are pointed out below

Sphagnum teres - Carex lasiocarpa - Calliergon- ella cuspidata Community Type (Alkaline Lake Edge Zone S ) (Figs 1718 27)

This community type is present only around those lakes which have a relatively high pH that is Mud Lake (pH 78) Little Lake 16 (pH 75) and Ryerse Lake (pH 73) This zone was also observed by Conway (1949) in Minnesota and referred to as the sedge mat Within this com- munity type pH values and Ca2+ and Mg2+ cation concentrations are high The mean pH is 67 the mean calcium content is 77 ppm and the mean magnesium content is 18 ppm (Table 2) There is very little microtopographic differ-

entiation and the zone is extremely wet The width of this community type varies from a few decimeters at Ryerse Lake to more than 20 m on the southeastern side of Mud Lake It does not always form a continuous perimeter around the lake often being present only on one or two sides possibly in connection with inflow or outflow channels of the lake False bottoms were present only in lakes that supported this com- munity type

Ericaceous shrubs such as Chamaedaphne calyculata and Andromeda glaucophylla are only a minor component and trees are absent except for occasional islands of Alnus rugosa There is a high diversity of vascular plant species with 10 of these entirely restricted to this zone (Table 4) Carex lasiocarpa (Fig 5) is the dominant species but Cladium mariscoides Dulichium arundina- ceum Hypericum virginicum Lysimachia terres- tris Pogonia ophioglossoides Campanula apari- noides and Potentilla palustris are also very characteristic (Fig 27)

Two species of Sphagnum S teres (Fig 5) and S subsecundum (sensu lato) are found commonly in this community type with S papillosum S magellanicum and S recurvum occasionally present in small amounts Sphagnum teres often forms a continuous mat in this zone and is characteristic of areas of high light intensity high moisture high pH and high cation con- centrations Sphagnum subsecundum although never abundant was present in almost every quadrat particularly in those that fell at the waters edge Moerkia hibernica Campylium stellatum Calliergonella cuspidata and Pellia epiphylla have high frequencies in this com- munity type although Calliergonella cuspidata is the only species that is abundant

Sphagnum cuspidatum - S papillosum - Carex limosa - C paupercula Community Type (Acidic Lake Edge Zone A) (Figs 19 28)

This community type is found at the edge of lakes having pH between 50 and 70 The zone may be less than 1 m wide or absent in a few areas as at Livingston Bog or a continuous mat 2-4 m wide as at Stutsmanville Bog The pH as well as the Ca2+ and Mg2+ concentrations is much lower than in the alkaline lake edge zone (Table 2) Like the alkaline lake edge this community type is a floating mat and hence is very wet The topography is slight and there is little shade Trees are absent although Picea and Larix

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CAN J BOT VOL 53 1975

FIG 4 Dendrogram of minimum variance cluster analysis of the 47 stands sampled Letters refer to com- munity types as designated in the ordination (Fig 3)

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VITT AND SLACK BOG VEGETATION ANALYSIS 339

TABLE 2 Mean pH and cation concentrations k standard error in each of the seven community types Number of analyses in parentheses below data the number of analyses for CaZ+ is the same as for MgZ+ K + and Na A acidic edge C open mat F marginal moat H Tsuga-dominated forest M closed mat S alkaline edge T Tlr~rja-dominated

forest AC acid lake water AL alkaline lake water

A C F H M S T AC AL

seedlings are occasionally present Ericaceous shrubs are present with Andromeda glaucophylla and ChamaecIclphne calyculata reaching high importance values Rhynchospora alba (Fig 6) is the dominant sedge however since it is also an important part of the open mat community type it is not considered as an indicator species Three species of Carex are usually present C paupercula and C limosa both of which are mostly restricted to this zone and C oligosperma which is present in two other zones

~ Ten species of Sphagnum were recorded in this community type of which two form most of the cover Sphagnum cuspidatum (Fig 6) is found submerged or emergent and is the dominant species in the wettest areas while S papillosum

I becomes dominant when there is less moisture Other bryophytes are rare in this zone and un- like the alkaline lake edge zone they make up a very small portion of the vegetation When the two edge zones are compared there is almost no overlap in species composition (Table 3) The type of community which forms next to the water of a kettle-hole lake according to our data depends largely on the physical features of the lake of which the most important may be pH and cation concentrations

The two types of edge communities also differ markedly in species diversity The mean number of vascular plants for the acidic lake edge is 15 as compared with a mean of 29 for the alkaline lake edge The opposite relationship is seen for Sphagnum a mean of 37 and a total of 10 different species for the acidic lake edge but a mean of only 18 and a total of 4 species for the alkaline Much higher diversity of other bryo- phytes in the alkaline lake edge community has already been noted It can be seen here and in the other community types as well that neither the vascular plant diversity nor the Sphagnum

diversity alone nor that of other bryophytes provides a good index to the overall plant diver- sity of these communities Each segment of the community responds differently to changes in pH cation concentration etc Similar results have previously been found for bryophyte and vascu- lar plant diversity in forest communities (Slack 1971)

Sphagnum capillaceum - Carex exilis - Vaccini- um oxycoccos Community Type (Open Mat Zone C ) (Figs 20 29 32)

The open mat community is present surround- ing alkaline lakes adjacent to the edge zone It often forms extensive mats up to 20 m wide It is wet contains very few or no trees and has little topographic variation The mean pH is 49 and the Ca2+ + Mg2+ concentration is low (53 ppm Table 2) The dominant vascular plant is Vaccinium oxycoccos Habenaria blepharo- glottis Carex exilis and Scheuchzeria palustris are also characteristic parts of this community type and show high constancy

Sphagnum capillaceum (= S capillaceum var tenellum or S rubellum in this zone) is the domi- nant Sphagnum most often forming extensive pure mats Sometimes other Sphagna become im- portant parts of this zone ie S papillosum and more rarely S cuspidatum Sphagnum magellani- cum like Chamaedaphne calyculata is present to some extent in all of the community types

The open mat community type becomes well developed only in the alkaline lake bogs In Weber Lake Bog which presently has no open water and no Sphagnum teres - Carex lasiocarpa zone there is an extensive open mat community type This seemed an anomaly until we noted that ecology classes from the University of Michigan Biological Station under Dr A Cooper had mapped Weber Lake Bog in 1967 when there was

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TABLE 3 Species present in kettle-hole bog lakes in northern Michigan with mean importance value for each of the seven com- munity types recognized in this study Number of stands indicated in parentheses See Fig 3 for abbreviations of

community types

Sphagnum Sphqftr~itn annlilatum var porosum (Warnst) Maass 3 7

capillaceum (Weiss) Schrank (= netnoreurn) 1 1 3 cetrtrale C Jens cuspidatum Ehrh ex Hoffm 4 4 5 fiiscutn (Schimp) Klinggr 0 5 girgensohtrii Russ magellanic~im Brid 2 3 9 nlajiis (Russ) C Jens 101 papillosrttn Lindb 5 3 5 pnlcl~r~it~ (Lindb) Warnst 0 5 recnruum P Beauv (sensu lato) 3 5 ~lissowii Warnst sqliarroslml Crom slibseclmdlm~ Nees in Sturm (sensu lato)

teres (Schimp) Angstr in Hartm warnstorfii Roell wlilfianlln~ Girg

Hepatics Calypogeia sphagnicola (Arn amp Perss)

Warnst amp Loeske Cepl~fflozia media Lind b

pleniceps (Aust) Lindb Ceplraloziella sp Cladopodiella flliitatrs (Nees) Buch ~opl~ocolea hamperoplrylla ( ~ i h r a d ) Dumort Loplrozia ssp Moerkia hibernica Gott Mylia anomala (Hook) Gray PeNia epiphylla (L) Corda Pellia neesiat~a (Gott) Limpr Riccardia pit~gliis (L) Gray

Lichens Cladha rangiferit~a (L) Harm Clalpnia cetlotea (Ach) Schaer

cl~loropllaea (Florke ex Somm) Spreng coniocraea (Florke) Spreng

Mosses Alilacomniiitn pallistre (Hedw) Schwaegr 1 O Bracll~l~eci~itr rritablillim (Hedw) BSG

salebrosum (Web amp Mohr) BSG Bryum psaidotriq~ietr~itn (Hedw) Schwaegr Callicladi~im l~aldaniat~ltm (Grev) Crum Callgon corclifolilitn (Hedw) Kindb

stran~inelim (Brid) Kindb Calliergot~ella clispidata (Hedw) Loeske Can~lt~l iun~ polygamrim (BSG) C Jens

stellatlitn (Hedw) C Jens Dic~~t ir im flagellare Hedw

jirscescetrs Turn polysetlim Sw

rmdlilatlrm Schrad ex Brid Drepar~pclad~ts exannulatlrs (BSG) Warnst

fluitat~s (Hedw) Warnst Hyptrlitn itnponens Hedw

lindbergii Mitt Plagion~nilrm medium BSG Plagiothecilrm denticulatlrm (Hedw) BSG Plelrrozilrm schreberi (Brid) Mitt 0 4 Pohlia nutans (Hedw) Lindb Polytricl~lrm juniperit~~im Willd ex Hedw 0 3 Rhynchostegium serrulatum (Hedw) Jaeg amp Sauerb Tetraphis pellucida Hed w

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VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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VITT AND SLACK BOG VEGETATION ANALYSIS

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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7 Jake F Weltzin Jason K Keller Scott D Bridgham J Pastor P B Allen J Chen 2005 Litter controls plant communitycomposition in a northern fen Oikos 110101111oik2005110issue-3 537-546 [CrossRef]

8 Heather E Whitehouse Suzanne E Bayley 2005 Vegetation patterns and biodiversity of peatland plant communities surroundingmid-boreal wetland ponds in Alberta Canada Canadian Journal of Botany 836 621-637 [Abstract] [PDF] [PDF Plus]

9 L D Gignac R Gauthier L Rochefort J Bubier 2004 Distribution and habitat niches of 37 peatland Cyperaceae species acrossa broad geographic range in Canada Canadian Journal of Botany 829 1292-1313 [Abstract] [PDF] [PDF Plus]

10 P Oacutedor T Standovaacuter 2002 Substrate specificity and community structure of bryophyte vegetation in a near-natural montanebeech forest Community Ecology 3 39-49 [CrossRef]

11 Michal Haacutejek Petr Hekera Petra Haacutejkovaacute 2002 Spring fen vegetation and water chemistry in the Western Carpathian flyschzone Folia Geobotanica 37 205-224 [CrossRef]

12 Brian R Hall Dudley J Raynal Donald J Leopold 2001 Environmental influences on plant species composition in ground-water seeps in the Catskill Mountains of New York Wetlands 21 125-134 [CrossRef]

13 Barbara L Bedford Mark R Walbridge Allison Aldous 1999 PATTERNS IN NUTRIENT AVAILABILITY AND PLANTDIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS Ecology 80 2151-2169 [CrossRef]

14 L Bragazza 1999 Spatial patterns of plant species in a poor mire on the Southern Alps (Italy) Plant Biosystems - An InternationalJournal Dealing with all Aspects of Plant Biology 133 83-92 [CrossRef]

15 Dennis S Anderson Ronald B Davis 1997 The vegetation and its environments in Maine peatlands Canadian Journal ofBotany 7510 1785-1805 [Abstract] [PDF] [PDF Plus]

16 R Diggelen WJ Molenaar AM Kooijman 1996 Vegetation succession in a floating mire in relation to management andhydrology Journal of Vegetation Science 7 809-820 [CrossRef]

17 Andrew S Fisher Gregory S Podniesinski Donald J Leopold 1996 Effects of drainage ditches on vegetation patterns inAbandoned agricultural peatlands in central New York Wetlands 16 397-409 [CrossRef]

18 B J Nicholson L D Gignac S E Bayley 1996 Peatland distibution along a north-south transect in the Mackenzie RiverBasin in relation to climatic and environmental gradients Vegetatio 126 119-133 [CrossRef]

19 Gay M Donaldson 1996 Oribatida (Acari) associated with three species of Sphagnum at Spruce Hole Bog New HampshireUSA Canadian Journal of Zoology 749 1706-1712 [Abstract] [PDF] [PDF Plus]

20 J Bradley Johnson 1996 Phytosociology and gradient analysis of a subalpine treed fen in Rocky Mountain National ParkColorado Canadian Journal of Botany 748 1203-1213 [Abstract] [PDF] [PDF Plus]

21 Joslashrn-Frode Nordbakken 1996 Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expansea numerical approach Nordic Journal of Botany 16 197-209 [CrossRef]

22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 2: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG

descriptive or phytosociological methods largely of the Ziirich-Montpellier school and describe a variety of bog and fen communities based primarily on vascular plants Significant papers by Aldrich (1943) Catenhuisen (1950) Conway (1949) Heinselman (1963 1965 1970) Janssen (1967) Moss (1953) Nichols (1919) Osvald (1933) Sjors (1959 1963) and Transeau (1905- 1906) have added to the knowledge of peatlands in North America In Europe and Asia where peatlands have been studied intensively regional studies by Eurola (1962) Katz (1926) Kulczynski (1949) Malmer (1962) Osvald (1925 1949) Paasio (1 933) Rose (1953) Ruuhijarvi (1 960) Sjors (1948 1950) and Tolonen (1967) have greatly increased understanding of peatlands

The term bog lake in a restricted sense has been used by some authors to include only those bodies of water enclosed by ombrotrophic peat- land As compared with bogs fens would include peatlands with minerotrophic conditions Since in the area we studied truly ombrotrophic con- ditions do not appear to be present the kettle- hole lakes are strictly speaking fen lakes His- torically the peatland-surrounded basin lakes of northern Michigan have been called kettle-hole bogs or lakes We have used the term kettle-hole bog using bog in the sense of Sphagnum- dominated habitats including potentially ombro- trophic and minerotrophic conditions The bogs studied here are limited to those areas that have a Sphagnum mat at least partially surrounding a lake formed in a kettle-hole or similar depression The objectives of this study are (I) to analyze the composition structure and pattern of the vegeta- tion surrounding the open water of kettle-hole lakes (2) to determine the influence of certain physical parameters on the bog vegetation and (3) to determine plant gradient patterns and correlations of species in Sphagnunz-dominated vegetation Both gradient analysis and com- munity approaches to vegetation have been used

Any type of vegetation may be viewed as a group of communities each characterized by particular species populations vegetation struc- ture and in most cases environmental factors Such communities may or may not be separated be ecotonal areas Phytosociological methods reinforce this view and have been widely used in studies of bog vegetation on both sides of the Atlantic

Alternately one can view a type of vegetation as a continuum in which species populations and

[EGETATION ANALYSIS 333

community structure change in relation to en- vironmental gradients Although each species has its own physiological requirements differing to some extent from those of any other species groups of species with similar responses to environmental gradients may be recognized as community types Methods of vegetational analysis using this continuum viewpoint by some authors called gradient analysis have been developed by Ramensky (1930) and in North America by Whittaker (1948 1956 and later) and by Curtis and associates (195 1 and follow- ing) These techniques including ordination have been widely used in mountain vegetation and in the forests of Wisconsin and elsewhere For example ordination methods have recently been used to analyze corticolous lichen com- munities (Jesberger and Sheard 1973) marsh vegetation (Auclair et al 1973 Walker and Coupland 1968) and peatlands (Jeglum 1971 Dirschl and Coupland 1972) but they have not been used in this type of Sphagnum bog vegeta- tion We have used several of these methods including both indirect and direct gradient analysis (Whittaker 1967) In the former vegeta- tion samples are compared on the basis of species composition and arranged in an ordination along axes which may represent environmental gradi- ents In direct gradient analysis samples are arranged along actual environmental gradients Environmental gradients that may be important in bogs include light water level cation con- centration and pH all of which may in part be influenced by the Sphagnunz itself

Study Area The kettle-hole bogs sampled in this study are

located between 4520 and 4625 N latitude and 8515 and 84201 W longitude in the Great Lakes - St Lawrence forest region (Rowe 1972) The elevation of the area is about 200-220 m above sea level and does not vary to any con- siderable degree

The upland vegetation of the area is at present a mosaic of beech-maple forests and aspen woods with some areas dominated by oak white pine and other trees The lowland tree vegetation includes white cedar swamps and black spruce forests with a few areas dominated by larch and hemlock The region was extensively glaciated during Wisconsin times and the mineral soil is of glacial origin The bedrock is covered by glacial till and rock outcrops are rare Those that

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CAN J BOT VOL 53 1975

P R E C I P I T A T I O N I c m I

2 0

16

- U

- 1 2 -

W = 3 t

4 8 - p

U1 L

E W t

4

0

- 4

FIG 1 Climograph showing mean monthly precipitation and temperature at Mackinaw City in northern Michigan (solid line) and at Willis in southeastern Michigan (broken line) Months numbered from 1 to 12 beginning with January

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- - M A C K I N A W C I T Y

- - - - - - - - W l L L l S

2 I

3 4 5 6 7 8 9 10

are present are of limestone Gates (1912) has discussed the vegetation of the Douglas Lake region in the Lower Peninsula and Voss (1956) has given a history of the floristics of Emmet and Cheboygan Counties About 20-30 of the study region is covered by peatland vegetation most of this in the form of Tl~uja occidentalis - Picea mariana - Larix laricina swamps (Gates 1942)

Climate The climate of northern Michigan can be

considered cool temperate with mean precipita- tion of 840 mm and the mean monthly tempera- ture ranging from -7degC in February to 20degC in July and August Snowfall usually begins in October or November and may continue into April or early May averaging 1575 mm per year The absolute minimum temperature is - 39degC in February and the absolute maximum tempera- ture is 40degC in July (Gates 1942) The mean

monthly precipitation and temperature for 1970- 1971 are given in Fig 1 Temperatures peak in July and August while rainfall is highest in September In most years the maximum precipi- tation occurs in late summer whereas in southern Michigan the wettest season is spring and early summer (Fig 1) The relatively low annual precipitation and extremes of winter cold and summer heat are markedly different from the cool moist maritime climates of eg New- foundland and British Columbia

Methods Vegetation

A general reconnaissance of bog vegetation and bog distribution in the northern Michigan region was con- ducted in the summer of 1970 Based on this preliminary survey eight kettle-hole bogs were selected for study Two criteria were used in selecting the bogs for sampling (I) the presence of a Sphagnum mat and (2) the presence of open water forming a lake or pool in the center of the bog Five of the bogs are located in the Lower Peninsula

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 1

County range and township locality and stand numbers for the kettle-hole lakes in northern Michigan that were analyzed in this study

Kettle-hole lake County Range and township Stand No - -

Brown Lake Chippewa R5W T45N S6 SW-114 NE-114 8 9 10 Little Lake Mackinac R8W T44N S36 NE-114 E-118 6 26 27 Little Lake 16 (Fig 17) Cheboygan RIE T37N S16 SW-114 16 41 42 43 44 45 46 47 Livingston (Fig 15) Cheboygan R3W T36N S2 SW-114 NW-114 3 4 20 21 22 Mud Lake (Fig 18) Cheboygan R2W T37N S7 SE-114 S-118 5 7 23 24 25 28 29 30 Ryerse Lake Mackinac R7W T43N S9 SE-114 SE-114 11 12 13 31 32 33 34 35 Stutsmanville (Fig 16)

(or Hard Bottom Bog) Emmet R6W T36N S23 NE-114 SE-114 1 2 17 IS 19 Weber Lake Cheboygan R3W T34N S31 NW-114 SE-114 14 15 36 37 38 3940

FIG 2 Detail of five-county region in northern Michigan showing location of the eight kettle-hole lakes sampled Open circle is location of Mackinaw City from which precipitation and temperature were taken

in Emmet and Cheboygan Counties and three are located in the Upper Peninsula in Chippewa and Mackinac Counties The geographical coordinates for the bogs studied are given in Table 1 and the locations are shown in Fig 2

Quantitative data were gathered along transects oriented both parallel to the lake shore and perpendicular to it The parallel transects consisted of 10 quadrats while the perpendicular transects ran from open water to bog forest Within each transect I-mZ quadrats were chosen randomly from each 10-m interval Ocular esti- mates of percentage cover and frequency were recorded for each species Samples of bog water from natural depressions were taken along each transect and analyzed in the laboratory (within 6 h of sampling) with a Corning Model No 5 pH meter They were frozen and later analyzed on an atomic absorption spectrophotometer at theUniversity of Alberta for Ca2 + Na + K + and MgZ+

The data were compiled to give importance-value

indices (IV) based on a maximum of 200 by adding the relative frequency to the relative mean percentage cover Both frequency and percentage cover were calculated separately for the vascular plants and separately for Sphagnum plus other associated bryophytes and lichens

Authority names and nomenclature for the vascular plants follow Fernald (1950) Crum et 01 (1973) for the Sphagnum and mosses Schuster (1953) for the hepatics and Hale and Culberson (1970) for the lichens Voucher specimens are deposited in the herbaria of the University of Alberta (ALTA) and of Russell Sage College Troy New York

Ordination and Cluster Analysis Indirect gradient-analysis as well as direct gradient-

analysis methods were used in evaluating the data from the 300 quadrats Importance-value indices for all species of Sphagnum mosses hepatics lichens and vascular plants were used to calculate coefficients of

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336 CAN J BOT VOL 53 1975

FIG 3 Two-dimensional ordination of the 47 stands sampled Lines delimit community types S Sphagnum teres - Carex lasiocarpa - Calliergonella ctispidata community type A S clispidatlim - S papillos~rtn - Carex liniosa - C patiperctila community type C S capillacelim - Carex exilis - Vaccinitinz oxycoccos community type M S magellaniclin~ - S rectirurini- Ledtim groenlandictim - Picea mariarla community type F Zlex oerticillata- Netnopanthlis mlicronata community type T Tiirija occidentalis - S ~varnstorji community type H Tstiga canadensis - S centrale community type

similarity between stands using the formula C = [2w ( a + b ) ] x 100 where C is the coefficient of similarity rv is the sum of the lowest common importance values of the two stands and a and b are the sums of the impor- tance values of the stands A and B Species were not adjusted in relation to their maximum values of frequency and cover as in Bray and Curtis (1957) A two-dimen- sional ordination of communities was constructed from the dissimilarity values (1 - C ) and yielded a patterning for interpretation of the sampled stands (Bray and Curtis 1957 Beals 1960) A computer program was developed and all calculations were completed on the MTS and

Calcomp computer facilities at the University of Alberta The Bray-Curtis ordination was considered reliable for showing the relationships of the vegetation (see Gauch and Whittaker 1972 for a discussion)

Using the ordination the stands were grouped by visual inspection of axis 1 and 2 to reflect the spatial association of the ordination and from this grouping the mean importance values shown in Table 3 were obtained

Community types as delineated in the ordination (Fig 3 ) were named indicating the dominant vascular plant and dominant Sphagntint species as well as their most

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VITT AND SLACK BOG VEGETATION ANALYSIS 337

salient feature or distribution within the bog Bog com- munities often occur as overlapping vegetation zones and well-separated species groupings may or may not be present The associations of points of the ordination are not meant to imply that bog communities are found as distinct entities however they are presented to more easily discuss the vegetation

T o obtain another point of view of community struc- ture and to assess the validity of the community types derived from our ordination the same data as were used in the ordination were analyzed by a cluster analysis (Fig 4) Importance values of all species were used in a computer program using minimum variance clustering (agglomerative least squares (Pritchard and Anderson 1971) computer program obtained from A J B Ander- son)

Results and Discussion An analysis of the data indicates that two

distinct types of kettle-hole lakes exist in northern Michigan In the vegetation surrounding each of these two types of lakes there is a gradient of plant species from open water to bog forest zones Four Sphagnum-mat community types are described below as occurring around kettle-hole lakes in northern Michigan The outer marginal moat zone often termed a lagg or marginal fen is treated as a variable unit in which several alternative community types can be recognized The occurrence of two bog forest community types is discussed in relation to the Sphagnum- mat communities The two-dimensional ordina- tion is shown in Fig 3 illustrating the pattern of distribution of the community types The ordina- tion was constructed solely from dissimilarity values between sampled stands nevertheless the stand groupings indicate environmental gradi- ents which are considered important in bog vegetation The cluster analysis distinguishes community types similar to those of the ordina- tion while some of the differences are pointed out below

Sphagnum teres - Carex lasiocarpa - Calliergon- ella cuspidata Community Type (Alkaline Lake Edge Zone S ) (Figs 1718 27)

This community type is present only around those lakes which have a relatively high pH that is Mud Lake (pH 78) Little Lake 16 (pH 75) and Ryerse Lake (pH 73) This zone was also observed by Conway (1949) in Minnesota and referred to as the sedge mat Within this com- munity type pH values and Ca2+ and Mg2+ cation concentrations are high The mean pH is 67 the mean calcium content is 77 ppm and the mean magnesium content is 18 ppm (Table 2) There is very little microtopographic differ-

entiation and the zone is extremely wet The width of this community type varies from a few decimeters at Ryerse Lake to more than 20 m on the southeastern side of Mud Lake It does not always form a continuous perimeter around the lake often being present only on one or two sides possibly in connection with inflow or outflow channels of the lake False bottoms were present only in lakes that supported this com- munity type

Ericaceous shrubs such as Chamaedaphne calyculata and Andromeda glaucophylla are only a minor component and trees are absent except for occasional islands of Alnus rugosa There is a high diversity of vascular plant species with 10 of these entirely restricted to this zone (Table 4) Carex lasiocarpa (Fig 5) is the dominant species but Cladium mariscoides Dulichium arundina- ceum Hypericum virginicum Lysimachia terres- tris Pogonia ophioglossoides Campanula apari- noides and Potentilla palustris are also very characteristic (Fig 27)

Two species of Sphagnum S teres (Fig 5) and S subsecundum (sensu lato) are found commonly in this community type with S papillosum S magellanicum and S recurvum occasionally present in small amounts Sphagnum teres often forms a continuous mat in this zone and is characteristic of areas of high light intensity high moisture high pH and high cation con- centrations Sphagnum subsecundum although never abundant was present in almost every quadrat particularly in those that fell at the waters edge Moerkia hibernica Campylium stellatum Calliergonella cuspidata and Pellia epiphylla have high frequencies in this com- munity type although Calliergonella cuspidata is the only species that is abundant

Sphagnum cuspidatum - S papillosum - Carex limosa - C paupercula Community Type (Acidic Lake Edge Zone A) (Figs 19 28)

This community type is found at the edge of lakes having pH between 50 and 70 The zone may be less than 1 m wide or absent in a few areas as at Livingston Bog or a continuous mat 2-4 m wide as at Stutsmanville Bog The pH as well as the Ca2+ and Mg2+ concentrations is much lower than in the alkaline lake edge zone (Table 2) Like the alkaline lake edge this community type is a floating mat and hence is very wet The topography is slight and there is little shade Trees are absent although Picea and Larix

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CAN J BOT VOL 53 1975

FIG 4 Dendrogram of minimum variance cluster analysis of the 47 stands sampled Letters refer to com- munity types as designated in the ordination (Fig 3)

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VITT AND SLACK BOG VEGETATION ANALYSIS 339

TABLE 2 Mean pH and cation concentrations k standard error in each of the seven community types Number of analyses in parentheses below data the number of analyses for CaZ+ is the same as for MgZ+ K + and Na A acidic edge C open mat F marginal moat H Tsuga-dominated forest M closed mat S alkaline edge T Tlr~rja-dominated

forest AC acid lake water AL alkaline lake water

A C F H M S T AC AL

seedlings are occasionally present Ericaceous shrubs are present with Andromeda glaucophylla and ChamaecIclphne calyculata reaching high importance values Rhynchospora alba (Fig 6) is the dominant sedge however since it is also an important part of the open mat community type it is not considered as an indicator species Three species of Carex are usually present C paupercula and C limosa both of which are mostly restricted to this zone and C oligosperma which is present in two other zones

~ Ten species of Sphagnum were recorded in this community type of which two form most of the cover Sphagnum cuspidatum (Fig 6) is found submerged or emergent and is the dominant species in the wettest areas while S papillosum

I becomes dominant when there is less moisture Other bryophytes are rare in this zone and un- like the alkaline lake edge zone they make up a very small portion of the vegetation When the two edge zones are compared there is almost no overlap in species composition (Table 3) The type of community which forms next to the water of a kettle-hole lake according to our data depends largely on the physical features of the lake of which the most important may be pH and cation concentrations

The two types of edge communities also differ markedly in species diversity The mean number of vascular plants for the acidic lake edge is 15 as compared with a mean of 29 for the alkaline lake edge The opposite relationship is seen for Sphagnum a mean of 37 and a total of 10 different species for the acidic lake edge but a mean of only 18 and a total of 4 species for the alkaline Much higher diversity of other bryo- phytes in the alkaline lake edge community has already been noted It can be seen here and in the other community types as well that neither the vascular plant diversity nor the Sphagnum

diversity alone nor that of other bryophytes provides a good index to the overall plant diver- sity of these communities Each segment of the community responds differently to changes in pH cation concentration etc Similar results have previously been found for bryophyte and vascu- lar plant diversity in forest communities (Slack 1971)

Sphagnum capillaceum - Carex exilis - Vaccini- um oxycoccos Community Type (Open Mat Zone C ) (Figs 20 29 32)

The open mat community is present surround- ing alkaline lakes adjacent to the edge zone It often forms extensive mats up to 20 m wide It is wet contains very few or no trees and has little topographic variation The mean pH is 49 and the Ca2+ + Mg2+ concentration is low (53 ppm Table 2) The dominant vascular plant is Vaccinium oxycoccos Habenaria blepharo- glottis Carex exilis and Scheuchzeria palustris are also characteristic parts of this community type and show high constancy

Sphagnum capillaceum (= S capillaceum var tenellum or S rubellum in this zone) is the domi- nant Sphagnum most often forming extensive pure mats Sometimes other Sphagna become im- portant parts of this zone ie S papillosum and more rarely S cuspidatum Sphagnum magellani- cum like Chamaedaphne calyculata is present to some extent in all of the community types

The open mat community type becomes well developed only in the alkaline lake bogs In Weber Lake Bog which presently has no open water and no Sphagnum teres - Carex lasiocarpa zone there is an extensive open mat community type This seemed an anomaly until we noted that ecology classes from the University of Michigan Biological Station under Dr A Cooper had mapped Weber Lake Bog in 1967 when there was

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340 CAN J BOT VOL 53 1975

TABLE 3 Species present in kettle-hole bog lakes in northern Michigan with mean importance value for each of the seven com- munity types recognized in this study Number of stands indicated in parentheses See Fig 3 for abbreviations of

community types

Sphagnum Sphqftr~itn annlilatum var porosum (Warnst) Maass 3 7

capillaceum (Weiss) Schrank (= netnoreurn) 1 1 3 cetrtrale C Jens cuspidatum Ehrh ex Hoffm 4 4 5 fiiscutn (Schimp) Klinggr 0 5 girgensohtrii Russ magellanic~im Brid 2 3 9 nlajiis (Russ) C Jens 101 papillosrttn Lindb 5 3 5 pnlcl~r~it~ (Lindb) Warnst 0 5 recnruum P Beauv (sensu lato) 3 5 ~lissowii Warnst sqliarroslml Crom slibseclmdlm~ Nees in Sturm (sensu lato)

teres (Schimp) Angstr in Hartm warnstorfii Roell wlilfianlln~ Girg

Hepatics Calypogeia sphagnicola (Arn amp Perss)

Warnst amp Loeske Cepl~fflozia media Lind b

pleniceps (Aust) Lindb Ceplraloziella sp Cladopodiella flliitatrs (Nees) Buch ~opl~ocolea hamperoplrylla ( ~ i h r a d ) Dumort Loplrozia ssp Moerkia hibernica Gott Mylia anomala (Hook) Gray PeNia epiphylla (L) Corda Pellia neesiat~a (Gott) Limpr Riccardia pit~gliis (L) Gray

Lichens Cladha rangiferit~a (L) Harm Clalpnia cetlotea (Ach) Schaer

cl~loropllaea (Florke ex Somm) Spreng coniocraea (Florke) Spreng

Mosses Alilacomniiitn pallistre (Hedw) Schwaegr 1 O Bracll~l~eci~itr rritablillim (Hedw) BSG

salebrosum (Web amp Mohr) BSG Bryum psaidotriq~ietr~itn (Hedw) Schwaegr Callicladi~im l~aldaniat~ltm (Grev) Crum Callgon corclifolilitn (Hedw) Kindb

stran~inelim (Brid) Kindb Calliergot~ella clispidata (Hedw) Loeske Can~lt~l iun~ polygamrim (BSG) C Jens

stellatlitn (Hedw) C Jens Dic~~t ir im flagellare Hedw

jirscescetrs Turn polysetlim Sw

rmdlilatlrm Schrad ex Brid Drepar~pclad~ts exannulatlrs (BSG) Warnst

fluitat~s (Hedw) Warnst Hyptrlitn itnponens Hedw

lindbergii Mitt Plagion~nilrm medium BSG Plagiothecilrm denticulatlrm (Hedw) BSG Plelrrozilrm schreberi (Brid) Mitt 0 4 Pohlia nutans (Hedw) Lindb Polytricl~lrm juniperit~~im Willd ex Hedw 0 3 Rhynchostegium serrulatum (Hedw) Jaeg amp Sauerb Tetraphis pellucida Hed w

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VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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VITT AND SLACK BOG VEGETATION ANALYSIS

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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9 L D Gignac R Gauthier L Rochefort J Bubier 2004 Distribution and habitat niches of 37 peatland Cyperaceae species acrossa broad geographic range in Canada Canadian Journal of Botany 829 1292-1313 [Abstract] [PDF] [PDF Plus]

10 P Oacutedor T Standovaacuter 2002 Substrate specificity and community structure of bryophyte vegetation in a near-natural montanebeech forest Community Ecology 3 39-49 [CrossRef]

11 Michal Haacutejek Petr Hekera Petra Haacutejkovaacute 2002 Spring fen vegetation and water chemistry in the Western Carpathian flyschzone Folia Geobotanica 37 205-224 [CrossRef]

12 Brian R Hall Dudley J Raynal Donald J Leopold 2001 Environmental influences on plant species composition in ground-water seeps in the Catskill Mountains of New York Wetlands 21 125-134 [CrossRef]

13 Barbara L Bedford Mark R Walbridge Allison Aldous 1999 PATTERNS IN NUTRIENT AVAILABILITY AND PLANTDIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS Ecology 80 2151-2169 [CrossRef]

14 L Bragazza 1999 Spatial patterns of plant species in a poor mire on the Southern Alps (Italy) Plant Biosystems - An InternationalJournal Dealing with all Aspects of Plant Biology 133 83-92 [CrossRef]

15 Dennis S Anderson Ronald B Davis 1997 The vegetation and its environments in Maine peatlands Canadian Journal ofBotany 7510 1785-1805 [Abstract] [PDF] [PDF Plus]

16 R Diggelen WJ Molenaar AM Kooijman 1996 Vegetation succession in a floating mire in relation to management andhydrology Journal of Vegetation Science 7 809-820 [CrossRef]

17 Andrew S Fisher Gregory S Podniesinski Donald J Leopold 1996 Effects of drainage ditches on vegetation patterns inAbandoned agricultural peatlands in central New York Wetlands 16 397-409 [CrossRef]

18 B J Nicholson L D Gignac S E Bayley 1996 Peatland distibution along a north-south transect in the Mackenzie RiverBasin in relation to climatic and environmental gradients Vegetatio 126 119-133 [CrossRef]

19 Gay M Donaldson 1996 Oribatida (Acari) associated with three species of Sphagnum at Spruce Hole Bog New HampshireUSA Canadian Journal of Zoology 749 1706-1712 [Abstract] [PDF] [PDF Plus]

20 J Bradley Johnson 1996 Phytosociology and gradient analysis of a subalpine treed fen in Rocky Mountain National ParkColorado Canadian Journal of Botany 748 1203-1213 [Abstract] [PDF] [PDF Plus]

21 Joslashrn-Frode Nordbakken 1996 Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expansea numerical approach Nordic Journal of Botany 16 197-209 [CrossRef]

22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 3: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

CAN J BOT VOL 53 1975

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FIG 1 Climograph showing mean monthly precipitation and temperature at Mackinaw City in northern Michigan (solid line) and at Willis in southeastern Michigan (broken line) Months numbered from 1 to 12 beginning with January

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are present are of limestone Gates (1912) has discussed the vegetation of the Douglas Lake region in the Lower Peninsula and Voss (1956) has given a history of the floristics of Emmet and Cheboygan Counties About 20-30 of the study region is covered by peatland vegetation most of this in the form of Tl~uja occidentalis - Picea mariana - Larix laricina swamps (Gates 1942)

Climate The climate of northern Michigan can be

considered cool temperate with mean precipita- tion of 840 mm and the mean monthly tempera- ture ranging from -7degC in February to 20degC in July and August Snowfall usually begins in October or November and may continue into April or early May averaging 1575 mm per year The absolute minimum temperature is - 39degC in February and the absolute maximum tempera- ture is 40degC in July (Gates 1942) The mean

monthly precipitation and temperature for 1970- 1971 are given in Fig 1 Temperatures peak in July and August while rainfall is highest in September In most years the maximum precipi- tation occurs in late summer whereas in southern Michigan the wettest season is spring and early summer (Fig 1) The relatively low annual precipitation and extremes of winter cold and summer heat are markedly different from the cool moist maritime climates of eg New- foundland and British Columbia

Methods Vegetation

A general reconnaissance of bog vegetation and bog distribution in the northern Michigan region was con- ducted in the summer of 1970 Based on this preliminary survey eight kettle-hole bogs were selected for study Two criteria were used in selecting the bogs for sampling (I) the presence of a Sphagnum mat and (2) the presence of open water forming a lake or pool in the center of the bog Five of the bogs are located in the Lower Peninsula

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 1

County range and township locality and stand numbers for the kettle-hole lakes in northern Michigan that were analyzed in this study

Kettle-hole lake County Range and township Stand No - -

Brown Lake Chippewa R5W T45N S6 SW-114 NE-114 8 9 10 Little Lake Mackinac R8W T44N S36 NE-114 E-118 6 26 27 Little Lake 16 (Fig 17) Cheboygan RIE T37N S16 SW-114 16 41 42 43 44 45 46 47 Livingston (Fig 15) Cheboygan R3W T36N S2 SW-114 NW-114 3 4 20 21 22 Mud Lake (Fig 18) Cheboygan R2W T37N S7 SE-114 S-118 5 7 23 24 25 28 29 30 Ryerse Lake Mackinac R7W T43N S9 SE-114 SE-114 11 12 13 31 32 33 34 35 Stutsmanville (Fig 16)

(or Hard Bottom Bog) Emmet R6W T36N S23 NE-114 SE-114 1 2 17 IS 19 Weber Lake Cheboygan R3W T34N S31 NW-114 SE-114 14 15 36 37 38 3940

FIG 2 Detail of five-county region in northern Michigan showing location of the eight kettle-hole lakes sampled Open circle is location of Mackinaw City from which precipitation and temperature were taken

in Emmet and Cheboygan Counties and three are located in the Upper Peninsula in Chippewa and Mackinac Counties The geographical coordinates for the bogs studied are given in Table 1 and the locations are shown in Fig 2

Quantitative data were gathered along transects oriented both parallel to the lake shore and perpendicular to it The parallel transects consisted of 10 quadrats while the perpendicular transects ran from open water to bog forest Within each transect I-mZ quadrats were chosen randomly from each 10-m interval Ocular esti- mates of percentage cover and frequency were recorded for each species Samples of bog water from natural depressions were taken along each transect and analyzed in the laboratory (within 6 h of sampling) with a Corning Model No 5 pH meter They were frozen and later analyzed on an atomic absorption spectrophotometer at theUniversity of Alberta for Ca2 + Na + K + and MgZ+

The data were compiled to give importance-value

indices (IV) based on a maximum of 200 by adding the relative frequency to the relative mean percentage cover Both frequency and percentage cover were calculated separately for the vascular plants and separately for Sphagnum plus other associated bryophytes and lichens

Authority names and nomenclature for the vascular plants follow Fernald (1950) Crum et 01 (1973) for the Sphagnum and mosses Schuster (1953) for the hepatics and Hale and Culberson (1970) for the lichens Voucher specimens are deposited in the herbaria of the University of Alberta (ALTA) and of Russell Sage College Troy New York

Ordination and Cluster Analysis Indirect gradient-analysis as well as direct gradient-

analysis methods were used in evaluating the data from the 300 quadrats Importance-value indices for all species of Sphagnum mosses hepatics lichens and vascular plants were used to calculate coefficients of

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336 CAN J BOT VOL 53 1975

FIG 3 Two-dimensional ordination of the 47 stands sampled Lines delimit community types S Sphagnum teres - Carex lasiocarpa - Calliergonella ctispidata community type A S clispidatlim - S papillos~rtn - Carex liniosa - C patiperctila community type C S capillacelim - Carex exilis - Vaccinitinz oxycoccos community type M S magellaniclin~ - S rectirurini- Ledtim groenlandictim - Picea mariarla community type F Zlex oerticillata- Netnopanthlis mlicronata community type T Tiirija occidentalis - S ~varnstorji community type H Tstiga canadensis - S centrale community type

similarity between stands using the formula C = [2w ( a + b ) ] x 100 where C is the coefficient of similarity rv is the sum of the lowest common importance values of the two stands and a and b are the sums of the impor- tance values of the stands A and B Species were not adjusted in relation to their maximum values of frequency and cover as in Bray and Curtis (1957) A two-dimen- sional ordination of communities was constructed from the dissimilarity values (1 - C ) and yielded a patterning for interpretation of the sampled stands (Bray and Curtis 1957 Beals 1960) A computer program was developed and all calculations were completed on the MTS and

Calcomp computer facilities at the University of Alberta The Bray-Curtis ordination was considered reliable for showing the relationships of the vegetation (see Gauch and Whittaker 1972 for a discussion)

Using the ordination the stands were grouped by visual inspection of axis 1 and 2 to reflect the spatial association of the ordination and from this grouping the mean importance values shown in Table 3 were obtained

Community types as delineated in the ordination (Fig 3 ) were named indicating the dominant vascular plant and dominant Sphagntint species as well as their most

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VITT AND SLACK BOG VEGETATION ANALYSIS 337

salient feature or distribution within the bog Bog com- munities often occur as overlapping vegetation zones and well-separated species groupings may or may not be present The associations of points of the ordination are not meant to imply that bog communities are found as distinct entities however they are presented to more easily discuss the vegetation

T o obtain another point of view of community struc- ture and to assess the validity of the community types derived from our ordination the same data as were used in the ordination were analyzed by a cluster analysis (Fig 4) Importance values of all species were used in a computer program using minimum variance clustering (agglomerative least squares (Pritchard and Anderson 1971) computer program obtained from A J B Ander- son)

Results and Discussion An analysis of the data indicates that two

distinct types of kettle-hole lakes exist in northern Michigan In the vegetation surrounding each of these two types of lakes there is a gradient of plant species from open water to bog forest zones Four Sphagnum-mat community types are described below as occurring around kettle-hole lakes in northern Michigan The outer marginal moat zone often termed a lagg or marginal fen is treated as a variable unit in which several alternative community types can be recognized The occurrence of two bog forest community types is discussed in relation to the Sphagnum- mat communities The two-dimensional ordina- tion is shown in Fig 3 illustrating the pattern of distribution of the community types The ordina- tion was constructed solely from dissimilarity values between sampled stands nevertheless the stand groupings indicate environmental gradi- ents which are considered important in bog vegetation The cluster analysis distinguishes community types similar to those of the ordina- tion while some of the differences are pointed out below

Sphagnum teres - Carex lasiocarpa - Calliergon- ella cuspidata Community Type (Alkaline Lake Edge Zone S ) (Figs 1718 27)

This community type is present only around those lakes which have a relatively high pH that is Mud Lake (pH 78) Little Lake 16 (pH 75) and Ryerse Lake (pH 73) This zone was also observed by Conway (1949) in Minnesota and referred to as the sedge mat Within this com- munity type pH values and Ca2+ and Mg2+ cation concentrations are high The mean pH is 67 the mean calcium content is 77 ppm and the mean magnesium content is 18 ppm (Table 2) There is very little microtopographic differ-

entiation and the zone is extremely wet The width of this community type varies from a few decimeters at Ryerse Lake to more than 20 m on the southeastern side of Mud Lake It does not always form a continuous perimeter around the lake often being present only on one or two sides possibly in connection with inflow or outflow channels of the lake False bottoms were present only in lakes that supported this com- munity type

Ericaceous shrubs such as Chamaedaphne calyculata and Andromeda glaucophylla are only a minor component and trees are absent except for occasional islands of Alnus rugosa There is a high diversity of vascular plant species with 10 of these entirely restricted to this zone (Table 4) Carex lasiocarpa (Fig 5) is the dominant species but Cladium mariscoides Dulichium arundina- ceum Hypericum virginicum Lysimachia terres- tris Pogonia ophioglossoides Campanula apari- noides and Potentilla palustris are also very characteristic (Fig 27)

Two species of Sphagnum S teres (Fig 5) and S subsecundum (sensu lato) are found commonly in this community type with S papillosum S magellanicum and S recurvum occasionally present in small amounts Sphagnum teres often forms a continuous mat in this zone and is characteristic of areas of high light intensity high moisture high pH and high cation con- centrations Sphagnum subsecundum although never abundant was present in almost every quadrat particularly in those that fell at the waters edge Moerkia hibernica Campylium stellatum Calliergonella cuspidata and Pellia epiphylla have high frequencies in this com- munity type although Calliergonella cuspidata is the only species that is abundant

Sphagnum cuspidatum - S papillosum - Carex limosa - C paupercula Community Type (Acidic Lake Edge Zone A) (Figs 19 28)

This community type is found at the edge of lakes having pH between 50 and 70 The zone may be less than 1 m wide or absent in a few areas as at Livingston Bog or a continuous mat 2-4 m wide as at Stutsmanville Bog The pH as well as the Ca2+ and Mg2+ concentrations is much lower than in the alkaline lake edge zone (Table 2) Like the alkaline lake edge this community type is a floating mat and hence is very wet The topography is slight and there is little shade Trees are absent although Picea and Larix

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CAN J BOT VOL 53 1975

FIG 4 Dendrogram of minimum variance cluster analysis of the 47 stands sampled Letters refer to com- munity types as designated in the ordination (Fig 3)

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VITT AND SLACK BOG VEGETATION ANALYSIS 339

TABLE 2 Mean pH and cation concentrations k standard error in each of the seven community types Number of analyses in parentheses below data the number of analyses for CaZ+ is the same as for MgZ+ K + and Na A acidic edge C open mat F marginal moat H Tsuga-dominated forest M closed mat S alkaline edge T Tlr~rja-dominated

forest AC acid lake water AL alkaline lake water

A C F H M S T AC AL

seedlings are occasionally present Ericaceous shrubs are present with Andromeda glaucophylla and ChamaecIclphne calyculata reaching high importance values Rhynchospora alba (Fig 6) is the dominant sedge however since it is also an important part of the open mat community type it is not considered as an indicator species Three species of Carex are usually present C paupercula and C limosa both of which are mostly restricted to this zone and C oligosperma which is present in two other zones

~ Ten species of Sphagnum were recorded in this community type of which two form most of the cover Sphagnum cuspidatum (Fig 6) is found submerged or emergent and is the dominant species in the wettest areas while S papillosum

I becomes dominant when there is less moisture Other bryophytes are rare in this zone and un- like the alkaline lake edge zone they make up a very small portion of the vegetation When the two edge zones are compared there is almost no overlap in species composition (Table 3) The type of community which forms next to the water of a kettle-hole lake according to our data depends largely on the physical features of the lake of which the most important may be pH and cation concentrations

The two types of edge communities also differ markedly in species diversity The mean number of vascular plants for the acidic lake edge is 15 as compared with a mean of 29 for the alkaline lake edge The opposite relationship is seen for Sphagnum a mean of 37 and a total of 10 different species for the acidic lake edge but a mean of only 18 and a total of 4 species for the alkaline Much higher diversity of other bryo- phytes in the alkaline lake edge community has already been noted It can be seen here and in the other community types as well that neither the vascular plant diversity nor the Sphagnum

diversity alone nor that of other bryophytes provides a good index to the overall plant diver- sity of these communities Each segment of the community responds differently to changes in pH cation concentration etc Similar results have previously been found for bryophyte and vascu- lar plant diversity in forest communities (Slack 1971)

Sphagnum capillaceum - Carex exilis - Vaccini- um oxycoccos Community Type (Open Mat Zone C ) (Figs 20 29 32)

The open mat community is present surround- ing alkaline lakes adjacent to the edge zone It often forms extensive mats up to 20 m wide It is wet contains very few or no trees and has little topographic variation The mean pH is 49 and the Ca2+ + Mg2+ concentration is low (53 ppm Table 2) The dominant vascular plant is Vaccinium oxycoccos Habenaria blepharo- glottis Carex exilis and Scheuchzeria palustris are also characteristic parts of this community type and show high constancy

Sphagnum capillaceum (= S capillaceum var tenellum or S rubellum in this zone) is the domi- nant Sphagnum most often forming extensive pure mats Sometimes other Sphagna become im- portant parts of this zone ie S papillosum and more rarely S cuspidatum Sphagnum magellani- cum like Chamaedaphne calyculata is present to some extent in all of the community types

The open mat community type becomes well developed only in the alkaline lake bogs In Weber Lake Bog which presently has no open water and no Sphagnum teres - Carex lasiocarpa zone there is an extensive open mat community type This seemed an anomaly until we noted that ecology classes from the University of Michigan Biological Station under Dr A Cooper had mapped Weber Lake Bog in 1967 when there was

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340 CAN J BOT VOL 53 1975

TABLE 3 Species present in kettle-hole bog lakes in northern Michigan with mean importance value for each of the seven com- munity types recognized in this study Number of stands indicated in parentheses See Fig 3 for abbreviations of

community types

Sphagnum Sphqftr~itn annlilatum var porosum (Warnst) Maass 3 7

capillaceum (Weiss) Schrank (= netnoreurn) 1 1 3 cetrtrale C Jens cuspidatum Ehrh ex Hoffm 4 4 5 fiiscutn (Schimp) Klinggr 0 5 girgensohtrii Russ magellanic~im Brid 2 3 9 nlajiis (Russ) C Jens 101 papillosrttn Lindb 5 3 5 pnlcl~r~it~ (Lindb) Warnst 0 5 recnruum P Beauv (sensu lato) 3 5 ~lissowii Warnst sqliarroslml Crom slibseclmdlm~ Nees in Sturm (sensu lato)

teres (Schimp) Angstr in Hartm warnstorfii Roell wlilfianlln~ Girg

Hepatics Calypogeia sphagnicola (Arn amp Perss)

Warnst amp Loeske Cepl~fflozia media Lind b

pleniceps (Aust) Lindb Ceplraloziella sp Cladopodiella flliitatrs (Nees) Buch ~opl~ocolea hamperoplrylla ( ~ i h r a d ) Dumort Loplrozia ssp Moerkia hibernica Gott Mylia anomala (Hook) Gray PeNia epiphylla (L) Corda Pellia neesiat~a (Gott) Limpr Riccardia pit~gliis (L) Gray

Lichens Cladha rangiferit~a (L) Harm Clalpnia cetlotea (Ach) Schaer

cl~loropllaea (Florke ex Somm) Spreng coniocraea (Florke) Spreng

Mosses Alilacomniiitn pallistre (Hedw) Schwaegr 1 O Bracll~l~eci~itr rritablillim (Hedw) BSG

salebrosum (Web amp Mohr) BSG Bryum psaidotriq~ietr~itn (Hedw) Schwaegr Callicladi~im l~aldaniat~ltm (Grev) Crum Callgon corclifolilitn (Hedw) Kindb

stran~inelim (Brid) Kindb Calliergot~ella clispidata (Hedw) Loeske Can~lt~l iun~ polygamrim (BSG) C Jens

stellatlitn (Hedw) C Jens Dic~~t ir im flagellare Hedw

jirscescetrs Turn polysetlim Sw

rmdlilatlrm Schrad ex Brid Drepar~pclad~ts exannulatlrs (BSG) Warnst

fluitat~s (Hedw) Warnst Hyptrlitn itnponens Hedw

lindbergii Mitt Plagion~nilrm medium BSG Plagiothecilrm denticulatlrm (Hedw) BSG Plelrrozilrm schreberi (Brid) Mitt 0 4 Pohlia nutans (Hedw) Lindb Polytricl~lrm juniperit~~im Willd ex Hedw 0 3 Rhynchostegium serrulatum (Hedw) Jaeg amp Sauerb Tetraphis pellucida Hed w

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VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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342 CAN J BOT VOL 53 1975

TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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344 CAN J BOT VOL 53 1975

Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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CAN J BOT VOL 53 1975

0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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VITT AND SLACK BOG VEGETATION ANALYSIS

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 4: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 1

County range and township locality and stand numbers for the kettle-hole lakes in northern Michigan that were analyzed in this study

Kettle-hole lake County Range and township Stand No - -

Brown Lake Chippewa R5W T45N S6 SW-114 NE-114 8 9 10 Little Lake Mackinac R8W T44N S36 NE-114 E-118 6 26 27 Little Lake 16 (Fig 17) Cheboygan RIE T37N S16 SW-114 16 41 42 43 44 45 46 47 Livingston (Fig 15) Cheboygan R3W T36N S2 SW-114 NW-114 3 4 20 21 22 Mud Lake (Fig 18) Cheboygan R2W T37N S7 SE-114 S-118 5 7 23 24 25 28 29 30 Ryerse Lake Mackinac R7W T43N S9 SE-114 SE-114 11 12 13 31 32 33 34 35 Stutsmanville (Fig 16)

(or Hard Bottom Bog) Emmet R6W T36N S23 NE-114 SE-114 1 2 17 IS 19 Weber Lake Cheboygan R3W T34N S31 NW-114 SE-114 14 15 36 37 38 3940

FIG 2 Detail of five-county region in northern Michigan showing location of the eight kettle-hole lakes sampled Open circle is location of Mackinaw City from which precipitation and temperature were taken

in Emmet and Cheboygan Counties and three are located in the Upper Peninsula in Chippewa and Mackinac Counties The geographical coordinates for the bogs studied are given in Table 1 and the locations are shown in Fig 2

Quantitative data were gathered along transects oriented both parallel to the lake shore and perpendicular to it The parallel transects consisted of 10 quadrats while the perpendicular transects ran from open water to bog forest Within each transect I-mZ quadrats were chosen randomly from each 10-m interval Ocular esti- mates of percentage cover and frequency were recorded for each species Samples of bog water from natural depressions were taken along each transect and analyzed in the laboratory (within 6 h of sampling) with a Corning Model No 5 pH meter They were frozen and later analyzed on an atomic absorption spectrophotometer at theUniversity of Alberta for Ca2 + Na + K + and MgZ+

The data were compiled to give importance-value

indices (IV) based on a maximum of 200 by adding the relative frequency to the relative mean percentage cover Both frequency and percentage cover were calculated separately for the vascular plants and separately for Sphagnum plus other associated bryophytes and lichens

Authority names and nomenclature for the vascular plants follow Fernald (1950) Crum et 01 (1973) for the Sphagnum and mosses Schuster (1953) for the hepatics and Hale and Culberson (1970) for the lichens Voucher specimens are deposited in the herbaria of the University of Alberta (ALTA) and of Russell Sage College Troy New York

Ordination and Cluster Analysis Indirect gradient-analysis as well as direct gradient-

analysis methods were used in evaluating the data from the 300 quadrats Importance-value indices for all species of Sphagnum mosses hepatics lichens and vascular plants were used to calculate coefficients of

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336 CAN J BOT VOL 53 1975

FIG 3 Two-dimensional ordination of the 47 stands sampled Lines delimit community types S Sphagnum teres - Carex lasiocarpa - Calliergonella ctispidata community type A S clispidatlim - S papillos~rtn - Carex liniosa - C patiperctila community type C S capillacelim - Carex exilis - Vaccinitinz oxycoccos community type M S magellaniclin~ - S rectirurini- Ledtim groenlandictim - Picea mariarla community type F Zlex oerticillata- Netnopanthlis mlicronata community type T Tiirija occidentalis - S ~varnstorji community type H Tstiga canadensis - S centrale community type

similarity between stands using the formula C = [2w ( a + b ) ] x 100 where C is the coefficient of similarity rv is the sum of the lowest common importance values of the two stands and a and b are the sums of the impor- tance values of the stands A and B Species were not adjusted in relation to their maximum values of frequency and cover as in Bray and Curtis (1957) A two-dimen- sional ordination of communities was constructed from the dissimilarity values (1 - C ) and yielded a patterning for interpretation of the sampled stands (Bray and Curtis 1957 Beals 1960) A computer program was developed and all calculations were completed on the MTS and

Calcomp computer facilities at the University of Alberta The Bray-Curtis ordination was considered reliable for showing the relationships of the vegetation (see Gauch and Whittaker 1972 for a discussion)

Using the ordination the stands were grouped by visual inspection of axis 1 and 2 to reflect the spatial association of the ordination and from this grouping the mean importance values shown in Table 3 were obtained

Community types as delineated in the ordination (Fig 3 ) were named indicating the dominant vascular plant and dominant Sphagntint species as well as their most

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VITT AND SLACK BOG VEGETATION ANALYSIS 337

salient feature or distribution within the bog Bog com- munities often occur as overlapping vegetation zones and well-separated species groupings may or may not be present The associations of points of the ordination are not meant to imply that bog communities are found as distinct entities however they are presented to more easily discuss the vegetation

T o obtain another point of view of community struc- ture and to assess the validity of the community types derived from our ordination the same data as were used in the ordination were analyzed by a cluster analysis (Fig 4) Importance values of all species were used in a computer program using minimum variance clustering (agglomerative least squares (Pritchard and Anderson 1971) computer program obtained from A J B Ander- son)

Results and Discussion An analysis of the data indicates that two

distinct types of kettle-hole lakes exist in northern Michigan In the vegetation surrounding each of these two types of lakes there is a gradient of plant species from open water to bog forest zones Four Sphagnum-mat community types are described below as occurring around kettle-hole lakes in northern Michigan The outer marginal moat zone often termed a lagg or marginal fen is treated as a variable unit in which several alternative community types can be recognized The occurrence of two bog forest community types is discussed in relation to the Sphagnum- mat communities The two-dimensional ordina- tion is shown in Fig 3 illustrating the pattern of distribution of the community types The ordina- tion was constructed solely from dissimilarity values between sampled stands nevertheless the stand groupings indicate environmental gradi- ents which are considered important in bog vegetation The cluster analysis distinguishes community types similar to those of the ordina- tion while some of the differences are pointed out below

Sphagnum teres - Carex lasiocarpa - Calliergon- ella cuspidata Community Type (Alkaline Lake Edge Zone S ) (Figs 1718 27)

This community type is present only around those lakes which have a relatively high pH that is Mud Lake (pH 78) Little Lake 16 (pH 75) and Ryerse Lake (pH 73) This zone was also observed by Conway (1949) in Minnesota and referred to as the sedge mat Within this com- munity type pH values and Ca2+ and Mg2+ cation concentrations are high The mean pH is 67 the mean calcium content is 77 ppm and the mean magnesium content is 18 ppm (Table 2) There is very little microtopographic differ-

entiation and the zone is extremely wet The width of this community type varies from a few decimeters at Ryerse Lake to more than 20 m on the southeastern side of Mud Lake It does not always form a continuous perimeter around the lake often being present only on one or two sides possibly in connection with inflow or outflow channels of the lake False bottoms were present only in lakes that supported this com- munity type

Ericaceous shrubs such as Chamaedaphne calyculata and Andromeda glaucophylla are only a minor component and trees are absent except for occasional islands of Alnus rugosa There is a high diversity of vascular plant species with 10 of these entirely restricted to this zone (Table 4) Carex lasiocarpa (Fig 5) is the dominant species but Cladium mariscoides Dulichium arundina- ceum Hypericum virginicum Lysimachia terres- tris Pogonia ophioglossoides Campanula apari- noides and Potentilla palustris are also very characteristic (Fig 27)

Two species of Sphagnum S teres (Fig 5) and S subsecundum (sensu lato) are found commonly in this community type with S papillosum S magellanicum and S recurvum occasionally present in small amounts Sphagnum teres often forms a continuous mat in this zone and is characteristic of areas of high light intensity high moisture high pH and high cation con- centrations Sphagnum subsecundum although never abundant was present in almost every quadrat particularly in those that fell at the waters edge Moerkia hibernica Campylium stellatum Calliergonella cuspidata and Pellia epiphylla have high frequencies in this com- munity type although Calliergonella cuspidata is the only species that is abundant

Sphagnum cuspidatum - S papillosum - Carex limosa - C paupercula Community Type (Acidic Lake Edge Zone A) (Figs 19 28)

This community type is found at the edge of lakes having pH between 50 and 70 The zone may be less than 1 m wide or absent in a few areas as at Livingston Bog or a continuous mat 2-4 m wide as at Stutsmanville Bog The pH as well as the Ca2+ and Mg2+ concentrations is much lower than in the alkaline lake edge zone (Table 2) Like the alkaline lake edge this community type is a floating mat and hence is very wet The topography is slight and there is little shade Trees are absent although Picea and Larix

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CAN J BOT VOL 53 1975

FIG 4 Dendrogram of minimum variance cluster analysis of the 47 stands sampled Letters refer to com- munity types as designated in the ordination (Fig 3)

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VITT AND SLACK BOG VEGETATION ANALYSIS 339

TABLE 2 Mean pH and cation concentrations k standard error in each of the seven community types Number of analyses in parentheses below data the number of analyses for CaZ+ is the same as for MgZ+ K + and Na A acidic edge C open mat F marginal moat H Tsuga-dominated forest M closed mat S alkaline edge T Tlr~rja-dominated

forest AC acid lake water AL alkaline lake water

A C F H M S T AC AL

seedlings are occasionally present Ericaceous shrubs are present with Andromeda glaucophylla and ChamaecIclphne calyculata reaching high importance values Rhynchospora alba (Fig 6) is the dominant sedge however since it is also an important part of the open mat community type it is not considered as an indicator species Three species of Carex are usually present C paupercula and C limosa both of which are mostly restricted to this zone and C oligosperma which is present in two other zones

~ Ten species of Sphagnum were recorded in this community type of which two form most of the cover Sphagnum cuspidatum (Fig 6) is found submerged or emergent and is the dominant species in the wettest areas while S papillosum

I becomes dominant when there is less moisture Other bryophytes are rare in this zone and un- like the alkaline lake edge zone they make up a very small portion of the vegetation When the two edge zones are compared there is almost no overlap in species composition (Table 3) The type of community which forms next to the water of a kettle-hole lake according to our data depends largely on the physical features of the lake of which the most important may be pH and cation concentrations

The two types of edge communities also differ markedly in species diversity The mean number of vascular plants for the acidic lake edge is 15 as compared with a mean of 29 for the alkaline lake edge The opposite relationship is seen for Sphagnum a mean of 37 and a total of 10 different species for the acidic lake edge but a mean of only 18 and a total of 4 species for the alkaline Much higher diversity of other bryo- phytes in the alkaline lake edge community has already been noted It can be seen here and in the other community types as well that neither the vascular plant diversity nor the Sphagnum

diversity alone nor that of other bryophytes provides a good index to the overall plant diver- sity of these communities Each segment of the community responds differently to changes in pH cation concentration etc Similar results have previously been found for bryophyte and vascu- lar plant diversity in forest communities (Slack 1971)

Sphagnum capillaceum - Carex exilis - Vaccini- um oxycoccos Community Type (Open Mat Zone C ) (Figs 20 29 32)

The open mat community is present surround- ing alkaline lakes adjacent to the edge zone It often forms extensive mats up to 20 m wide It is wet contains very few or no trees and has little topographic variation The mean pH is 49 and the Ca2+ + Mg2+ concentration is low (53 ppm Table 2) The dominant vascular plant is Vaccinium oxycoccos Habenaria blepharo- glottis Carex exilis and Scheuchzeria palustris are also characteristic parts of this community type and show high constancy

Sphagnum capillaceum (= S capillaceum var tenellum or S rubellum in this zone) is the domi- nant Sphagnum most often forming extensive pure mats Sometimes other Sphagna become im- portant parts of this zone ie S papillosum and more rarely S cuspidatum Sphagnum magellani- cum like Chamaedaphne calyculata is present to some extent in all of the community types

The open mat community type becomes well developed only in the alkaline lake bogs In Weber Lake Bog which presently has no open water and no Sphagnum teres - Carex lasiocarpa zone there is an extensive open mat community type This seemed an anomaly until we noted that ecology classes from the University of Michigan Biological Station under Dr A Cooper had mapped Weber Lake Bog in 1967 when there was

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340 CAN J BOT VOL 53 1975

TABLE 3 Species present in kettle-hole bog lakes in northern Michigan with mean importance value for each of the seven com- munity types recognized in this study Number of stands indicated in parentheses See Fig 3 for abbreviations of

community types

Sphagnum Sphqftr~itn annlilatum var porosum (Warnst) Maass 3 7

capillaceum (Weiss) Schrank (= netnoreurn) 1 1 3 cetrtrale C Jens cuspidatum Ehrh ex Hoffm 4 4 5 fiiscutn (Schimp) Klinggr 0 5 girgensohtrii Russ magellanic~im Brid 2 3 9 nlajiis (Russ) C Jens 101 papillosrttn Lindb 5 3 5 pnlcl~r~it~ (Lindb) Warnst 0 5 recnruum P Beauv (sensu lato) 3 5 ~lissowii Warnst sqliarroslml Crom slibseclmdlm~ Nees in Sturm (sensu lato)

teres (Schimp) Angstr in Hartm warnstorfii Roell wlilfianlln~ Girg

Hepatics Calypogeia sphagnicola (Arn amp Perss)

Warnst amp Loeske Cepl~fflozia media Lind b

pleniceps (Aust) Lindb Ceplraloziella sp Cladopodiella flliitatrs (Nees) Buch ~opl~ocolea hamperoplrylla ( ~ i h r a d ) Dumort Loplrozia ssp Moerkia hibernica Gott Mylia anomala (Hook) Gray PeNia epiphylla (L) Corda Pellia neesiat~a (Gott) Limpr Riccardia pit~gliis (L) Gray

Lichens Cladha rangiferit~a (L) Harm Clalpnia cetlotea (Ach) Schaer

cl~loropllaea (Florke ex Somm) Spreng coniocraea (Florke) Spreng

Mosses Alilacomniiitn pallistre (Hedw) Schwaegr 1 O Bracll~l~eci~itr rritablillim (Hedw) BSG

salebrosum (Web amp Mohr) BSG Bryum psaidotriq~ietr~itn (Hedw) Schwaegr Callicladi~im l~aldaniat~ltm (Grev) Crum Callgon corclifolilitn (Hedw) Kindb

stran~inelim (Brid) Kindb Calliergot~ella clispidata (Hedw) Loeske Can~lt~l iun~ polygamrim (BSG) C Jens

stellatlitn (Hedw) C Jens Dic~~t ir im flagellare Hedw

jirscescetrs Turn polysetlim Sw

rmdlilatlrm Schrad ex Brid Drepar~pclad~ts exannulatlrs (BSG) Warnst

fluitat~s (Hedw) Warnst Hyptrlitn itnponens Hedw

lindbergii Mitt Plagion~nilrm medium BSG Plagiothecilrm denticulatlrm (Hedw) BSG Plelrrozilrm schreberi (Brid) Mitt 0 4 Pohlia nutans (Hedw) Lindb Polytricl~lrm juniperit~~im Willd ex Hedw 0 3 Rhynchostegium serrulatum (Hedw) Jaeg amp Sauerb Tetraphis pellucida Hed w

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VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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342 CAN J BOT VOL 53 1975

TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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344 CAN J BOT VOL 53 1975

Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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VITT AND SLACK BOG VEGETATION ANALYSIS

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

Huntington marsh Quebec can J Bot 51 1231-12471 BEALS E 1960 Forest bird communities in the Apostle

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BRAY J R and J T CURTIS 1957 An ordination of upland forest communities of southern Wisconsin Ecol Monogr 27 325-349

CATENHUISEN J 1950 Secondary successions on the peatlands of glacial Lake Wisconsin Trans Wis Acad Sci Arts Lett 40 29-48

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

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KULCZYNSKI S 1949 TorfowiskaPolesia Mem LAcad Pol Sci Lett CI Sci Math Nat Ser B Sci Nat 15 1-356

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in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 5: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

336 CAN J BOT VOL 53 1975

FIG 3 Two-dimensional ordination of the 47 stands sampled Lines delimit community types S Sphagnum teres - Carex lasiocarpa - Calliergonella ctispidata community type A S clispidatlim - S papillos~rtn - Carex liniosa - C patiperctila community type C S capillacelim - Carex exilis - Vaccinitinz oxycoccos community type M S magellaniclin~ - S rectirurini- Ledtim groenlandictim - Picea mariarla community type F Zlex oerticillata- Netnopanthlis mlicronata community type T Tiirija occidentalis - S ~varnstorji community type H Tstiga canadensis - S centrale community type

similarity between stands using the formula C = [2w ( a + b ) ] x 100 where C is the coefficient of similarity rv is the sum of the lowest common importance values of the two stands and a and b are the sums of the impor- tance values of the stands A and B Species were not adjusted in relation to their maximum values of frequency and cover as in Bray and Curtis (1957) A two-dimen- sional ordination of communities was constructed from the dissimilarity values (1 - C ) and yielded a patterning for interpretation of the sampled stands (Bray and Curtis 1957 Beals 1960) A computer program was developed and all calculations were completed on the MTS and

Calcomp computer facilities at the University of Alberta The Bray-Curtis ordination was considered reliable for showing the relationships of the vegetation (see Gauch and Whittaker 1972 for a discussion)

Using the ordination the stands were grouped by visual inspection of axis 1 and 2 to reflect the spatial association of the ordination and from this grouping the mean importance values shown in Table 3 were obtained

Community types as delineated in the ordination (Fig 3 ) were named indicating the dominant vascular plant and dominant Sphagntint species as well as their most

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VITT AND SLACK BOG VEGETATION ANALYSIS 337

salient feature or distribution within the bog Bog com- munities often occur as overlapping vegetation zones and well-separated species groupings may or may not be present The associations of points of the ordination are not meant to imply that bog communities are found as distinct entities however they are presented to more easily discuss the vegetation

T o obtain another point of view of community struc- ture and to assess the validity of the community types derived from our ordination the same data as were used in the ordination were analyzed by a cluster analysis (Fig 4) Importance values of all species were used in a computer program using minimum variance clustering (agglomerative least squares (Pritchard and Anderson 1971) computer program obtained from A J B Ander- son)

Results and Discussion An analysis of the data indicates that two

distinct types of kettle-hole lakes exist in northern Michigan In the vegetation surrounding each of these two types of lakes there is a gradient of plant species from open water to bog forest zones Four Sphagnum-mat community types are described below as occurring around kettle-hole lakes in northern Michigan The outer marginal moat zone often termed a lagg or marginal fen is treated as a variable unit in which several alternative community types can be recognized The occurrence of two bog forest community types is discussed in relation to the Sphagnum- mat communities The two-dimensional ordina- tion is shown in Fig 3 illustrating the pattern of distribution of the community types The ordina- tion was constructed solely from dissimilarity values between sampled stands nevertheless the stand groupings indicate environmental gradi- ents which are considered important in bog vegetation The cluster analysis distinguishes community types similar to those of the ordina- tion while some of the differences are pointed out below

Sphagnum teres - Carex lasiocarpa - Calliergon- ella cuspidata Community Type (Alkaline Lake Edge Zone S ) (Figs 1718 27)

This community type is present only around those lakes which have a relatively high pH that is Mud Lake (pH 78) Little Lake 16 (pH 75) and Ryerse Lake (pH 73) This zone was also observed by Conway (1949) in Minnesota and referred to as the sedge mat Within this com- munity type pH values and Ca2+ and Mg2+ cation concentrations are high The mean pH is 67 the mean calcium content is 77 ppm and the mean magnesium content is 18 ppm (Table 2) There is very little microtopographic differ-

entiation and the zone is extremely wet The width of this community type varies from a few decimeters at Ryerse Lake to more than 20 m on the southeastern side of Mud Lake It does not always form a continuous perimeter around the lake often being present only on one or two sides possibly in connection with inflow or outflow channels of the lake False bottoms were present only in lakes that supported this com- munity type

Ericaceous shrubs such as Chamaedaphne calyculata and Andromeda glaucophylla are only a minor component and trees are absent except for occasional islands of Alnus rugosa There is a high diversity of vascular plant species with 10 of these entirely restricted to this zone (Table 4) Carex lasiocarpa (Fig 5) is the dominant species but Cladium mariscoides Dulichium arundina- ceum Hypericum virginicum Lysimachia terres- tris Pogonia ophioglossoides Campanula apari- noides and Potentilla palustris are also very characteristic (Fig 27)

Two species of Sphagnum S teres (Fig 5) and S subsecundum (sensu lato) are found commonly in this community type with S papillosum S magellanicum and S recurvum occasionally present in small amounts Sphagnum teres often forms a continuous mat in this zone and is characteristic of areas of high light intensity high moisture high pH and high cation con- centrations Sphagnum subsecundum although never abundant was present in almost every quadrat particularly in those that fell at the waters edge Moerkia hibernica Campylium stellatum Calliergonella cuspidata and Pellia epiphylla have high frequencies in this com- munity type although Calliergonella cuspidata is the only species that is abundant

Sphagnum cuspidatum - S papillosum - Carex limosa - C paupercula Community Type (Acidic Lake Edge Zone A) (Figs 19 28)

This community type is found at the edge of lakes having pH between 50 and 70 The zone may be less than 1 m wide or absent in a few areas as at Livingston Bog or a continuous mat 2-4 m wide as at Stutsmanville Bog The pH as well as the Ca2+ and Mg2+ concentrations is much lower than in the alkaline lake edge zone (Table 2) Like the alkaline lake edge this community type is a floating mat and hence is very wet The topography is slight and there is little shade Trees are absent although Picea and Larix

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FIG 4 Dendrogram of minimum variance cluster analysis of the 47 stands sampled Letters refer to com- munity types as designated in the ordination (Fig 3)

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VITT AND SLACK BOG VEGETATION ANALYSIS 339

TABLE 2 Mean pH and cation concentrations k standard error in each of the seven community types Number of analyses in parentheses below data the number of analyses for CaZ+ is the same as for MgZ+ K + and Na A acidic edge C open mat F marginal moat H Tsuga-dominated forest M closed mat S alkaline edge T Tlr~rja-dominated

forest AC acid lake water AL alkaline lake water

A C F H M S T AC AL

seedlings are occasionally present Ericaceous shrubs are present with Andromeda glaucophylla and ChamaecIclphne calyculata reaching high importance values Rhynchospora alba (Fig 6) is the dominant sedge however since it is also an important part of the open mat community type it is not considered as an indicator species Three species of Carex are usually present C paupercula and C limosa both of which are mostly restricted to this zone and C oligosperma which is present in two other zones

~ Ten species of Sphagnum were recorded in this community type of which two form most of the cover Sphagnum cuspidatum (Fig 6) is found submerged or emergent and is the dominant species in the wettest areas while S papillosum

I becomes dominant when there is less moisture Other bryophytes are rare in this zone and un- like the alkaline lake edge zone they make up a very small portion of the vegetation When the two edge zones are compared there is almost no overlap in species composition (Table 3) The type of community which forms next to the water of a kettle-hole lake according to our data depends largely on the physical features of the lake of which the most important may be pH and cation concentrations

The two types of edge communities also differ markedly in species diversity The mean number of vascular plants for the acidic lake edge is 15 as compared with a mean of 29 for the alkaline lake edge The opposite relationship is seen for Sphagnum a mean of 37 and a total of 10 different species for the acidic lake edge but a mean of only 18 and a total of 4 species for the alkaline Much higher diversity of other bryo- phytes in the alkaline lake edge community has already been noted It can be seen here and in the other community types as well that neither the vascular plant diversity nor the Sphagnum

diversity alone nor that of other bryophytes provides a good index to the overall plant diver- sity of these communities Each segment of the community responds differently to changes in pH cation concentration etc Similar results have previously been found for bryophyte and vascu- lar plant diversity in forest communities (Slack 1971)

Sphagnum capillaceum - Carex exilis - Vaccini- um oxycoccos Community Type (Open Mat Zone C ) (Figs 20 29 32)

The open mat community is present surround- ing alkaline lakes adjacent to the edge zone It often forms extensive mats up to 20 m wide It is wet contains very few or no trees and has little topographic variation The mean pH is 49 and the Ca2+ + Mg2+ concentration is low (53 ppm Table 2) The dominant vascular plant is Vaccinium oxycoccos Habenaria blepharo- glottis Carex exilis and Scheuchzeria palustris are also characteristic parts of this community type and show high constancy

Sphagnum capillaceum (= S capillaceum var tenellum or S rubellum in this zone) is the domi- nant Sphagnum most often forming extensive pure mats Sometimes other Sphagna become im- portant parts of this zone ie S papillosum and more rarely S cuspidatum Sphagnum magellani- cum like Chamaedaphne calyculata is present to some extent in all of the community types

The open mat community type becomes well developed only in the alkaline lake bogs In Weber Lake Bog which presently has no open water and no Sphagnum teres - Carex lasiocarpa zone there is an extensive open mat community type This seemed an anomaly until we noted that ecology classes from the University of Michigan Biological Station under Dr A Cooper had mapped Weber Lake Bog in 1967 when there was

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TABLE 3 Species present in kettle-hole bog lakes in northern Michigan with mean importance value for each of the seven com- munity types recognized in this study Number of stands indicated in parentheses See Fig 3 for abbreviations of

community types

Sphagnum Sphqftr~itn annlilatum var porosum (Warnst) Maass 3 7

capillaceum (Weiss) Schrank (= netnoreurn) 1 1 3 cetrtrale C Jens cuspidatum Ehrh ex Hoffm 4 4 5 fiiscutn (Schimp) Klinggr 0 5 girgensohtrii Russ magellanic~im Brid 2 3 9 nlajiis (Russ) C Jens 101 papillosrttn Lindb 5 3 5 pnlcl~r~it~ (Lindb) Warnst 0 5 recnruum P Beauv (sensu lato) 3 5 ~lissowii Warnst sqliarroslml Crom slibseclmdlm~ Nees in Sturm (sensu lato)

teres (Schimp) Angstr in Hartm warnstorfii Roell wlilfianlln~ Girg

Hepatics Calypogeia sphagnicola (Arn amp Perss)

Warnst amp Loeske Cepl~fflozia media Lind b

pleniceps (Aust) Lindb Ceplraloziella sp Cladopodiella flliitatrs (Nees) Buch ~opl~ocolea hamperoplrylla ( ~ i h r a d ) Dumort Loplrozia ssp Moerkia hibernica Gott Mylia anomala (Hook) Gray PeNia epiphylla (L) Corda Pellia neesiat~a (Gott) Limpr Riccardia pit~gliis (L) Gray

Lichens Cladha rangiferit~a (L) Harm Clalpnia cetlotea (Ach) Schaer

cl~loropllaea (Florke ex Somm) Spreng coniocraea (Florke) Spreng

Mosses Alilacomniiitn pallistre (Hedw) Schwaegr 1 O Bracll~l~eci~itr rritablillim (Hedw) BSG

salebrosum (Web amp Mohr) BSG Bryum psaidotriq~ietr~itn (Hedw) Schwaegr Callicladi~im l~aldaniat~ltm (Grev) Crum Callgon corclifolilitn (Hedw) Kindb

stran~inelim (Brid) Kindb Calliergot~ella clispidata (Hedw) Loeske Can~lt~l iun~ polygamrim (BSG) C Jens

stellatlitn (Hedw) C Jens Dic~~t ir im flagellare Hedw

jirscescetrs Turn polysetlim Sw

rmdlilatlrm Schrad ex Brid Drepar~pclad~ts exannulatlrs (BSG) Warnst

fluitat~s (Hedw) Warnst Hyptrlitn itnponens Hedw

lindbergii Mitt Plagion~nilrm medium BSG Plagiothecilrm denticulatlrm (Hedw) BSG Plelrrozilrm schreberi (Brid) Mitt 0 4 Pohlia nutans (Hedw) Lindb Polytricl~lrm juniperit~~im Willd ex Hedw 0 3 Rhynchostegium serrulatum (Hedw) Jaeg amp Sauerb Tetraphis pellucida Hed w

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VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

Huntington marsh Quebec can J Bot 51 1231-12471 BEALS E 1960 Forest bird communities in the Apostle

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

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30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

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34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

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41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 6: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS 337

salient feature or distribution within the bog Bog com- munities often occur as overlapping vegetation zones and well-separated species groupings may or may not be present The associations of points of the ordination are not meant to imply that bog communities are found as distinct entities however they are presented to more easily discuss the vegetation

T o obtain another point of view of community struc- ture and to assess the validity of the community types derived from our ordination the same data as were used in the ordination were analyzed by a cluster analysis (Fig 4) Importance values of all species were used in a computer program using minimum variance clustering (agglomerative least squares (Pritchard and Anderson 1971) computer program obtained from A J B Ander- son)

Results and Discussion An analysis of the data indicates that two

distinct types of kettle-hole lakes exist in northern Michigan In the vegetation surrounding each of these two types of lakes there is a gradient of plant species from open water to bog forest zones Four Sphagnum-mat community types are described below as occurring around kettle-hole lakes in northern Michigan The outer marginal moat zone often termed a lagg or marginal fen is treated as a variable unit in which several alternative community types can be recognized The occurrence of two bog forest community types is discussed in relation to the Sphagnum- mat communities The two-dimensional ordina- tion is shown in Fig 3 illustrating the pattern of distribution of the community types The ordina- tion was constructed solely from dissimilarity values between sampled stands nevertheless the stand groupings indicate environmental gradi- ents which are considered important in bog vegetation The cluster analysis distinguishes community types similar to those of the ordina- tion while some of the differences are pointed out below

Sphagnum teres - Carex lasiocarpa - Calliergon- ella cuspidata Community Type (Alkaline Lake Edge Zone S ) (Figs 1718 27)

This community type is present only around those lakes which have a relatively high pH that is Mud Lake (pH 78) Little Lake 16 (pH 75) and Ryerse Lake (pH 73) This zone was also observed by Conway (1949) in Minnesota and referred to as the sedge mat Within this com- munity type pH values and Ca2+ and Mg2+ cation concentrations are high The mean pH is 67 the mean calcium content is 77 ppm and the mean magnesium content is 18 ppm (Table 2) There is very little microtopographic differ-

entiation and the zone is extremely wet The width of this community type varies from a few decimeters at Ryerse Lake to more than 20 m on the southeastern side of Mud Lake It does not always form a continuous perimeter around the lake often being present only on one or two sides possibly in connection with inflow or outflow channels of the lake False bottoms were present only in lakes that supported this com- munity type

Ericaceous shrubs such as Chamaedaphne calyculata and Andromeda glaucophylla are only a minor component and trees are absent except for occasional islands of Alnus rugosa There is a high diversity of vascular plant species with 10 of these entirely restricted to this zone (Table 4) Carex lasiocarpa (Fig 5) is the dominant species but Cladium mariscoides Dulichium arundina- ceum Hypericum virginicum Lysimachia terres- tris Pogonia ophioglossoides Campanula apari- noides and Potentilla palustris are also very characteristic (Fig 27)

Two species of Sphagnum S teres (Fig 5) and S subsecundum (sensu lato) are found commonly in this community type with S papillosum S magellanicum and S recurvum occasionally present in small amounts Sphagnum teres often forms a continuous mat in this zone and is characteristic of areas of high light intensity high moisture high pH and high cation con- centrations Sphagnum subsecundum although never abundant was present in almost every quadrat particularly in those that fell at the waters edge Moerkia hibernica Campylium stellatum Calliergonella cuspidata and Pellia epiphylla have high frequencies in this com- munity type although Calliergonella cuspidata is the only species that is abundant

Sphagnum cuspidatum - S papillosum - Carex limosa - C paupercula Community Type (Acidic Lake Edge Zone A) (Figs 19 28)

This community type is found at the edge of lakes having pH between 50 and 70 The zone may be less than 1 m wide or absent in a few areas as at Livingston Bog or a continuous mat 2-4 m wide as at Stutsmanville Bog The pH as well as the Ca2+ and Mg2+ concentrations is much lower than in the alkaline lake edge zone (Table 2) Like the alkaline lake edge this community type is a floating mat and hence is very wet The topography is slight and there is little shade Trees are absent although Picea and Larix

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CAN J BOT VOL 53 1975

FIG 4 Dendrogram of minimum variance cluster analysis of the 47 stands sampled Letters refer to com- munity types as designated in the ordination (Fig 3)

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VITT AND SLACK BOG VEGETATION ANALYSIS 339

TABLE 2 Mean pH and cation concentrations k standard error in each of the seven community types Number of analyses in parentheses below data the number of analyses for CaZ+ is the same as for MgZ+ K + and Na A acidic edge C open mat F marginal moat H Tsuga-dominated forest M closed mat S alkaline edge T Tlr~rja-dominated

forest AC acid lake water AL alkaline lake water

A C F H M S T AC AL

seedlings are occasionally present Ericaceous shrubs are present with Andromeda glaucophylla and ChamaecIclphne calyculata reaching high importance values Rhynchospora alba (Fig 6) is the dominant sedge however since it is also an important part of the open mat community type it is not considered as an indicator species Three species of Carex are usually present C paupercula and C limosa both of which are mostly restricted to this zone and C oligosperma which is present in two other zones

~ Ten species of Sphagnum were recorded in this community type of which two form most of the cover Sphagnum cuspidatum (Fig 6) is found submerged or emergent and is the dominant species in the wettest areas while S papillosum

I becomes dominant when there is less moisture Other bryophytes are rare in this zone and un- like the alkaline lake edge zone they make up a very small portion of the vegetation When the two edge zones are compared there is almost no overlap in species composition (Table 3) The type of community which forms next to the water of a kettle-hole lake according to our data depends largely on the physical features of the lake of which the most important may be pH and cation concentrations

The two types of edge communities also differ markedly in species diversity The mean number of vascular plants for the acidic lake edge is 15 as compared with a mean of 29 for the alkaline lake edge The opposite relationship is seen for Sphagnum a mean of 37 and a total of 10 different species for the acidic lake edge but a mean of only 18 and a total of 4 species for the alkaline Much higher diversity of other bryo- phytes in the alkaline lake edge community has already been noted It can be seen here and in the other community types as well that neither the vascular plant diversity nor the Sphagnum

diversity alone nor that of other bryophytes provides a good index to the overall plant diver- sity of these communities Each segment of the community responds differently to changes in pH cation concentration etc Similar results have previously been found for bryophyte and vascu- lar plant diversity in forest communities (Slack 1971)

Sphagnum capillaceum - Carex exilis - Vaccini- um oxycoccos Community Type (Open Mat Zone C ) (Figs 20 29 32)

The open mat community is present surround- ing alkaline lakes adjacent to the edge zone It often forms extensive mats up to 20 m wide It is wet contains very few or no trees and has little topographic variation The mean pH is 49 and the Ca2+ + Mg2+ concentration is low (53 ppm Table 2) The dominant vascular plant is Vaccinium oxycoccos Habenaria blepharo- glottis Carex exilis and Scheuchzeria palustris are also characteristic parts of this community type and show high constancy

Sphagnum capillaceum (= S capillaceum var tenellum or S rubellum in this zone) is the domi- nant Sphagnum most often forming extensive pure mats Sometimes other Sphagna become im- portant parts of this zone ie S papillosum and more rarely S cuspidatum Sphagnum magellani- cum like Chamaedaphne calyculata is present to some extent in all of the community types

The open mat community type becomes well developed only in the alkaline lake bogs In Weber Lake Bog which presently has no open water and no Sphagnum teres - Carex lasiocarpa zone there is an extensive open mat community type This seemed an anomaly until we noted that ecology classes from the University of Michigan Biological Station under Dr A Cooper had mapped Weber Lake Bog in 1967 when there was

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340 CAN J BOT VOL 53 1975

TABLE 3 Species present in kettle-hole bog lakes in northern Michigan with mean importance value for each of the seven com- munity types recognized in this study Number of stands indicated in parentheses See Fig 3 for abbreviations of

community types

Sphagnum Sphqftr~itn annlilatum var porosum (Warnst) Maass 3 7

capillaceum (Weiss) Schrank (= netnoreurn) 1 1 3 cetrtrale C Jens cuspidatum Ehrh ex Hoffm 4 4 5 fiiscutn (Schimp) Klinggr 0 5 girgensohtrii Russ magellanic~im Brid 2 3 9 nlajiis (Russ) C Jens 101 papillosrttn Lindb 5 3 5 pnlcl~r~it~ (Lindb) Warnst 0 5 recnruum P Beauv (sensu lato) 3 5 ~lissowii Warnst sqliarroslml Crom slibseclmdlm~ Nees in Sturm (sensu lato)

teres (Schimp) Angstr in Hartm warnstorfii Roell wlilfianlln~ Girg

Hepatics Calypogeia sphagnicola (Arn amp Perss)

Warnst amp Loeske Cepl~fflozia media Lind b

pleniceps (Aust) Lindb Ceplraloziella sp Cladopodiella flliitatrs (Nees) Buch ~opl~ocolea hamperoplrylla ( ~ i h r a d ) Dumort Loplrozia ssp Moerkia hibernica Gott Mylia anomala (Hook) Gray PeNia epiphylla (L) Corda Pellia neesiat~a (Gott) Limpr Riccardia pit~gliis (L) Gray

Lichens Cladha rangiferit~a (L) Harm Clalpnia cetlotea (Ach) Schaer

cl~loropllaea (Florke ex Somm) Spreng coniocraea (Florke) Spreng

Mosses Alilacomniiitn pallistre (Hedw) Schwaegr 1 O Bracll~l~eci~itr rritablillim (Hedw) BSG

salebrosum (Web amp Mohr) BSG Bryum psaidotriq~ietr~itn (Hedw) Schwaegr Callicladi~im l~aldaniat~ltm (Grev) Crum Callgon corclifolilitn (Hedw) Kindb

stran~inelim (Brid) Kindb Calliergot~ella clispidata (Hedw) Loeske Can~lt~l iun~ polygamrim (BSG) C Jens

stellatlitn (Hedw) C Jens Dic~~t ir im flagellare Hedw

jirscescetrs Turn polysetlim Sw

rmdlilatlrm Schrad ex Brid Drepar~pclad~ts exannulatlrs (BSG) Warnst

fluitat~s (Hedw) Warnst Hyptrlitn itnponens Hedw

lindbergii Mitt Plagion~nilrm medium BSG Plagiothecilrm denticulatlrm (Hedw) BSG Plelrrozilrm schreberi (Brid) Mitt 0 4 Pohlia nutans (Hedw) Lindb Polytricl~lrm juniperit~~im Willd ex Hedw 0 3 Rhynchostegium serrulatum (Hedw) Jaeg amp Sauerb Tetraphis pellucida Hed w

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VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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CAN J BOT VOL 53 1975

0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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VITT AND SLACK BOG VEGETATION ANALYSIS

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 7: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

CAN J BOT VOL 53 1975

FIG 4 Dendrogram of minimum variance cluster analysis of the 47 stands sampled Letters refer to com- munity types as designated in the ordination (Fig 3)

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VITT AND SLACK BOG VEGETATION ANALYSIS 339

TABLE 2 Mean pH and cation concentrations k standard error in each of the seven community types Number of analyses in parentheses below data the number of analyses for CaZ+ is the same as for MgZ+ K + and Na A acidic edge C open mat F marginal moat H Tsuga-dominated forest M closed mat S alkaline edge T Tlr~rja-dominated

forest AC acid lake water AL alkaline lake water

A C F H M S T AC AL

seedlings are occasionally present Ericaceous shrubs are present with Andromeda glaucophylla and ChamaecIclphne calyculata reaching high importance values Rhynchospora alba (Fig 6) is the dominant sedge however since it is also an important part of the open mat community type it is not considered as an indicator species Three species of Carex are usually present C paupercula and C limosa both of which are mostly restricted to this zone and C oligosperma which is present in two other zones

~ Ten species of Sphagnum were recorded in this community type of which two form most of the cover Sphagnum cuspidatum (Fig 6) is found submerged or emergent and is the dominant species in the wettest areas while S papillosum

I becomes dominant when there is less moisture Other bryophytes are rare in this zone and un- like the alkaline lake edge zone they make up a very small portion of the vegetation When the two edge zones are compared there is almost no overlap in species composition (Table 3) The type of community which forms next to the water of a kettle-hole lake according to our data depends largely on the physical features of the lake of which the most important may be pH and cation concentrations

The two types of edge communities also differ markedly in species diversity The mean number of vascular plants for the acidic lake edge is 15 as compared with a mean of 29 for the alkaline lake edge The opposite relationship is seen for Sphagnum a mean of 37 and a total of 10 different species for the acidic lake edge but a mean of only 18 and a total of 4 species for the alkaline Much higher diversity of other bryo- phytes in the alkaline lake edge community has already been noted It can be seen here and in the other community types as well that neither the vascular plant diversity nor the Sphagnum

diversity alone nor that of other bryophytes provides a good index to the overall plant diver- sity of these communities Each segment of the community responds differently to changes in pH cation concentration etc Similar results have previously been found for bryophyte and vascu- lar plant diversity in forest communities (Slack 1971)

Sphagnum capillaceum - Carex exilis - Vaccini- um oxycoccos Community Type (Open Mat Zone C ) (Figs 20 29 32)

The open mat community is present surround- ing alkaline lakes adjacent to the edge zone It often forms extensive mats up to 20 m wide It is wet contains very few or no trees and has little topographic variation The mean pH is 49 and the Ca2+ + Mg2+ concentration is low (53 ppm Table 2) The dominant vascular plant is Vaccinium oxycoccos Habenaria blepharo- glottis Carex exilis and Scheuchzeria palustris are also characteristic parts of this community type and show high constancy

Sphagnum capillaceum (= S capillaceum var tenellum or S rubellum in this zone) is the domi- nant Sphagnum most often forming extensive pure mats Sometimes other Sphagna become im- portant parts of this zone ie S papillosum and more rarely S cuspidatum Sphagnum magellani- cum like Chamaedaphne calyculata is present to some extent in all of the community types

The open mat community type becomes well developed only in the alkaline lake bogs In Weber Lake Bog which presently has no open water and no Sphagnum teres - Carex lasiocarpa zone there is an extensive open mat community type This seemed an anomaly until we noted that ecology classes from the University of Michigan Biological Station under Dr A Cooper had mapped Weber Lake Bog in 1967 when there was

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340 CAN J BOT VOL 53 1975

TABLE 3 Species present in kettle-hole bog lakes in northern Michigan with mean importance value for each of the seven com- munity types recognized in this study Number of stands indicated in parentheses See Fig 3 for abbreviations of

community types

Sphagnum Sphqftr~itn annlilatum var porosum (Warnst) Maass 3 7

capillaceum (Weiss) Schrank (= netnoreurn) 1 1 3 cetrtrale C Jens cuspidatum Ehrh ex Hoffm 4 4 5 fiiscutn (Schimp) Klinggr 0 5 girgensohtrii Russ magellanic~im Brid 2 3 9 nlajiis (Russ) C Jens 101 papillosrttn Lindb 5 3 5 pnlcl~r~it~ (Lindb) Warnst 0 5 recnruum P Beauv (sensu lato) 3 5 ~lissowii Warnst sqliarroslml Crom slibseclmdlm~ Nees in Sturm (sensu lato)

teres (Schimp) Angstr in Hartm warnstorfii Roell wlilfianlln~ Girg

Hepatics Calypogeia sphagnicola (Arn amp Perss)

Warnst amp Loeske Cepl~fflozia media Lind b

pleniceps (Aust) Lindb Ceplraloziella sp Cladopodiella flliitatrs (Nees) Buch ~opl~ocolea hamperoplrylla ( ~ i h r a d ) Dumort Loplrozia ssp Moerkia hibernica Gott Mylia anomala (Hook) Gray PeNia epiphylla (L) Corda Pellia neesiat~a (Gott) Limpr Riccardia pit~gliis (L) Gray

Lichens Cladha rangiferit~a (L) Harm Clalpnia cetlotea (Ach) Schaer

cl~loropllaea (Florke ex Somm) Spreng coniocraea (Florke) Spreng

Mosses Alilacomniiitn pallistre (Hedw) Schwaegr 1 O Bracll~l~eci~itr rritablillim (Hedw) BSG

salebrosum (Web amp Mohr) BSG Bryum psaidotriq~ietr~itn (Hedw) Schwaegr Callicladi~im l~aldaniat~ltm (Grev) Crum Callgon corclifolilitn (Hedw) Kindb

stran~inelim (Brid) Kindb Calliergot~ella clispidata (Hedw) Loeske Can~lt~l iun~ polygamrim (BSG) C Jens

stellatlitn (Hedw) C Jens Dic~~t ir im flagellare Hedw

jirscescetrs Turn polysetlim Sw

rmdlilatlrm Schrad ex Brid Drepar~pclad~ts exannulatlrs (BSG) Warnst

fluitat~s (Hedw) Warnst Hyptrlitn itnponens Hedw

lindbergii Mitt Plagion~nilrm medium BSG Plagiothecilrm denticulatlrm (Hedw) BSG Plelrrozilrm schreberi (Brid) Mitt 0 4 Pohlia nutans (Hedw) Lindb Polytricl~lrm juniperit~~im Willd ex Hedw 0 3 Rhynchostegium serrulatum (Hedw) Jaeg amp Sauerb Tetraphis pellucida Hed w

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VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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342 CAN J BOT VOL 53 1975

TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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344 CAN J BOT VOL 53 1975

Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

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35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

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in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 8: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS 339

TABLE 2 Mean pH and cation concentrations k standard error in each of the seven community types Number of analyses in parentheses below data the number of analyses for CaZ+ is the same as for MgZ+ K + and Na A acidic edge C open mat F marginal moat H Tsuga-dominated forest M closed mat S alkaline edge T Tlr~rja-dominated

forest AC acid lake water AL alkaline lake water

A C F H M S T AC AL

seedlings are occasionally present Ericaceous shrubs are present with Andromeda glaucophylla and ChamaecIclphne calyculata reaching high importance values Rhynchospora alba (Fig 6) is the dominant sedge however since it is also an important part of the open mat community type it is not considered as an indicator species Three species of Carex are usually present C paupercula and C limosa both of which are mostly restricted to this zone and C oligosperma which is present in two other zones

~ Ten species of Sphagnum were recorded in this community type of which two form most of the cover Sphagnum cuspidatum (Fig 6) is found submerged or emergent and is the dominant species in the wettest areas while S papillosum

I becomes dominant when there is less moisture Other bryophytes are rare in this zone and un- like the alkaline lake edge zone they make up a very small portion of the vegetation When the two edge zones are compared there is almost no overlap in species composition (Table 3) The type of community which forms next to the water of a kettle-hole lake according to our data depends largely on the physical features of the lake of which the most important may be pH and cation concentrations

The two types of edge communities also differ markedly in species diversity The mean number of vascular plants for the acidic lake edge is 15 as compared with a mean of 29 for the alkaline lake edge The opposite relationship is seen for Sphagnum a mean of 37 and a total of 10 different species for the acidic lake edge but a mean of only 18 and a total of 4 species for the alkaline Much higher diversity of other bryo- phytes in the alkaline lake edge community has already been noted It can be seen here and in the other community types as well that neither the vascular plant diversity nor the Sphagnum

diversity alone nor that of other bryophytes provides a good index to the overall plant diver- sity of these communities Each segment of the community responds differently to changes in pH cation concentration etc Similar results have previously been found for bryophyte and vascu- lar plant diversity in forest communities (Slack 1971)

Sphagnum capillaceum - Carex exilis - Vaccini- um oxycoccos Community Type (Open Mat Zone C ) (Figs 20 29 32)

The open mat community is present surround- ing alkaline lakes adjacent to the edge zone It often forms extensive mats up to 20 m wide It is wet contains very few or no trees and has little topographic variation The mean pH is 49 and the Ca2+ + Mg2+ concentration is low (53 ppm Table 2) The dominant vascular plant is Vaccinium oxycoccos Habenaria blepharo- glottis Carex exilis and Scheuchzeria palustris are also characteristic parts of this community type and show high constancy

Sphagnum capillaceum (= S capillaceum var tenellum or S rubellum in this zone) is the domi- nant Sphagnum most often forming extensive pure mats Sometimes other Sphagna become im- portant parts of this zone ie S papillosum and more rarely S cuspidatum Sphagnum magellani- cum like Chamaedaphne calyculata is present to some extent in all of the community types

The open mat community type becomes well developed only in the alkaline lake bogs In Weber Lake Bog which presently has no open water and no Sphagnum teres - Carex lasiocarpa zone there is an extensive open mat community type This seemed an anomaly until we noted that ecology classes from the University of Michigan Biological Station under Dr A Cooper had mapped Weber Lake Bog in 1967 when there was

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340 CAN J BOT VOL 53 1975

TABLE 3 Species present in kettle-hole bog lakes in northern Michigan with mean importance value for each of the seven com- munity types recognized in this study Number of stands indicated in parentheses See Fig 3 for abbreviations of

community types

Sphagnum Sphqftr~itn annlilatum var porosum (Warnst) Maass 3 7

capillaceum (Weiss) Schrank (= netnoreurn) 1 1 3 cetrtrale C Jens cuspidatum Ehrh ex Hoffm 4 4 5 fiiscutn (Schimp) Klinggr 0 5 girgensohtrii Russ magellanic~im Brid 2 3 9 nlajiis (Russ) C Jens 101 papillosrttn Lindb 5 3 5 pnlcl~r~it~ (Lindb) Warnst 0 5 recnruum P Beauv (sensu lato) 3 5 ~lissowii Warnst sqliarroslml Crom slibseclmdlm~ Nees in Sturm (sensu lato)

teres (Schimp) Angstr in Hartm warnstorfii Roell wlilfianlln~ Girg

Hepatics Calypogeia sphagnicola (Arn amp Perss)

Warnst amp Loeske Cepl~fflozia media Lind b

pleniceps (Aust) Lindb Ceplraloziella sp Cladopodiella flliitatrs (Nees) Buch ~opl~ocolea hamperoplrylla ( ~ i h r a d ) Dumort Loplrozia ssp Moerkia hibernica Gott Mylia anomala (Hook) Gray PeNia epiphylla (L) Corda Pellia neesiat~a (Gott) Limpr Riccardia pit~gliis (L) Gray

Lichens Cladha rangiferit~a (L) Harm Clalpnia cetlotea (Ach) Schaer

cl~loropllaea (Florke ex Somm) Spreng coniocraea (Florke) Spreng

Mosses Alilacomniiitn pallistre (Hedw) Schwaegr 1 O Bracll~l~eci~itr rritablillim (Hedw) BSG

salebrosum (Web amp Mohr) BSG Bryum psaidotriq~ietr~itn (Hedw) Schwaegr Callicladi~im l~aldaniat~ltm (Grev) Crum Callgon corclifolilitn (Hedw) Kindb

stran~inelim (Brid) Kindb Calliergot~ella clispidata (Hedw) Loeske Can~lt~l iun~ polygamrim (BSG) C Jens

stellatlitn (Hedw) C Jens Dic~~t ir im flagellare Hedw

jirscescetrs Turn polysetlim Sw

rmdlilatlrm Schrad ex Brid Drepar~pclad~ts exannulatlrs (BSG) Warnst

fluitat~s (Hedw) Warnst Hyptrlitn itnponens Hedw

lindbergii Mitt Plagion~nilrm medium BSG Plagiothecilrm denticulatlrm (Hedw) BSG Plelrrozilrm schreberi (Brid) Mitt 0 4 Pohlia nutans (Hedw) Lindb Polytricl~lrm juniperit~~im Willd ex Hedw 0 3 Rhynchostegium serrulatum (Hedw) Jaeg amp Sauerb Tetraphis pellucida Hed w

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VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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342 CAN J BOT VOL 53 1975

TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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344 CAN J BOT VOL 53 1975

Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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CAN J BOT VOL 53 1975

0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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340 CAN J BOT VOL 53 1975

TABLE 3 Species present in kettle-hole bog lakes in northern Michigan with mean importance value for each of the seven com- munity types recognized in this study Number of stands indicated in parentheses See Fig 3 for abbreviations of

community types

Sphagnum Sphqftr~itn annlilatum var porosum (Warnst) Maass 3 7

capillaceum (Weiss) Schrank (= netnoreurn) 1 1 3 cetrtrale C Jens cuspidatum Ehrh ex Hoffm 4 4 5 fiiscutn (Schimp) Klinggr 0 5 girgensohtrii Russ magellanic~im Brid 2 3 9 nlajiis (Russ) C Jens 101 papillosrttn Lindb 5 3 5 pnlcl~r~it~ (Lindb) Warnst 0 5 recnruum P Beauv (sensu lato) 3 5 ~lissowii Warnst sqliarroslml Crom slibseclmdlm~ Nees in Sturm (sensu lato)

teres (Schimp) Angstr in Hartm warnstorfii Roell wlilfianlln~ Girg

Hepatics Calypogeia sphagnicola (Arn amp Perss)

Warnst amp Loeske Cepl~fflozia media Lind b

pleniceps (Aust) Lindb Ceplraloziella sp Cladopodiella flliitatrs (Nees) Buch ~opl~ocolea hamperoplrylla ( ~ i h r a d ) Dumort Loplrozia ssp Moerkia hibernica Gott Mylia anomala (Hook) Gray PeNia epiphylla (L) Corda Pellia neesiat~a (Gott) Limpr Riccardia pit~gliis (L) Gray

Lichens Cladha rangiferit~a (L) Harm Clalpnia cetlotea (Ach) Schaer

cl~loropllaea (Florke ex Somm) Spreng coniocraea (Florke) Spreng

Mosses Alilacomniiitn pallistre (Hedw) Schwaegr 1 O Bracll~l~eci~itr rritablillim (Hedw) BSG

salebrosum (Web amp Mohr) BSG Bryum psaidotriq~ietr~itn (Hedw) Schwaegr Callicladi~im l~aldaniat~ltm (Grev) Crum Callgon corclifolilitn (Hedw) Kindb

stran~inelim (Brid) Kindb Calliergot~ella clispidata (Hedw) Loeske Can~lt~l iun~ polygamrim (BSG) C Jens

stellatlitn (Hedw) C Jens Dic~~t ir im flagellare Hedw

jirscescetrs Turn polysetlim Sw

rmdlilatlrm Schrad ex Brid Drepar~pclad~ts exannulatlrs (BSG) Warnst

fluitat~s (Hedw) Warnst Hyptrlitn itnponens Hedw

lindbergii Mitt Plagion~nilrm medium BSG Plagiothecilrm denticulatlrm (Hedw) BSG Plelrrozilrm schreberi (Brid) Mitt 0 4 Pohlia nutans (Hedw) Lindb Polytricl~lrm juniperit~~im Willd ex Hedw 0 3 Rhynchostegium serrulatum (Hedw) Jaeg amp Sauerb Tetraphis pellucida Hed w

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VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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342 CAN J BOT VOL 53 1975

TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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344 CAN J BOT VOL 53 1975

Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

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in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

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Page 10: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS 341

TABLE 3 (Continued)

Abies balsamea (L) Mill Acer rrrbrum L Alnus rugosa (Du Roi) Spreng Andromeda glaucophylla Link Arisaema triphyllum (L) Schott Aster junciformis Rydb Betula lrrtea Michx f Bidens aristosa (Michx) Britt Boehmeria cylindrica (L) Sw Calamfigrostis canadensis (Michx) Nutt

inexpansa Gray Calla palustris L Camqpnula aparinoides Pursh

uliginosa Rydb C a ~ e x comosa Boott

exilis Dew lasiocarpa Ehrh leptalea Wahlenb limosa L oligosperma Michx paucryora Lightf pa~rpercrtla Michx trisperma Dew

Chamaedaphne calycrilata (L) Moench Cinna latifolia (Trev) Griseb Cladium mariscoides (Muhl) Torr Clintonia borealis (Ait) Raf Coptis groenlandica (Oeder) Fern Cypripedirrm acarile Ait Drqsera intermedia Hayne

I rotrrndifolia L Dryqfteris spinrilosa ( 0 F Muell) Watt

I thelypteris (L) Gray Drrlichium arrrndinaceum (L) Britt Eleacharis elliptica Kunth Epilobium leptophyllrrm Raf E r i o p F m angustifolirim Honckeny

spissrrm Fern virginicrrm L

E~rpatorirrm maculatum L Fagus grandifolia Ehrh Fraxinris nigra Marsh Galjim palustre L

trifdum L GauI~$eh hispidula (L) Bigel

procumbens L Gaylrrssacia baccata (Wang) K Koch Glyceria stricta (Lam) Hitchc Hubenaria blephariglottis (Willd) Hook Hyperic~rm virginicum L Ilex verticillata (L) Gray Iris versicolor L Juncrrs pelocarp~rs Mey Kalmia polifolia Wang Larix laricina (Du Roi) K Koch Ledum groenlandicrrm Oeder Lemna minor L Linnaea borealis L Lycoqpdium annotinum L

clavatum L inundatrrm L lucidulum Michx

Lycopus rrniporus Michx Lysimachia terrestris (L) BSP Maianthemum canadense Desf

Vascular Plants

135 104

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342 CAN J BOT VOL 53 1975

TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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344 CAN J BOT VOL 53 1975

Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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JESBERGER J A and J W SHEARD 1973 A quantitative

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

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sellschaftsmorphologie Ber Int Symp Vegetations kunde Rinteln 1966 The Hague Edited by R Tuxen pp 39-62

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35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

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in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 11: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

342 CAN J BOT VOL 53 1975

TABLE 3 (Concluded)

Medeola uirginiana L Melatnpyrum lineare Desr Menyanfhes trifoliafa L Monofropa 1mj4ora L Mr~ehlenbergia glonlerafa (Willd) Trin Nemopanfhrrs mtrcronafa (L) Trel Nuphar variegatum Engelm Nymphaea odorata Ait Osmritrda regalis L Picea mariana (Mill) BSP Pituis strobus L Pogonia ophioglossoides (L) Ker Polygonr~m amphibium L Popr~b~s tremuloides Michx Po fenfilla palr~stris (L) Sco p Pyrr~sfloribrinda Lindl Qr~ercr~s rr~bra L Rhynchospora alba (L) Vahl Rosa palrstris Marsh Rr~brts pr~bescens Raf Sagiffaria lafifolia Willd Salix pedicellaris Pursh Sarracenia purpurea L Sclier~chzeria palustris L S c i ~ r r s cyperinr~s (L) Kunth

sr~bferminalis Torr Sirm sriaue Walt Stniflcita racemosa (L) Desf

trifolia (L) Desf Thrija occidet~talis L Trienfalis borealis Raf Tsr~ga carraderrsis (L) Carr T v ~ h a tarifotia L LiicrilarA cornr~ta Michx 13

vr~lgaris L Vaccinir~nr angristifoliritn Ait

maoocarpotr Ait 133 3 5 0 6 6 4 nryrrilloides Michx 1 O 103 1 7 6 0 1 3 oxycoccos L 128 273 0 7 94 13

Vibtirtrritn cassinoides L 0 8 99 2 0 Viola ~replrropl~ylla Greene 24 21 27 Xyris nrorrtana Ries 16 0 3

Several taxonomic and nomenclatl~ral problems exist in the S capillaccroi~ group In this study we have considered S fcncrrrnr as an Atlantic- coastal-plain endemic not present in Michigan The correct name for S capi l laccur ~-r~cr~rorc~~~r~ is currently in doubt because o f nomenclature problems we have hampe followed Crum cl a (1973) and used S rapil laccur~ In all probability S ~ I ~ ~ c N I ~ I (= S rapillacc~trrr var r r ~ ~ e l l u r n ) is a distinct taxon however we have included it in the concept o f S cupillocc~rnr in this studylf it were recognized then S rrr6cllrrrrt would be present in community types A C and S while S copillorerr~n (= ro~rorcu~r) would predominate in F H and M

open water present and had found Carex lasio- carpa which is no longer present Assuming that this record is accurate Weber Lake Bog once had an alkaline edge zone which subsequent to filling in has disappeared but the S capillaceum - C exilis - Vaccinium oxycoccos zone still remains The presence at Weber Lake Bog of a remnant population of Dulickium arundinaceum char- acteristic of the alkaline edge zone gives addi- tional support to this view

As the cluster analysis shows (Fig 4) the S capillaceum - C exilis - V oxycoccos com- munity type (C) is segregated from the S cuspi- datum - S papillosum - C limosa - C pauper- cula community type (A) at about the 35 dis-

similarity level-the lowest of any of the mat community types The overlap of several impor- tant species probably explains this similarity Rhynchospora alba Carex oligosperma and Sphagnum papillosunz are found commonly in both zones The two comlnunity types do differ however in chemical properties (Table 2) and high constancy shown by several species as dis- cussed above

Sphagnum magellanicum - S recurvum - Ledurn groenlarzdicutn - Picea mariana Community Type (Closed Mat Zone M ) (Figs 16 21 22 31)

All of the bog complexes studied contained

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VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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344 CAN J BOT VOL 53 1975

Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

Huntington marsh Quebec can J Bot 51 1231-12471 BEALS E 1960 Forest bird communities in the Apostle

Islands of Wisconsin Wilson Bull 72 156-181 BEVIS F B 1960 Phytosociological study of Inverness

Mud Lake Bog Cheboygan County Michigan Pap Mich Acad Sci Arts Lett 45 61-75

BRAY J R and J T CURTIS 1957 An ordination of upland forest communities of southern Wisconsin Ecol Monogr 27 325-349

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Page 12: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS

TABLE 4 Comparison of the vascular-plant components of the acidic and alkaline edge community types

Data are given as absolute frequency

Alkaline lakes Acid lakes

Vascular plant Little Lake 16 Mud Ryerse Stutsmanville Brown Little

Chaniaedaphne calyc~rlata Andromeda gla~rcophylla Rhynchospora alba Vaccinium macrocarpon Drosera rotundifolia Carex lasiocarpa Cladilrm mariscoides Dulichiirni arundinaceum Hypericurn virginiclrm Lysimachia terrestris Pogonia ophioglossoides Potentilla palustris Viola nephrophylla Muhlenbergia glomerata Campanula aparinoides Carex palipercula Eriophorum virginicum Carex oligosperma Vaccinilrm oxycoccos Kalniia polifolia

this community type It is characterized by the presence of Picea nzariana and Larix laricina (as seedlings or mature trees) as well as a well- developed ericaceous shrub layer of Ledum groenlandicurn and Chanzaedaphne calyculata Sphagnum magellanicum and S recurvum (sensu lato) (Fig 31) are found in large amounts in this zone Topographic differences are often large with the formation of well-developed hummocks The hummocks are drier than the surrounding - hollows and consist mainly of S capillaceum (var capillaceum = S nemoreunz) and S fuscum On the dry hummock tops the mosses Aulacom- nium palustre and Pohlia nutans are often found Shade is an important factor and is due to both the tree layer and the presence of tall ericaceous shrubs Although other ericaceous shrubs have high importance values in this zone Andromeda glaucophylla is only a minor con~ponent being characteristic of the acidic edge community type

In some instances the tree layer may become quite extensive with large individuals of Picea and Larix (1 5- to 20-cm diameter at breast height (dbh)) This superficially resembles a Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) The latter is formed on mineral soil with a shallow unincorporated humus layer and has a rich feather moss carpet of Hylo- comium splendens Pleurozium schreberi and

Ptilium crista-castrensis with Sphagnum present only as isolated hummocks The closed mat zone has a thick organic soil layer and an almost con- tinuous cover of S nzagellanicztrn and S recurvum Likewise there are similarities with what might be termed a Picea mariana swamp forest with slow movement of mineral water and abundant Sphagnum species such as Sphagnum girgen- sohnii S wulJianum S russowii and S squar- rosum

Although the presence of trees in this zone is characteristic the exact importance of the tree layer in each stand is variable (Figs 10 21 22) Some stands are fairly open and much of the shade comes from the ericaceous shrubs while others are almost fully shaded by the tree layer In the former case Sphagnum magellanicum often dominates while in the latter S recurvum becomes the dominant Sphagnum Carex tri- sperma is characteristic of the more shaded portions of this zone most often occurring with S recurvum

The development of this community type seems to be independent of the type of lake and also of the surrounding soil conditions I t is characterized by low p H and low calcium and magnesium ion concentrations (Table 2) and can be considered slightly minerotrophic or possibly a n ombrotrophic community type

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344 CAN J BOT VOL 53 1975

Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

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35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

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in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 13: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

344 CAN J BOT VOL 53 1975

Ilex verticillata - Nemopanthus mucronata Com- munity Type (Marginal Moat Zone F) (Fig 23)

This community type when present lies between the bog proper and surrounding upland forest It is quite variable and further studies might serve to divide this zone into several alter- native community types In the four marginal moats we studied the dominant vascular plants were Nemopanthus mucronata Ilex verticillata Viburnum cassinoides or sometimes Osmunda regalis is dominant and a shrub canopy is lack- ing Because of the variability encountered no one species of Sphagnum can be considered characteristic However several species are com- monly found in the moat area If the moat is wet then either Sphagnum majus or Sphagnum squar- rosum may occur In drier sites Sphagnum rus- sowii S recurvum S girgensohnii and S magel- lanicum are often found The moss Pleurozium schreberi is sometimes present although it is common in many other habitats as well

The mean pH for this zone is 59 with a range between 54 and 64 which is intermediate between the extremes we recorded for the various bog habitats

Lowland Forests Lowland forests in the straits region of Michi-

gan are many and varied and are outside the scope of this paper A limited amount of samp- ling was done at the edge of several of these forests and for comparative purposes we have included our data From our observations it appears that there are three types of lowland forests in the straits region Thuja occidentalis is the dominant tree wherever a minerotrophic substrate is present while Picea mariana is dominant wherever more acidic and less minero- trophic situations occur The third type domi- nated by Tsuga canadensis is rare in the region but was present at one of our study areas (Living- ston Bog) as well as being present in upland vegetation The Picea mariana lowland forest (La Roi 1967 La Roi and Stringer 1975) was not sampled as it did not occur in any of our study areas The following two community types can be differentiated by our study

Thuja occidentalis - Sphagnum warnstorfii Community Type ( T ) (Fig 26)

This community type was sampled at Mud Lake and Ryerse Lake and was associated in

both cases with drainage areas (No pH or cation analyses were done) The vascular plants most characteristic of the sample areas were Thuja occidentalis Glyceria stria fa Rosapalustris Galium trijidum and Dryopteris thelypteris Almost all of these were to some extent also found in the Sphagnum teres - Carex lasiocarpa community type and in general indicate minero- trophic conditions

Two species of Sphagnum were characteristic S warnstorfii and S wulfianum The former is a species characteristic of minerotrophic wooded habitats the latter is found most often in wooded habitats but not necessarily minerotrophic The Thuja-dominated community type is common in the straits region but is not necessarily associated with kettle-hole lakes It contains several other Sphagnum species and a detailed study of this community type is needed

Tsuga canadensis - Sphagnum centrale Com- munity Type ( H ) (Figs 24 and 25)

Two stands dominated by Tsuga canadensis were sampled at Livingston Bog Cation and pH data (Table 2) indicate a weakly minerotrophic area Other common vascular plants in the stand were Coptis groenlandicum Gaultheria procum- bens Maianthemum canadense Osmunda regalis and Smilacina trfolia Scattered plants of Nemo- panthus mucronata and Viburnum cassinoides indicated that the stands contained elements of a marginal moat

The Sphagnum species present were typical in our experience of wooded habitats Sphagnum centrale S girgensohnii S wulfianum (Fig 25) and S squarrosum are all species of heavily shaded swamps S squarrosum perhaps is also characteristic of moat habitats

Community Segregation Indirect gradient analysis is one method of

assessing relationships between plant communi- ties and physical factors The two-dimensional ordination (Fig 3) shows environmental gradi- ents superimposed onto vegetational gradients found in the kettle-hole bog stands Three of the most important physical factors controlling bog vegetation are cation concentrations pH of the bog water and soil moisture which in kettle-hole bog community types is largely dependent on height of the vegetation above the water level

The ordination graphically represents the relationship of bog pattern to pH Ca2+ and

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VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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1 Alex W Ireland Robert K Booth Sara C Hotchkiss Jennifer E Schmitz 2012 Drought as a Trigger for Rapid State Shifts inKettle Ecosystems Implications for Ecosystem Responses to Climate Change Wetlands 32 989-1000 [CrossRef]

2 R Andersen M Poulin D Borcard R Laiho J Laine H Vasander E-T Tuittila 2011 Environmental control and spatialstructures in peatland vegetation Journal of Vegetation Science 22101111jvs201122issue-5 878-890 [CrossRef]

3 Juan J Gaitaacuten Carlos R Loacutepez Donaldo E Bran 2011 Vegetation composition and its relationship with the environment inmallines of north Patagonia Argentina Wetlands Ecology and Management 19 121-130 [CrossRef]

4 Mare Nazaire Garrett E Crow 2008 A study of the vegetation and floristic diversity of two peatland complexes of post-settlement origin in Lake Umbagog National Wildlife Refuge Coos County New Hampshire Rhodora 110 296-344 [CrossRef]

5 Jill LBubierJL Bubier Tim RMooreTR Moore GarethCrosbyG Crosby 2006 Fine-scale vegetation distribution in a cooltemperate peatland Canadian Journal of Botany 846 910-923 [Abstract] [Full Text] [PDF] [PDF Plus]

6 DI Siegel PH Glaser J So DR Janecky 2006 The dynamic balance between organic acids and circumneutral groundwaterin a large boreal peat basin Journal of Hydrology 320 421-431 [CrossRef]

7 Jake F Weltzin Jason K Keller Scott D Bridgham J Pastor P B Allen J Chen 2005 Litter controls plant communitycomposition in a northern fen Oikos 110101111oik2005110issue-3 537-546 [CrossRef]

8 Heather E Whitehouse Suzanne E Bayley 2005 Vegetation patterns and biodiversity of peatland plant communities surroundingmid-boreal wetland ponds in Alberta Canada Canadian Journal of Botany 836 621-637 [Abstract] [PDF] [PDF Plus]

9 L D Gignac R Gauthier L Rochefort J Bubier 2004 Distribution and habitat niches of 37 peatland Cyperaceae species acrossa broad geographic range in Canada Canadian Journal of Botany 829 1292-1313 [Abstract] [PDF] [PDF Plus]

10 P Oacutedor T Standovaacuter 2002 Substrate specificity and community structure of bryophyte vegetation in a near-natural montanebeech forest Community Ecology 3 39-49 [CrossRef]

11 Michal Haacutejek Petr Hekera Petra Haacutejkovaacute 2002 Spring fen vegetation and water chemistry in the Western Carpathian flyschzone Folia Geobotanica 37 205-224 [CrossRef]

12 Brian R Hall Dudley J Raynal Donald J Leopold 2001 Environmental influences on plant species composition in ground-water seeps in the Catskill Mountains of New York Wetlands 21 125-134 [CrossRef]

13 Barbara L Bedford Mark R Walbridge Allison Aldous 1999 PATTERNS IN NUTRIENT AVAILABILITY AND PLANTDIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS Ecology 80 2151-2169 [CrossRef]

14 L Bragazza 1999 Spatial patterns of plant species in a poor mire on the Southern Alps (Italy) Plant Biosystems - An InternationalJournal Dealing with all Aspects of Plant Biology 133 83-92 [CrossRef]

15 Dennis S Anderson Ronald B Davis 1997 The vegetation and its environments in Maine peatlands Canadian Journal ofBotany 7510 1785-1805 [Abstract] [PDF] [PDF Plus]

16 R Diggelen WJ Molenaar AM Kooijman 1996 Vegetation succession in a floating mire in relation to management andhydrology Journal of Vegetation Science 7 809-820 [CrossRef]

17 Andrew S Fisher Gregory S Podniesinski Donald J Leopold 1996 Effects of drainage ditches on vegetation patterns inAbandoned agricultural peatlands in central New York Wetlands 16 397-409 [CrossRef]

18 B J Nicholson L D Gignac S E Bayley 1996 Peatland distibution along a north-south transect in the Mackenzie RiverBasin in relation to climatic and environmental gradients Vegetatio 126 119-133 [CrossRef]

19 Gay M Donaldson 1996 Oribatida (Acari) associated with three species of Sphagnum at Spruce Hole Bog New HampshireUSA Canadian Journal of Zoology 749 1706-1712 [Abstract] [PDF] [PDF Plus]

20 J Bradley Johnson 1996 Phytosociology and gradient analysis of a subalpine treed fen in Rocky Mountain National ParkColorado Canadian Journal of Botany 748 1203-1213 [Abstract] [PDF] [PDF Plus]

21 Joslashrn-Frode Nordbakken 1996 Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expansea numerical approach Nordic Journal of Botany 16 197-209 [CrossRef]

22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 14: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS 345

Mg2+ concentrations and moisture regime of the peat which have been fitted to the ordination from the stand data Cation and pH data are given in detail in Table 2

As the moisture gradient moves toward drier conditions with buildup of peat and a progres- sion away from the open water of the lake shade- producing species of trees and (or) high shrubs begin to appear These shade-producing species such as Picea mariana Larix laricina Thuja occidentalis and Nemopanthus mucronata in turn affect the understory If this X axis representing a moisture gradient were the only environ- mental factor considered then the concept of hydrarch succession would be upheld However other factors are of eaual if nolt of more imuor- tance in spatial distribution of bog communities

The positions of stands and communities along the X axis is related to two gradients which parallel one another pH and Ca2+ and Mg2+ concentrations Thus the Sphagnum teres- Carex lasiocarpa community type (S) is charac-

~ -

terized by high pH minerotrophic conditions and wet habitats while the Sphagnum capil- laceum - Carex exilis - Vaccinium oxycoccos type (C) is found in wet areas with low p H and slightly minerotrophic conditions

The vertical spread of community types at the right of the ordination (ie those communities near the lake edge) indicates that these types are primarily controlled by p H and (or) cation con- centrations whereas the largely horizontal spread of the remaining communities indicates primary control by peat moisture A well-lighted to well-shaded gradient largely parallels the wet to dry moisture gradient and interacts with it For example when conditions are sufficiently dry for large ericaceous shrubs the shade they pro- duce appears to eliminate Vaccinium oxycoccos exceut from less-shaded hummock summits

The results of the minimum variance cluster analysis are shown in Fig 4 If the 35 dis- similarity level is used the clusters that are formed follow the ordination in most respects The alkaline edge zone is separated as a distinct cluster The open mat with two exceptions is separated from the acidic edge zone (see previous discussion) The closed mat is largely considered as a unit Unlike the ordination however the cluster analysis considers the various surround- ing zones as one instead of three Further sampling of moat and surrounding forest zones might resolve this difference

Direct Gradient Analysis Direct gradient analysis (Whittaker 1967

1970) can be conducted by actually measuring changing environmental factors eg p H and cation concentrations in this study and also by sampling vegetation at fixed intervals along a changing complex environmental gradient eg a mountain slope These bogs show such complex gradients and along these gradients transects were laid out and quadrats were studied at 10-m intervals Figures 11 and 12 and Tables 5-8 show the distribution of the dominant Sphagnum and vascular-plant species for five of the bogs

Populations of species along environmental gradients often take the form of bell-shaped curves with each species distributed according to its own unique response to the environment thus the composition of communities changes continuously along environmental gradients and species populations may form a complex popu- lation continuum along the gradient Table 6 shows quantitative changes in species popula- tions including several species ie Chamae- daphne calyculata Vaccinium oxycoccos and Sphagnum magellanicum which are present over a large portion of the bog Figure 12 shows changes in populations of 14 important vascular- plant species across Weber Lake Bog Shapes of curves differ for some species as Vaccinium oxycoccos and Kalmia polifolia curves are about bell-shaped Other patterns are also seen as for example Carex oligosperma which peaks late but is a minor component of community types closer to the center of the bog Similarly quanti- tative changes are shown for Sphagnum species in Fig 1 1

Even though species occur as shown in these figures as a continuum of changing populations some species have their population centers close together indicating similar responses to environ- mental stimuli These can be grouped into com- munity types From our data it appears that a complex environmental gradient exists from open water to bog forest and that along this gradient there is a vegetational gradient divisible into community types (see ordination and cluster analysis Figs 3 and 4) This complex gradient includes at least four different but interacting environmental gradients which were measured in this study namely p H (Fig 13 for Little Lake 16 and Tables 5-8 for three other lakes) Ca2+ and Mg2+ concentrations (Fig 13 for Little Lake 16) shade as determined by percentage

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346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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CAN J BOT VOL 53 1975

0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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VITT AND SLACK BOG VEGETATION ANALYSIS

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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Page 15: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

346 CAN J BOT VOL 53 1975

TABLE 5

Percentage cover for major plant species along transect from lake edge to beginning of closed mat zone at Mud Lake Bog Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a

scale of 0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphagnurn reres Sphagnlrm papilloslrnl Sphagnum capillaceum Sphagnum magellaniclrm Sphagrllrm rec~rru~rrn Drrlichirrnl arrmdirlacerrm 15 Carex lasiocarpa 25 Rlgvnchospora alba Vaccitlirrrn oxycoccos Carex parrciora Merryanrhes rrifoliara Haberraria blephariglorris Scheuchzeria palrrsrris Chatnaedaphrze calycrrlata Carex oligospernza Carex Irisperma

PH 55 58 54 45 46 51 46 45 41 50 51 Shade index 0 0 0 0 0 0 1 2 0 1 0 0 0 1 5 3 Community type S S S C C C C C C C C C C M M M

TABLE 6

Percentage cover for major plant species at Ryerse Lake Bog along transect from lake edge to marginal moat Quadrats positioned every I0 m For Ca2+ and Mg2+ concentrations see Fig13 Shade based on a scale of 0-5 for total percentage

cover for trees (see text for details)

Quadrat no

Sphagnrrm reres Sphagnlrnl papillosum Sphagnlrrn capillacerrrn Sphagnrrm magellaniclrrn Sphagnum recurulrm Sphagnurn centvale Carex lasiocarpa Porenrilla palusrris Cladium mariscoides Vaccinium macrocarpon Rhynchospora alba Andromeda glalrcophylla Vaccinium oxycoccos Cl~amaedapkne calyculara Sarracenia purpurea Kalmia polifolia Ledum groenlandicum Eriophorlim virginicum Carex paucij4ora Carex oligospernla Carex trispernla Nemopanthlrs mucrorzara

pH Shade index Community type

70 50 48 68 56 0 0 1 3 3 0 0 0 5 S C C C C - M M M M M

5 + 10 + + 15 50

71 71 3 3 0 5 M M M T

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VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

Huntington marsh Quebec can J Bot 51 1231-12471 BEALS E 1960 Forest bird communities in the Apostle

Islands of Wisconsin Wilson Bull 72 156-181 BEVIS F B 1960 Phytosociological study of Inverness

Mud Lake Bog Cheboygan County Michigan Pap Mich Acad Sci Arts Lett 45 61-75

BRAY J R and J T CURTIS 1957 An ordination of upland forest communities of southern Wisconsin Ecol Monogr 27 325-349

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

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9-31 [CrossRef]

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Page 16: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS 347

TABLE 7

Percentage cover for major plant species at Stutsmanville Bog along transect from lake edge to marginal moat Quadrats positioned every 10 m pH readings were taken from the water within the quadrats Shade based on a scale of

0-5 for total percentage cover of trees (see text for details)

Quadrat no

Sphampnlinz ~r~agellnnic~~nr Sphagnlmz recrirulim Sphagnrinr r~isso+vii Rhynchospora alba Carex pal~perclila Carex oligosperma Picea tnariana Carex trispertna Vaccit~i~nn osj~coccos Vib~irnum cassirzoides Clzamaedaphne calyc~ilata Kalmia polifolia Led~inz groerzlandicm~z

pH Shade index Communitv tvoe

51 49 0 0 0 5

A A-M M M

47 47 46 46 3 4 5 4 0 3 3

M M M M M M F

TABLE 8

Percentage cover for the 16 major plant species at Little Lake 16 along transect from edge of lake to marginal moat Quadrats placed every 10 m For Sphagtz~in~ distribution see Fig 11 for pH and CaZ+ and MgZ+ concentra-

tions see Fig 13 Shade based on a scale of 0-5 for total percentage cover of trees

Quadrat no

1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4

I Carex lasiocarpa 10 Cladi~nn mariscoides 40 Vaccinirirr~ macrocarpon 10 Sche~ichzeriapal~istris + Rhynchospora alba 5 20 15 20 Carex pa~icipora 10 Atzdromeda glaricophylla + 5 + 15 5 25 Vaccitzi~irn oxycoccos + + 10 10 5 30 35 15 20 + 5 Chamaedaphne calyc~ilata 20 40 + + 50 60 70 25 75 75 75 65 Sarracenia p~irpurea 5 + 5 + + Kalmiapolifolia 5 + + + + 5 + 5 5 5 + Led~im groerzlandic~im 5 5 + 5 Eriopphor~it~z virginiclitn + 5 15 5 5 + Carex exilis + 10 40 40 50 20 + 10 + + Nernopanthlis mlicronata 15 10 Shade index 0 1 0 0 0 0 1 3 3 3 0 0 3 3 Community type S C C C C C C C C - M M M M M - F F

cover of tree species (based on a scale of 0-5 changes in Sphagnum and vascular-plant species 1 = 1 4 2 = 5-9 3 = 10-29 4 = 3049 populations the probable importance of each 5 = 50-100 (Tables 5-8)) Height of the of these factors singly or in combination in hollows of the Sphagnutn above the water level limiting the distribution of each of the species was measured and shown indirectly on the ordi- within a bog is discussed below nation (Fig 3) The gradients have been com- The succession of dominant Sphagnum species pared (in meters from the waters edge) with in alkaline lake bogs (Mud Lake Little Lake 16

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CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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CAN J BOT VOL 53 1975

0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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VITT AND SLACK BOG VEGETATION ANALYSIS

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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Page 17: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

CAN J BOT VOL 53 1975

sphgm ter 0 c 1iarpo 0

Sphognum ruspldatum 0 mhynchap~ro olb- 0

FIGS 5-9 Importance of the five major Sphagtrritn species and four major vascular-plant species plotted over stand position on the two-dimensional ordination A solid dot is placed over a stand from which a species is absent Circles refer to Sphagtutni species squares to vascular plants Circle and square size indicates impor- tance value of the species in a stand Largest circlesquare = IV gt 50 second largest circlesquare = IV 3 0 4 9 third largest circlesquare = IV 10-29 smallest circlesquare = IV lt 10

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FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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TOLONEN K 1967 Uber die Entwicklung der Moore im finnischen Nordkarelien Ann Bot Fenn 4 219416

TRANSEAU E N 1905-06 The bogs and bog flora of the Huron River Valley Bot Gaz40 251-37541844841 1742

Voss E G 1956 A history of floristics in the Douglas Lake Region (Emmet and Cheboygan Counties) Mich- igan with an account of rejected records J Sci Lab Denison Univ 44 16-75

WALKER B H and R T COUPLAND 1968 An analysis of vegetation-environment relationships in Saskatch- ewan sloughs Can J Bot 46 509-522

WHITTAKER R H 1948 A vegetational analysis of the Great Smoky Mountains PhD Thesis University of Illinois Urbana Illinois

1956 Vegetation of the Great Smoky Mountains Ecol Monogr 26 1-80

1967 Gradient analysis Biol Rev 49 207-264 1970 Population structure of vegetation In Ge-

sellschaftsmorphologie Ber Int Symp Vegetations kunde Rinteln 1966 The Hague Edited by R Tuxen pp 39-62

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26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

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30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

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41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 18: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS

FIGS 7 and 8

Ryerse Lake) beginning at the waters edge is (Fig 13) between the S teres - C lasiocar~a ( S ) Sphagnum teres S papillosum S capillaceum and S capillaceum - C exilis - V oxycoccos (C) S magellanicum S recurvum S rnajus S zones as is the boundary between the two com- centrale (Fig 11 for Little Lake 16) although one munity types However the changes of pH and or more of these species may be absent from a cation concentration between the open mat (C) particular bog The changes in pH (Figs 13 and and closed mat (M) are often gradual and over- 14) and in cation concentration are abrupt lapping species populations are often present

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CAN J BOT VOL 53 1975

0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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VITT AND SLACK BOG VEGETATION ANALYSIS

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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KATZ N J 1926 Sphagnitrn bogs in central Russia phytosociology ecology and succession J Ecol 14 177-202

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studies in the Boreal Spruce-Fir forests of the North American Taiga 11 Analysis of the bryophyte flora

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NICHOLS G E 1919 Raised bogs in eastern Maine Geogr Rev 7 159-167

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35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

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in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 19: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

CAN J BOT VOL 53 1975

0 3 0 - 5 0 IV

0 lt 3 0 IV NONE

FIG 10 Shade in terms of importance values of trees and tall shrubs plotted over stand position on the two-dimensional ordination A solid dot is placed in those stands where no trees o r tall shrubs were found Species included are Abies balsamea Acer robrum Alnlrs nrgosa Betula l~rtea Fagus grandifolia Fraxinrrs nigra Ilex verticillata Larix laricina Netnopanthus mucronata Picea mariana Pinus strobus Popuhrs tremll- loides Pyrus Joribunda Querclrs rubra Thuja occidentalis Tsuga canadensis and Viblrr~~lrm cassinoides

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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Page 20: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS

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352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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9 L D Gignac R Gauthier L Rochefort J Bubier 2004 Distribution and habitat niches of 37 peatland Cyperaceae species acrossa broad geographic range in Canada Canadian Journal of Botany 829 1292-1313 [Abstract] [PDF] [PDF Plus]

10 P Oacutedor T Standovaacuter 2002 Substrate specificity and community structure of bryophyte vegetation in a near-natural montanebeech forest Community Ecology 3 39-49 [CrossRef]

11 Michal Haacutejek Petr Hekera Petra Haacutejkovaacute 2002 Spring fen vegetation and water chemistry in the Western Carpathian flyschzone Folia Geobotanica 37 205-224 [CrossRef]

12 Brian R Hall Dudley J Raynal Donald J Leopold 2001 Environmental influences on plant species composition in ground-water seeps in the Catskill Mountains of New York Wetlands 21 125-134 [CrossRef]

13 Barbara L Bedford Mark R Walbridge Allison Aldous 1999 PATTERNS IN NUTRIENT AVAILABILITY AND PLANTDIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS Ecology 80 2151-2169 [CrossRef]

14 L Bragazza 1999 Spatial patterns of plant species in a poor mire on the Southern Alps (Italy) Plant Biosystems - An InternationalJournal Dealing with all Aspects of Plant Biology 133 83-92 [CrossRef]

15 Dennis S Anderson Ronald B Davis 1997 The vegetation and its environments in Maine peatlands Canadian Journal ofBotany 7510 1785-1805 [Abstract] [PDF] [PDF Plus]

16 R Diggelen WJ Molenaar AM Kooijman 1996 Vegetation succession in a floating mire in relation to management andhydrology Journal of Vegetation Science 7 809-820 [CrossRef]

17 Andrew S Fisher Gregory S Podniesinski Donald J Leopold 1996 Effects of drainage ditches on vegetation patterns inAbandoned agricultural peatlands in central New York Wetlands 16 397-409 [CrossRef]

18 B J Nicholson L D Gignac S E Bayley 1996 Peatland distibution along a north-south transect in the Mackenzie RiverBasin in relation to climatic and environmental gradients Vegetatio 126 119-133 [CrossRef]

19 Gay M Donaldson 1996 Oribatida (Acari) associated with three species of Sphagnum at Spruce Hole Bog New HampshireUSA Canadian Journal of Zoology 749 1706-1712 [Abstract] [PDF] [PDF Plus]

20 J Bradley Johnson 1996 Phytosociology and gradient analysis of a subalpine treed fen in Rocky Mountain National ParkColorado Canadian Journal of Botany 748 1203-1213 [Abstract] [PDF] [PDF Plus]

21 Joslashrn-Frode Nordbakken 1996 Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expansea numerical approach Nordic Journal of Botany 16 197-209 [CrossRef]

22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 21: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

352 CAN J BOT

Distribution of Important Sphagnum and Vas- cular Plant Species

The distributions of the five major species of Sphagnutn in kettle-hole bog communities in northern Michigan are plotted over the ordina- tion analysis ( ~ i ~ s 5-9) In any one community one or sometimes two species are dominant If there are two dominant species they are often spatially separated because of hummock-hollow development and vertical wet-dry gradients present in the microtopography of the stand

The feature that most strongly joins stands of any one community type together is the Sphag- num layer The vascular plants support these community type distinctions The three most important components of the vascular plants are (1) ericaceous shrubs (2) sedges and (3) the tree laver Distributions of four of the dominant vascular plants (two ericaceous species and two sedges) plus the total tree layer are plotted over the ordination analysis (Figs 5-79 and 10) and in three cases compared directly with closely associated Sphagnum species

An estimate of the amount of shade in each stand was made by plotting the importance values of the tree species over the ordination (Fig 10) as well as percentage cover on a scale of 0-5 in the perpendicular transects (Tables 5-8) The amount of shade present in the stands is related to the height of the Sphagnum (or peat surface) above the water level Thus it is often difficult to determine if moisture shade or both are among the controlling factors of plant distributions along the horizontal or X axis of the ordination

Sphagnum teres is most abundant in open non- shaded habitats where the calcium ion concentra- tion averages 7-8 ppm and the pH is between 60 and 70 (Fig 5) The extensive floating mat sites which seem to occur only surrounding alka- line lakes are wet and when walked on they sink rapidly Apparently the alkaline conditions inhibit all other Sphagnum species (except S warnstorfii which is a shade-tolerant species) and little direct competition between S teres and other Sphagnum species is present In almost all situations (Fig 5) Sphagnum teres is found with Carex lasiocarpa These two species form an important part of one of the most distinct com- munity types in vegetation surrounding northern Michigan kettle-hole lakes

Sphagnum capillaceum (var tenellum) and Vaccinium oxycoccos occur together (Fig 7) in

VOL 53 1975

OOJ f

ANGUSTIFOLIUM d P 30 yl

gt LARlX LARlClNA

0 OLIGOSPERMA PICEA MARIANA

20 CAREX LIMOSA

SCHEUCHZERIA

10

0 0 20 40 60 80 100 120 140

METERS FROM C E N T E R O F BOG

FIG 12 Smoothed curves of percentage cover of the major vascular plant species along a transect from the center of the bog to bog forest edge at Weber Lake Bog

open non-shaded habitats which have low pH and cation concentrations (Table 2 for open mat) Both are of major importance in com- munity types which are wet and have little hum- mock development However both species occur as minor constituents of hummocky stands In this case S capillaceum (var capillaceutn) occurs near or at the to^ of 40- to 80-cm hummocks under dry conditions as opposed to its occur- rence in small amounts in the acidic edge zone under emergent or almost submerged conditions In all these situations there is very little shade (compare Figs 7 and 10) and S capillaceum appears to be restricted by low light conditions and not by moisture

In northern Michigan fire is a factor in bog development (Gates 1942 and others) Fire often removes the trees of the closed mat zone without completely destroying the Sphagnum and shrub layers An example of a portion of a closed mat that has been burned is found east of Mud Lake In this instance the hummocks are highly devel- oped and the dominant shrubs are Vaccinium angustifolium and Chamaedaphne calyculata Sphagnum fuscum and S capillaceum become more common than they are in the S magellani- cum - S recurvum closed mat Vaccinium oxy- coccos is also primarily light-limited and reaches

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VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

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in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 22: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS

80 16

LW ( 5 I C I M I F

40 1 L O 0 2 0 4 0 6 0 80 100 120

METERS FROM WATERS EDGE

FIG 13 pH values (solid circles and line) and Ca2+ plus Mg2+ concentrations (open circles and broken line) along a transect from open water to the marginal moat a t Little Lake 16 Data were taken on August 18 1971 and in each case are the result of single water samples LW lake water S alkaline edge community type C open mat community type M closed mat community type and F marginal moat community type

A l LW

FIG 14 Mean values of pH from each of the com- munity types with standard deviation shown by longest line and standard error indicated by the inner vertical marks Al LW alkaline bog lake water S alkaline edge zone C open mat zone Ac LW acidic bog lake water A acidic edge zone M closed mat zone F marginal moat zone

-a-

Ac LW

its best development and highest productivity either in the S capillaceur7l - C exilis zone or on

--

tops of hummocks in the closed mat zone Another less-fertile growth form of V oxycoccos with larger flatter leaves is often found in more shaded conditions This shade form of Vacciniunz oxycoccos may have a parallel in Sphagnum capillaceum in that there is a hemi-isophyllous form of S capillaceum that may be a shade expression of the type variety

Sphagnum cuspidatum is found predominantly in wet (usually submerged) conditions which have a low p H and low cation concentration (Fig 6 and Table 2) It forms extensive mats in the acidic edge community type and when hollows are present which are below the water level in the closed mat zone it sometimes occurs there It is rarely if ever present under minero- trophic conditions where it is replaced by S majus Rhynchospora alba a sedge which is dominant in both the acidic edge and open mat community types has a similar distribution (Fig 6) In addition to being found in large quantities in community types which are charac- terized by weakly minerotrophic conditions it often occurs in lesser amounts in the alkaline edge community (Fig 6) From this information

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354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

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Page 23: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

354 CAN J BOT VOL 53 1975

FIG 15 Livingston Bog View over open water with beginning of closed mat zone visible in foreground The acidic edge zone is narrow and often present only in scattered areas FIG 16 Stut~manville Bog View over open water with well-developed closed mat zone in background FIG 17 Little Lake 16 Bog An alkaline lake bog with a well-developed alkaline edge zone dominated by Carex lasiocarpa and Cladium mariscoides FIG 18 Mud Lake Bog An alkaline lake bog with an edge zone dominated by Carex lasiocarpa Isolated stands of Typha larifolia are present Submerged is a large zone of Nymphaea odorata FIG 19 The acidic edge community at Stutsmanville Bog The dominant Sphagna are S cuspidarum and Spapillosum FIG 20 The open mat community at Mud Lake dominated by Sphagnum capillaceunz and Scheuchzeriapalustris Also present are Sarracenia purpurea and Habernaria blephariglottis

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VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

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36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

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in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 24: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS 355

FIG 21 The closed mat community at Little Lake 16 Bog with scattered individuals o f Larix laricina and Picea mariana The dominant shrub is Chamaedaphne calyculata FIG 22 The closed mat community at Weber Lake Bog with Larix laricina and Picea mariana as the dominant trees and Chamaedaphne calyculata as the dominant shrub FIG 23 The marginal moat at Little Lake 16 Bog dominated by several tall shrub species including IIex verticillata Viburnum cassinoides and Nemopanthus mucronata FIG 24 A hemlock(Tsuga canadensis)-dominated forest at Livingston Bog with Dryopteris spinulosa and various hardwood seedlings in the herb layer FIG 25 Sphagnum wulfianum with Coptis groenlandicum interspersed from the hemlock- dominated forest at Livingston Bog FIG 26 Thuja occidentalis- dominated bog forest on south side of Mud Lake Bog

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356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

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in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 25: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

356 CAN J BOT VOL 53 1975

it seems that Rhynchospora alba is more affected by moisture level or shade than by water chem- istry while all of these environmental parameters are important in controlling the distribution of S cuspidatum

Sphagnum magellanicum is a very common species in Sphagnum-dominated communities in boreal North America As Fig 8 shows it is found in most of the stands sampled but it only becomes dominant in stands positioned in the closed mat community type It grows on low hummocks or on the sides of large hummocks in areas of intermediate moisture and light (com- pare Figs 3 and 8) No vascular plant was found to correlate exactly with its distribution although Ledum groenlandicum would come close

Sphagnum recuruum a codominant with S magellanicum in the closed mat zone is an important species found in hollows and depres- sions It has wide tolerances for pH but only becomes dominant under conditions of low p H and low calcium and magnesium ion concen- trations (Figs 3 and 9) Within the closed mat zone its distribution appears to be controlled by two interacting parameters ie shade and moisture It is restricted to depressions just below Sphagnum magellanicum in the vertical (hummock-hollow) distribution pattern From comparison of Figs 3 8 and 9 and Table 7 it can be seen that as one progresses toward drier conditions and more shade S recuruum becomes more abundant and S magellanicum less abund- ant Thus it appears S recurvum has a high tolerance for shade much more so than S magel- lanicum which has higher importance values in open shrub-dominated areas It also might be suggested that one critical factor relating to the vertical distribution of these two species is water- holding ability Sphagnum recuruum holds much less water and retains it for a much shorter period of time than does S magellanicum This might well be related to the microtopographic distribution of S recuruum in hollows and S magellanicum on hummocks It also correlates

with the horizontal distribution with S recuruum growing in wet hollows in the mesophytic shaded portions of bogs where hummocks are not well developed while S magellanicum is most impor- tant in the shrub-dominated mostly open portion of the closed mat zone where conditions on the hummocks are drier and less shaded In this latter situation S recuruum occurs in the more shaded wet hollows

Carex trisperma is often associated with S recurvum (Fig 9) particularly in the highly shaded stands of the closed mat Its distribution seems to be shade-controlled It becomes more prominent whenever Larix and Picea begin to form an important part of the vegetation It is found in the marginal moat (as is S recuruum) and beyond the bog in the moist hemlock woods bordering Livingston Bog In New York state bogs it has been found in the closed mat zone with Sphagnum recurvum and in that area also appears to be a constant associate of the more shaded portions of the closed mat community type

In conclusion there are a number of species associations a few of which have been noted between particular vascular plants and particular species of Sphagnum These associations are a result of similar physiological tolerances of these species pairs resulting in similar distributions within the bog Where conditions are quite rigorous for either group for example high pH for Sphagnum the association may be very close Sphagnum teres and Carex lasiocarpa were always found together in this study although the impor- tance values of the species vary On the other hand where several factors interact in determin- ing a species distribution in the bog and this is the usual case there is no reason to expect com- plete association Some species pairs such as S cuspidatum and Rhynchospora alba may occur together in some areas in which both have high importance values they have similar physiologi- cal tolerances in these areas However each occurs separately in other areas of the bog

FIG 27 Alkaline edge community at Mud Lake Bog The dominant plants are Sphagnum teres Carex lasio- carpa Potentilla palrrstris and Vaccinir~m macrocarpon FIG 28 The acidic edge community at Stutsmanville Bog The dominant plants are Sphanum papillosrtm and Rhynchospora alba FIG 29 The open mat com- munity at Little Lake 16 Bog The dominant plants are Sphagnum capillacerrm and S papillosum Vaccinirrm oxycoccos Scheuchzeria palustris and Andromeda glarrcophylla FIG 30 The center of Weber Lake Bog The closed mat is visible towards the rear of the picture with trees gradually invading In the foreground is the open mat community FIG 31 Sphagnrrtn recrrruum and S magellanicum at Stutsmanville Bog These two Sphagna are the dominants in the closed mat community type Chamaedaphne calycrrlata is the dominant vascular plant FIG 32 The open mat community at Mud Lake Bog with Sphagnrrm capillacerrm Rl~ynchospora alba Andromeda glaucopl~ylla Habenaria blepl~ariglottis and Vaccinirrm oxycoccos dominating

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

Huntington marsh Quebec can J Bot 51 1231-12471 BEALS E 1960 Forest bird communities in the Apostle

Islands of Wisconsin Wilson Bull 72 156-181 BEVIS F B 1960 Phytosociological study of Inverness

Mud Lake Bog Cheboygan County Michigan Pap Mich Acad Sci Arts Lett 45 61-75

BRAY J R and J T CURTIS 1957 An ordination of upland forest communities of southern Wisconsin Ecol Monogr 27 325-349

CATENHUISEN J 1950 Secondary successions on the peatlands of glacial Lake Wisconsin Trans Wis Acad Sci Arts Lett 40 29-48

CONWAY V 1949 The bogs of central Minnesota Ecol Monogr 19(2) 173-206

COOPER W S 1913 The climax forest of Isle Royale Lake Superior and its development I Bot Gaz 55 1-44 115-140

CRUM H A W C STEERE and L ANDERSON 1973 A new list of mosses of North America North of Mexico Bryologist 76 85-130

CURTIS J T and R P MCINTOSH 1951 An upland forest continuum in the prairie-forest border region of Wiscon- sin Ecology 32 476-496

DANSEREAU P and F SEGADES-VIANNA 1952 Ecologi- cal study of the peat bogs of eastern North America I Structure and evolution of vegetation Can J Bot 30 490-520

DIRSCHL H J and R T COUPLAND 1972 Vegetation patterns and site relationships in the Saskatchewan River delta Can J Bot 50 647-676

EUROLA S 1962 Uber die regionale Einteilung der Siidfinnschen Moore Ann Bot Soc Zool Bot Fenn Vanamo 33 1-243

FERNALD M L 1950 Grays manual of botany 8th ed Amer Book Co NY pp 1-1632

GATES F C 1912 The vegetation of the region in the vicinity ofDouglasLake CheboyganCounty Michigan 191 1 Rep Mich Acad Sci 14 46-106

1926 Evaporation in vegetation at different heights Am J Bot 13 167-178

1942 The bogs of northern Lower Michigan Ecol Monogr 12(3) 2 13-254

GAUCH H G and R H W H I ~ A K E R 1972 Comparison of ordination techniques Ecology 53 868-875

HALE M E and W L CULBERSON 1970 A fourth checklist of the lichens of the continental United States and Canada Bryologist 73 499-543

HEINSELMAN H L 1963 Forest sites bog processes and peatland types in the Glacial Lake Agassiz Region Min- nesota Ecol Monogr 33 327-374

15 String bogs and other patterned organic ter- rain near Seney Upper Michigan Ecology 46 185-188 - 1970 Landscape evolution peatland types and the

environment in the Lake Agassiz peatlands Natural Area Minnesota Ecol Monogr 40 235-261

JANSSEN C R 1967 A floristic study of forests and bog vegetation northwestern Minnesota Ecology 48 751-765

JEGLUM J K 1971 Plant indicators of pH and water level in peatlands at Candle Lake Saskatchewan Can J Bot 49 1661-1676

JESBERGER J A and J W SHEARD 1973 A quantitative

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

study and multivariate analysis of corticolous lichen communities in the southern boreal forest of Saskatch- ewan Can J Bot 51 185-201

KATZ N J 1926 Sphagnitrn bogs in central Russia phytosociology ecology and succession J Ecol 14 177-202

KULCZYNSKI S 1949 TorfowiskaPolesia Mem LAcad Pol Sci Lett CI Sci Math Nat Ser B Sci Nat 15 1-356

LA ROI G H 1967 Ecological studies in the Boreal Spruce-Fir forests of the North American Taiga I Analvsis of the vascular flora Ecol Monogr 37 229-253

- LA Roi G H and M H L STRINGER 1975 Ecological

studies in the Boreal Spruce-Fir forests of the North American Taiga 11 Analysis of the bryophyte flora

MALMER N 1962 Studies on mire vegetation in the Ar- chaean area of southwestern Gotaland (south Sweden) I Vegetation and habitat conditions on the Akhult mire Opera Bot 7 1-322

Moss E H 1953 Marsh and bog vegetation in northwest- ern Alberta Can J Bot 31 448470

NICHOLS G E 1919 Raised bogs in eastern Maine Geogr Rev 7 159-167

OSVALD H 1925 Zur Vegetation der ozeanischen Hoch- moore in Nonvegen fuhrer fur die vierte IPE Sven Vaxtsoc Sallsk Handl 7 1-106 - 1933 Vegetation of the pacific coast bogs of North

America Acta Phytogeogr Suec 5 1-32 1949 Notes on the vegetation of British and Irish

mosses Acta Phytogeogr Suec 26 1-62 PAASIO I 1933 Uber die Vegetation der Hochmoore Finn-

lands Acta For Fenn 39 1-210 PRITCHARD N M and A J B ANDERSON 1971 Obser-

vations on the use of cluster analysis in botany with an ecological example J Ecol 59 727-747

RAMENSKY L G 1930 Zur Methodik der vergleichenden Bearbeitung und Ordung von Pflanzenlisten und anderen Objekten die durch mehrere verschiedenartig wir- kende Faktoren bestimmt werden Beitr Biol Pflanz 18 269-304

ROSE F 1953 A survey of the ecology of the British lowland bogs Roc Linn Soc Lond 164 186-21 1

ROWE J S 1972 Forest regions of Canada Dep Environ Can For Ser Publ 1300 Ottawa Canada

RUUHIJARVI R 1960 Uber die reeionale Einteilune der ~~- -~

~ordfinnischen Moore Ann ~ o t ~ o c ~ o o l ~ B o t Fenn Vanamo 31 1-360

SCHUSTER R M 1953 Boreal hepaticae a manual of the liverworts ofMinnesotaand adjacent regions Am Midl Nat 49 257-684

SJORS H 1948 Mvrvegetation in Bergslagen Acta Phv- - - togeogr Suec 2 i 1-299

1950 Regional studies in North Swedish mire vege- tation Bot Not 1950 173-222

1959 Bogs and fens in the Hudson Bay Lowlands Arctic 12(1) 1-19 - 1963 Bogs and fens on Attawapiskat River north-

ern Ontario Natl Mus Can Bull 186 45-133 SLACK N G 1971 Species diversity and community

structure in bryophytes PhD Thesis State Univ NY Albany New York

TOLONEN K 1967 Uber die Entwicklung der Moore im finnischen Nordkarelien Ann Bot Fenn 4 219416

TRANSEAU E N 1905-06 The bogs and bog flora of the Huron River Valley Bot Gaz40 251-37541844841 1742

Voss E G 1956 A history of floristics in the Douglas Lake Region (Emmet and Cheboygan Counties) Mich- igan with an account of rejected records J Sci Lab Denison Univ 44 16-75

WALKER B H and R T COUPLAND 1968 An analysis of vegetation-environment relationships in Saskatch- ewan sloughs Can J Bot 46 509-522

WHITTAKER R H 1948 A vegetational analysis of the Great Smoky Mountains PhD Thesis University of Illinois Urbana Illinois

1956 Vegetation of the Great Smoky Mountains Ecol Monogr 26 1-80

1967 Gradient analysis Biol Rev 49 207-264 1970 Population structure of vegetation In Ge-

sellschaftsmorphologie Ber Int Symp Vegetations kunde Rinteln 1966 The Hague Edited by R Tuxen pp 39-62

-

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29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

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35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

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41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 26: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

Huntington marsh Quebec can J Bot 51 1231-12471 BEALS E 1960 Forest bird communities in the Apostle

Islands of Wisconsin Wilson Bull 72 156-181 BEVIS F B 1960 Phytosociological study of Inverness

Mud Lake Bog Cheboygan County Michigan Pap Mich Acad Sci Arts Lett 45 61-75

BRAY J R and J T CURTIS 1957 An ordination of upland forest communities of southern Wisconsin Ecol Monogr 27 325-349

CATENHUISEN J 1950 Secondary successions on the peatlands of glacial Lake Wisconsin Trans Wis Acad Sci Arts Lett 40 29-48

CONWAY V 1949 The bogs of central Minnesota Ecol Monogr 19(2) 173-206

COOPER W S 1913 The climax forest of Isle Royale Lake Superior and its development I Bot Gaz 55 1-44 115-140

CRUM H A W C STEERE and L ANDERSON 1973 A new list of mosses of North America North of Mexico Bryologist 76 85-130

CURTIS J T and R P MCINTOSH 1951 An upland forest continuum in the prairie-forest border region of Wiscon- sin Ecology 32 476-496

DANSEREAU P and F SEGADES-VIANNA 1952 Ecologi- cal study of the peat bogs of eastern North America I Structure and evolution of vegetation Can J Bot 30 490-520

DIRSCHL H J and R T COUPLAND 1972 Vegetation patterns and site relationships in the Saskatchewan River delta Can J Bot 50 647-676

EUROLA S 1962 Uber die regionale Einteilung der Siidfinnschen Moore Ann Bot Soc Zool Bot Fenn Vanamo 33 1-243

FERNALD M L 1950 Grays manual of botany 8th ed Amer Book Co NY pp 1-1632

GATES F C 1912 The vegetation of the region in the vicinity ofDouglasLake CheboyganCounty Michigan 191 1 Rep Mich Acad Sci 14 46-106

1926 Evaporation in vegetation at different heights Am J Bot 13 167-178

1942 The bogs of northern Lower Michigan Ecol Monogr 12(3) 2 13-254

GAUCH H G and R H W H I ~ A K E R 1972 Comparison of ordination techniques Ecology 53 868-875

HALE M E and W L CULBERSON 1970 A fourth checklist of the lichens of the continental United States and Canada Bryologist 73 499-543

HEINSELMAN H L 1963 Forest sites bog processes and peatland types in the Glacial Lake Agassiz Region Min- nesota Ecol Monogr 33 327-374

15 String bogs and other patterned organic ter- rain near Seney Upper Michigan Ecology 46 185-188 - 1970 Landscape evolution peatland types and the

environment in the Lake Agassiz peatlands Natural Area Minnesota Ecol Monogr 40 235-261

JANSSEN C R 1967 A floristic study of forests and bog vegetation northwestern Minnesota Ecology 48 751-765

JEGLUM J K 1971 Plant indicators of pH and water level in peatlands at Candle Lake Saskatchewan Can J Bot 49 1661-1676

JESBERGER J A and J W SHEARD 1973 A quantitative

Can

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

study and multivariate analysis of corticolous lichen communities in the southern boreal forest of Saskatch- ewan Can J Bot 51 185-201

KATZ N J 1926 Sphagnitrn bogs in central Russia phytosociology ecology and succession J Ecol 14 177-202

KULCZYNSKI S 1949 TorfowiskaPolesia Mem LAcad Pol Sci Lett CI Sci Math Nat Ser B Sci Nat 15 1-356

LA ROI G H 1967 Ecological studies in the Boreal Spruce-Fir forests of the North American Taiga I Analvsis of the vascular flora Ecol Monogr 37 229-253

- LA Roi G H and M H L STRINGER 1975 Ecological

studies in the Boreal Spruce-Fir forests of the North American Taiga 11 Analysis of the bryophyte flora

MALMER N 1962 Studies on mire vegetation in the Ar- chaean area of southwestern Gotaland (south Sweden) I Vegetation and habitat conditions on the Akhult mire Opera Bot 7 1-322

Moss E H 1953 Marsh and bog vegetation in northwest- ern Alberta Can J Bot 31 448470

NICHOLS G E 1919 Raised bogs in eastern Maine Geogr Rev 7 159-167

OSVALD H 1925 Zur Vegetation der ozeanischen Hoch- moore in Nonvegen fuhrer fur die vierte IPE Sven Vaxtsoc Sallsk Handl 7 1-106 - 1933 Vegetation of the pacific coast bogs of North

America Acta Phytogeogr Suec 5 1-32 1949 Notes on the vegetation of British and Irish

mosses Acta Phytogeogr Suec 26 1-62 PAASIO I 1933 Uber die Vegetation der Hochmoore Finn-

lands Acta For Fenn 39 1-210 PRITCHARD N M and A J B ANDERSON 1971 Obser-

vations on the use of cluster analysis in botany with an ecological example J Ecol 59 727-747

RAMENSKY L G 1930 Zur Methodik der vergleichenden Bearbeitung und Ordung von Pflanzenlisten und anderen Objekten die durch mehrere verschiedenartig wir- kende Faktoren bestimmt werden Beitr Biol Pflanz 18 269-304

ROSE F 1953 A survey of the ecology of the British lowland bogs Roc Linn Soc Lond 164 186-21 1

ROWE J S 1972 Forest regions of Canada Dep Environ Can For Ser Publ 1300 Ottawa Canada

RUUHIJARVI R 1960 Uber die reeionale Einteilune der ~~- -~

~ordfinnischen Moore Ann ~ o t ~ o c ~ o o l ~ B o t Fenn Vanamo 31 1-360

SCHUSTER R M 1953 Boreal hepaticae a manual of the liverworts ofMinnesotaand adjacent regions Am Midl Nat 49 257-684

SJORS H 1948 Mvrvegetation in Bergslagen Acta Phv- - - togeogr Suec 2 i 1-299

1950 Regional studies in North Swedish mire vege- tation Bot Not 1950 173-222

1959 Bogs and fens in the Hudson Bay Lowlands Arctic 12(1) 1-19 - 1963 Bogs and fens on Attawapiskat River north-

ern Ontario Natl Mus Can Bull 186 45-133 SLACK N G 1971 Species diversity and community

structure in bryophytes PhD Thesis State Univ NY Albany New York

TOLONEN K 1967 Uber die Entwicklung der Moore im finnischen Nordkarelien Ann Bot Fenn 4 219416

TRANSEAU E N 1905-06 The bogs and bog flora of the Huron River Valley Bot Gaz40 251-37541844841 1742

Voss E G 1956 A history of floristics in the Douglas Lake Region (Emmet and Cheboygan Counties) Mich- igan with an account of rejected records J Sci Lab Denison Univ 44 16-75

WALKER B H and R T COUPLAND 1968 An analysis of vegetation-environment relationships in Saskatch- ewan sloughs Can J Bot 46 509-522

WHITTAKER R H 1948 A vegetational analysis of the Great Smoky Mountains PhD Thesis University of Illinois Urbana Illinois

1956 Vegetation of the Great Smoky Mountains Ecol Monogr 26 1-80

1967 Gradient analysis Biol Rev 49 207-264 1970 Population structure of vegetation In Ge-

sellschaftsmorphologie Ber Int Symp Vegetations kunde Rinteln 1966 The Hague Edited by R Tuxen pp 39-62

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This article has been cited by

1 Alex W Ireland Robert K Booth Sara C Hotchkiss Jennifer E Schmitz 2012 Drought as a Trigger for Rapid State Shifts inKettle Ecosystems Implications for Ecosystem Responses to Climate Change Wetlands 32 989-1000 [CrossRef]

2 R Andersen M Poulin D Borcard R Laiho J Laine H Vasander E-T Tuittila 2011 Environmental control and spatialstructures in peatland vegetation Journal of Vegetation Science 22101111jvs201122issue-5 878-890 [CrossRef]

3 Juan J Gaitaacuten Carlos R Loacutepez Donaldo E Bran 2011 Vegetation composition and its relationship with the environment inmallines of north Patagonia Argentina Wetlands Ecology and Management 19 121-130 [CrossRef]

4 Mare Nazaire Garrett E Crow 2008 A study of the vegetation and floristic diversity of two peatland complexes of post-settlement origin in Lake Umbagog National Wildlife Refuge Coos County New Hampshire Rhodora 110 296-344 [CrossRef]

5 Jill LBubierJL Bubier Tim RMooreTR Moore GarethCrosbyG Crosby 2006 Fine-scale vegetation distribution in a cooltemperate peatland Canadian Journal of Botany 846 910-923 [Abstract] [Full Text] [PDF] [PDF Plus]

6 DI Siegel PH Glaser J So DR Janecky 2006 The dynamic balance between organic acids and circumneutral groundwaterin a large boreal peat basin Journal of Hydrology 320 421-431 [CrossRef]

7 Jake F Weltzin Jason K Keller Scott D Bridgham J Pastor P B Allen J Chen 2005 Litter controls plant communitycomposition in a northern fen Oikos 110101111oik2005110issue-3 537-546 [CrossRef]

8 Heather E Whitehouse Suzanne E Bayley 2005 Vegetation patterns and biodiversity of peatland plant communities surroundingmid-boreal wetland ponds in Alberta Canada Canadian Journal of Botany 836 621-637 [Abstract] [PDF] [PDF Plus]

9 L D Gignac R Gauthier L Rochefort J Bubier 2004 Distribution and habitat niches of 37 peatland Cyperaceae species acrossa broad geographic range in Canada Canadian Journal of Botany 829 1292-1313 [Abstract] [PDF] [PDF Plus]

10 P Oacutedor T Standovaacuter 2002 Substrate specificity and community structure of bryophyte vegetation in a near-natural montanebeech forest Community Ecology 3 39-49 [CrossRef]

11 Michal Haacutejek Petr Hekera Petra Haacutejkovaacute 2002 Spring fen vegetation and water chemistry in the Western Carpathian flyschzone Folia Geobotanica 37 205-224 [CrossRef]

12 Brian R Hall Dudley J Raynal Donald J Leopold 2001 Environmental influences on plant species composition in ground-water seeps in the Catskill Mountains of New York Wetlands 21 125-134 [CrossRef]

13 Barbara L Bedford Mark R Walbridge Allison Aldous 1999 PATTERNS IN NUTRIENT AVAILABILITY AND PLANTDIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS Ecology 80 2151-2169 [CrossRef]

14 L Bragazza 1999 Spatial patterns of plant species in a poor mire on the Southern Alps (Italy) Plant Biosystems - An InternationalJournal Dealing with all Aspects of Plant Biology 133 83-92 [CrossRef]

15 Dennis S Anderson Ronald B Davis 1997 The vegetation and its environments in Maine peatlands Canadian Journal ofBotany 7510 1785-1805 [Abstract] [PDF] [PDF Plus]

16 R Diggelen WJ Molenaar AM Kooijman 1996 Vegetation succession in a floating mire in relation to management andhydrology Journal of Vegetation Science 7 809-820 [CrossRef]

17 Andrew S Fisher Gregory S Podniesinski Donald J Leopold 1996 Effects of drainage ditches on vegetation patterns inAbandoned agricultural peatlands in central New York Wetlands 16 397-409 [CrossRef]

18 B J Nicholson L D Gignac S E Bayley 1996 Peatland distibution along a north-south transect in the Mackenzie RiverBasin in relation to climatic and environmental gradients Vegetatio 126 119-133 [CrossRef]

19 Gay M Donaldson 1996 Oribatida (Acari) associated with three species of Sphagnum at Spruce Hole Bog New HampshireUSA Canadian Journal of Zoology 749 1706-1712 [Abstract] [PDF] [PDF Plus]

20 J Bradley Johnson 1996 Phytosociology and gradient analysis of a subalpine treed fen in Rocky Mountain National ParkColorado Canadian Journal of Botany 748 1203-1213 [Abstract] [PDF] [PDF Plus]

21 Joslashrn-Frode Nordbakken 1996 Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expansea numerical approach Nordic Journal of Botany 16 197-209 [CrossRef]

22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 27: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

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VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

Huntington marsh Quebec can J Bot 51 1231-12471 BEALS E 1960 Forest bird communities in the Apostle

Islands of Wisconsin Wilson Bull 72 156-181 BEVIS F B 1960 Phytosociological study of Inverness

Mud Lake Bog Cheboygan County Michigan Pap Mich Acad Sci Arts Lett 45 61-75

BRAY J R and J T CURTIS 1957 An ordination of upland forest communities of southern Wisconsin Ecol Monogr 27 325-349

CATENHUISEN J 1950 Secondary successions on the peatlands of glacial Lake Wisconsin Trans Wis Acad Sci Arts Lett 40 29-48

CONWAY V 1949 The bogs of central Minnesota Ecol Monogr 19(2) 173-206

COOPER W S 1913 The climax forest of Isle Royale Lake Superior and its development I Bot Gaz 55 1-44 115-140

CRUM H A W C STEERE and L ANDERSON 1973 A new list of mosses of North America North of Mexico Bryologist 76 85-130

CURTIS J T and R P MCINTOSH 1951 An upland forest continuum in the prairie-forest border region of Wiscon- sin Ecology 32 476-496

DANSEREAU P and F SEGADES-VIANNA 1952 Ecologi- cal study of the peat bogs of eastern North America I Structure and evolution of vegetation Can J Bot 30 490-520

DIRSCHL H J and R T COUPLAND 1972 Vegetation patterns and site relationships in the Saskatchewan River delta Can J Bot 50 647-676

EUROLA S 1962 Uber die regionale Einteilung der Siidfinnschen Moore Ann Bot Soc Zool Bot Fenn Vanamo 33 1-243

FERNALD M L 1950 Grays manual of botany 8th ed Amer Book Co NY pp 1-1632

GATES F C 1912 The vegetation of the region in the vicinity ofDouglasLake CheboyganCounty Michigan 191 1 Rep Mich Acad Sci 14 46-106

1926 Evaporation in vegetation at different heights Am J Bot 13 167-178

1942 The bogs of northern Lower Michigan Ecol Monogr 12(3) 2 13-254

GAUCH H G and R H W H I ~ A K E R 1972 Comparison of ordination techniques Ecology 53 868-875

HALE M E and W L CULBERSON 1970 A fourth checklist of the lichens of the continental United States and Canada Bryologist 73 499-543

HEINSELMAN H L 1963 Forest sites bog processes and peatland types in the Glacial Lake Agassiz Region Min- nesota Ecol Monogr 33 327-374

15 String bogs and other patterned organic ter- rain near Seney Upper Michigan Ecology 46 185-188 - 1970 Landscape evolution peatland types and the

environment in the Lake Agassiz peatlands Natural Area Minnesota Ecol Monogr 40 235-261

JANSSEN C R 1967 A floristic study of forests and bog vegetation northwestern Minnesota Ecology 48 751-765

JEGLUM J K 1971 Plant indicators of pH and water level in peatlands at Candle Lake Saskatchewan Can J Bot 49 1661-1676

JESBERGER J A and J W SHEARD 1973 A quantitative

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

study and multivariate analysis of corticolous lichen communities in the southern boreal forest of Saskatch- ewan Can J Bot 51 185-201

KATZ N J 1926 Sphagnitrn bogs in central Russia phytosociology ecology and succession J Ecol 14 177-202

KULCZYNSKI S 1949 TorfowiskaPolesia Mem LAcad Pol Sci Lett CI Sci Math Nat Ser B Sci Nat 15 1-356

LA ROI G H 1967 Ecological studies in the Boreal Spruce-Fir forests of the North American Taiga I Analvsis of the vascular flora Ecol Monogr 37 229-253

- LA Roi G H and M H L STRINGER 1975 Ecological

studies in the Boreal Spruce-Fir forests of the North American Taiga 11 Analysis of the bryophyte flora

MALMER N 1962 Studies on mire vegetation in the Ar- chaean area of southwestern Gotaland (south Sweden) I Vegetation and habitat conditions on the Akhult mire Opera Bot 7 1-322

Moss E H 1953 Marsh and bog vegetation in northwest- ern Alberta Can J Bot 31 448470

NICHOLS G E 1919 Raised bogs in eastern Maine Geogr Rev 7 159-167

OSVALD H 1925 Zur Vegetation der ozeanischen Hoch- moore in Nonvegen fuhrer fur die vierte IPE Sven Vaxtsoc Sallsk Handl 7 1-106 - 1933 Vegetation of the pacific coast bogs of North

America Acta Phytogeogr Suec 5 1-32 1949 Notes on the vegetation of British and Irish

mosses Acta Phytogeogr Suec 26 1-62 PAASIO I 1933 Uber die Vegetation der Hochmoore Finn-

lands Acta For Fenn 39 1-210 PRITCHARD N M and A J B ANDERSON 1971 Obser-

vations on the use of cluster analysis in botany with an ecological example J Ecol 59 727-747

RAMENSKY L G 1930 Zur Methodik der vergleichenden Bearbeitung und Ordung von Pflanzenlisten und anderen Objekten die durch mehrere verschiedenartig wir- kende Faktoren bestimmt werden Beitr Biol Pflanz 18 269-304

ROSE F 1953 A survey of the ecology of the British lowland bogs Roc Linn Soc Lond 164 186-21 1

ROWE J S 1972 Forest regions of Canada Dep Environ Can For Ser Publ 1300 Ottawa Canada

RUUHIJARVI R 1960 Uber die reeionale Einteilune der ~~- -~

~ordfinnischen Moore Ann ~ o t ~ o c ~ o o l ~ B o t Fenn Vanamo 31 1-360

SCHUSTER R M 1953 Boreal hepaticae a manual of the liverworts ofMinnesotaand adjacent regions Am Midl Nat 49 257-684

SJORS H 1948 Mvrvegetation in Bergslagen Acta Phv- - - togeogr Suec 2 i 1-299

1950 Regional studies in North Swedish mire vege- tation Bot Not 1950 173-222

1959 Bogs and fens in the Hudson Bay Lowlands Arctic 12(1) 1-19 - 1963 Bogs and fens on Attawapiskat River north-

ern Ontario Natl Mus Can Bull 186 45-133 SLACK N G 1971 Species diversity and community

structure in bryophytes PhD Thesis State Univ NY Albany New York

TOLONEN K 1967 Uber die Entwicklung der Moore im finnischen Nordkarelien Ann Bot Fenn 4 219416

TRANSEAU E N 1905-06 The bogs and bog flora of the Huron River Valley Bot Gaz40 251-37541844841 1742

Voss E G 1956 A history of floristics in the Douglas Lake Region (Emmet and Cheboygan Counties) Mich- igan with an account of rejected records J Sci Lab Denison Univ 44 16-75

WALKER B H and R T COUPLAND 1968 An analysis of vegetation-environment relationships in Saskatch- ewan sloughs Can J Bot 46 509-522

WHITTAKER R H 1948 A vegetational analysis of the Great Smoky Mountains PhD Thesis University of Illinois Urbana Illinois

1956 Vegetation of the Great Smoky Mountains Ecol Monogr 26 1-80

1967 Gradient analysis Biol Rev 49 207-264 1970 Population structure of vegetation In Ge-

sellschaftsmorphologie Ber Int Symp Vegetations kunde Rinteln 1966 The Hague Edited by R Tuxen pp 39-62

-

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This article has been cited by

1 Alex W Ireland Robert K Booth Sara C Hotchkiss Jennifer E Schmitz 2012 Drought as a Trigger for Rapid State Shifts inKettle Ecosystems Implications for Ecosystem Responses to Climate Change Wetlands 32 989-1000 [CrossRef]

2 R Andersen M Poulin D Borcard R Laiho J Laine H Vasander E-T Tuittila 2011 Environmental control and spatialstructures in peatland vegetation Journal of Vegetation Science 22101111jvs201122issue-5 878-890 [CrossRef]

3 Juan J Gaitaacuten Carlos R Loacutepez Donaldo E Bran 2011 Vegetation composition and its relationship with the environment inmallines of north Patagonia Argentina Wetlands Ecology and Management 19 121-130 [CrossRef]

4 Mare Nazaire Garrett E Crow 2008 A study of the vegetation and floristic diversity of two peatland complexes of post-settlement origin in Lake Umbagog National Wildlife Refuge Coos County New Hampshire Rhodora 110 296-344 [CrossRef]

5 Jill LBubierJL Bubier Tim RMooreTR Moore GarethCrosbyG Crosby 2006 Fine-scale vegetation distribution in a cooltemperate peatland Canadian Journal of Botany 846 910-923 [Abstract] [Full Text] [PDF] [PDF Plus]

6 DI Siegel PH Glaser J So DR Janecky 2006 The dynamic balance between organic acids and circumneutral groundwaterin a large boreal peat basin Journal of Hydrology 320 421-431 [CrossRef]

7 Jake F Weltzin Jason K Keller Scott D Bridgham J Pastor P B Allen J Chen 2005 Litter controls plant communitycomposition in a northern fen Oikos 110101111oik2005110issue-3 537-546 [CrossRef]

8 Heather E Whitehouse Suzanne E Bayley 2005 Vegetation patterns and biodiversity of peatland plant communities surroundingmid-boreal wetland ponds in Alberta Canada Canadian Journal of Botany 836 621-637 [Abstract] [PDF] [PDF Plus]

9 L D Gignac R Gauthier L Rochefort J Bubier 2004 Distribution and habitat niches of 37 peatland Cyperaceae species acrossa broad geographic range in Canada Canadian Journal of Botany 829 1292-1313 [Abstract] [PDF] [PDF Plus]

10 P Oacutedor T Standovaacuter 2002 Substrate specificity and community structure of bryophyte vegetation in a near-natural montanebeech forest Community Ecology 3 39-49 [CrossRef]

11 Michal Haacutejek Petr Hekera Petra Haacutejkovaacute 2002 Spring fen vegetation and water chemistry in the Western Carpathian flyschzone Folia Geobotanica 37 205-224 [CrossRef]

12 Brian R Hall Dudley J Raynal Donald J Leopold 2001 Environmental influences on plant species composition in ground-water seeps in the Catskill Mountains of New York Wetlands 21 125-134 [CrossRef]

13 Barbara L Bedford Mark R Walbridge Allison Aldous 1999 PATTERNS IN NUTRIENT AVAILABILITY AND PLANTDIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS Ecology 80 2151-2169 [CrossRef]

14 L Bragazza 1999 Spatial patterns of plant species in a poor mire on the Southern Alps (Italy) Plant Biosystems - An InternationalJournal Dealing with all Aspects of Plant Biology 133 83-92 [CrossRef]

15 Dennis S Anderson Ronald B Davis 1997 The vegetation and its environments in Maine peatlands Canadian Journal ofBotany 7510 1785-1805 [Abstract] [PDF] [PDF Plus]

16 R Diggelen WJ Molenaar AM Kooijman 1996 Vegetation succession in a floating mire in relation to management andhydrology Journal of Vegetation Science 7 809-820 [CrossRef]

17 Andrew S Fisher Gregory S Podniesinski Donald J Leopold 1996 Effects of drainage ditches on vegetation patterns inAbandoned agricultural peatlands in central New York Wetlands 16 397-409 [CrossRef]

18 B J Nicholson L D Gignac S E Bayley 1996 Peatland distibution along a north-south transect in the Mackenzie RiverBasin in relation to climatic and environmental gradients Vegetatio 126 119-133 [CrossRef]

19 Gay M Donaldson 1996 Oribatida (Acari) associated with three species of Sphagnum at Spruce Hole Bog New HampshireUSA Canadian Journal of Zoology 749 1706-1712 [Abstract] [PDF] [PDF Plus]

20 J Bradley Johnson 1996 Phytosociology and gradient analysis of a subalpine treed fen in Rocky Mountain National ParkColorado Canadian Journal of Botany 748 1203-1213 [Abstract] [PDF] [PDF Plus]

21 Joslashrn-Frode Nordbakken 1996 Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expansea numerical approach Nordic Journal of Botany 16 197-209 [CrossRef]

22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 28: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VlTT AND SLACK BOG VEGETATION ANALYSIS 357

indicating overlapping but not identical physio- logical tolerances for these species

Several species of vascular plants notably Chamaedaphne calyculata have such broad tolerances (within the range of conditions found in bogs) that they are found in all community types and are not especially associated with particular Sphagnum species Similarly a few Sphagnum species such as S magellanicum seem to have wider ecological tolerances than most Sphagna

Relations of Physical Parameters to Bog Development

The p H and cation content of the lake water greatly influence the vegetation in the proximity of the lake but do not affect the plant com- munities which are some distance removed from the open water The p H of the water of kettle- hole lakes in the study area is variable ranging from 47 in Stutsmanville Bog to 78 in Mud

Lake Three lakes (Ryerse Lake Little Lake 16 and Mud Lake) have p H levels above 74 and are decidedly alkaline All three of these lakes

have Sphagnum mats bounded on the water side I I by a Sphagnum teres - Carex lasiocarpa com-

munity type In contrast to this four of the bogs have a waters edge community having Sphagnum cuspidatum - S papiflosum - Carex limosa - C paupercula as characteristic species (The eighth bog that was studied Weber Lake Bog con- tained no open water and is discussed below) In none of these latter four bogs is there Carex lasiocarpa or Sphagnum teres The p H of the lake water of these ranges from 47 to-65 (an occa- sional reading would register 70) and these lakes can be considered acidophilous

Surrounding the alkaline lakes a second com- munity is found which is not present around the acidophilous lakes The community has as its dominant species Sphagnum capillaceum (var tenellum) Carex exilis and Vaccinium oxycoccos The mean p H of stands having these species as dominants is 49 while that of the Sphagnum teres - dominated zone is 67 This second zone of the alkaline lakes and the Sphagnum cuspi- datum - S papillosunz - Carex limosa - C pau- percula zone of the acidophilous lakes are both followed by a third community type which is present in both types of bog lakes This zone has as its dominants Sphagnum magellanicunz S

I recurvum Picea mariana Chamaedaphne caly-

culata and Ledum groenlandicum The mean p H of stands in this zone is 52 Table 2 illustrates the mean pH values for the various zones found in the two types of lakes Figure 13 shows p H values and Ca2+ and Mg2+ concentrations for a perpendicular transect through the com- munities surrounding Little Lake 16 The most significant result of these data is to point out that the principal of hydrarch succession in kettle-hole bogs is an over simplification of the situation Bog succession is dependent on several interrelated features of which p H and cation content are important

Calcium and magnesium ion contents (Table 2) in the open mat acidic edge and closed mat community types indicate that these zones are weakly to slightly minerotrophic The alkaline edge community type can be considered minero- trophic

These data suggest that when lake water is alkaline the plant communities formed at the waters edge include plants which are found in alkaline conditions These plants act as a buffer zone to the acidophilous~communities farther from the waters edge In this situation the acido- philous species of Sphagnum have been excluded from this zone because of the influence of the high pH of the lake water In contrast in the acidophilous bog lakes the edge zone is occupied by a relatively acidic plant community In all eight bogs the horizontal distance from the waters edge to the Sphagnum recurvum - S magellanicum - Ledum groenlandicunz zone is great enough that this community is unaffected by the pH or cation content of the lake water

Extensive observations were made resulting in the finding of additional bog lakes of both types For example Rexton Lake in Mackinac County is an acid lake (pH of lake water 69) while Galloway Lake (Emmet Co) is an alkaline lake (pH of lake water 80) At East Lake in Cheboygan County which was not visited in this study students from the University of Michigan Biological Station (under Dr E Phillips mimeo- UMBS) found a mat of Carex lasiocarpa Cala- rnagrostis canadensis Dulichium arundinaceum Hypericum virginicum and Potentilla palustris plus other species found in the alkaline edge zone They also measured a pH of 80 for lake water and 55 from water in the Carex lasiocarpa zone

Intensive sampling was conducted at Weber Lake Bog which a t the present does not have

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358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

Huntington marsh Quebec can J Bot 51 1231-12471 BEALS E 1960 Forest bird communities in the Apostle

Islands of Wisconsin Wilson Bull 72 156-181 BEVIS F B 1960 Phytosociological study of Inverness

Mud Lake Bog Cheboygan County Michigan Pap Mich Acad Sci Arts Lett 45 61-75

BRAY J R and J T CURTIS 1957 An ordination of upland forest communities of southern Wisconsin Ecol Monogr 27 325-349

CATENHUISEN J 1950 Secondary successions on the peatlands of glacial Lake Wisconsin Trans Wis Acad Sci Arts Lett 40 29-48

CONWAY V 1949 The bogs of central Minnesota Ecol Monogr 19(2) 173-206

COOPER W S 1913 The climax forest of Isle Royale Lake Superior and its development I Bot Gaz 55 1-44 115-140

CRUM H A W C STEERE and L ANDERSON 1973 A new list of mosses of North America North of Mexico Bryologist 76 85-130

CURTIS J T and R P MCINTOSH 1951 An upland forest continuum in the prairie-forest border region of Wiscon- sin Ecology 32 476-496

DANSEREAU P and F SEGADES-VIANNA 1952 Ecologi- cal study of the peat bogs of eastern North America I Structure and evolution of vegetation Can J Bot 30 490-520

DIRSCHL H J and R T COUPLAND 1972 Vegetation patterns and site relationships in the Saskatchewan River delta Can J Bot 50 647-676

EUROLA S 1962 Uber die regionale Einteilung der Siidfinnschen Moore Ann Bot Soc Zool Bot Fenn Vanamo 33 1-243

FERNALD M L 1950 Grays manual of botany 8th ed Amer Book Co NY pp 1-1632

GATES F C 1912 The vegetation of the region in the vicinity ofDouglasLake CheboyganCounty Michigan 191 1 Rep Mich Acad Sci 14 46-106

1926 Evaporation in vegetation at different heights Am J Bot 13 167-178

1942 The bogs of northern Lower Michigan Ecol Monogr 12(3) 2 13-254

GAUCH H G and R H W H I ~ A K E R 1972 Comparison of ordination techniques Ecology 53 868-875

HALE M E and W L CULBERSON 1970 A fourth checklist of the lichens of the continental United States and Canada Bryologist 73 499-543

HEINSELMAN H L 1963 Forest sites bog processes and peatland types in the Glacial Lake Agassiz Region Min- nesota Ecol Monogr 33 327-374

15 String bogs and other patterned organic ter- rain near Seney Upper Michigan Ecology 46 185-188 - 1970 Landscape evolution peatland types and the

environment in the Lake Agassiz peatlands Natural Area Minnesota Ecol Monogr 40 235-261

JANSSEN C R 1967 A floristic study of forests and bog vegetation northwestern Minnesota Ecology 48 751-765

JEGLUM J K 1971 Plant indicators of pH and water level in peatlands at Candle Lake Saskatchewan Can J Bot 49 1661-1676

JESBERGER J A and J W SHEARD 1973 A quantitative

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

study and multivariate analysis of corticolous lichen communities in the southern boreal forest of Saskatch- ewan Can J Bot 51 185-201

KATZ N J 1926 Sphagnitrn bogs in central Russia phytosociology ecology and succession J Ecol 14 177-202

KULCZYNSKI S 1949 TorfowiskaPolesia Mem LAcad Pol Sci Lett CI Sci Math Nat Ser B Sci Nat 15 1-356

LA ROI G H 1967 Ecological studies in the Boreal Spruce-Fir forests of the North American Taiga I Analvsis of the vascular flora Ecol Monogr 37 229-253

- LA Roi G H and M H L STRINGER 1975 Ecological

studies in the Boreal Spruce-Fir forests of the North American Taiga 11 Analysis of the bryophyte flora

MALMER N 1962 Studies on mire vegetation in the Ar- chaean area of southwestern Gotaland (south Sweden) I Vegetation and habitat conditions on the Akhult mire Opera Bot 7 1-322

Moss E H 1953 Marsh and bog vegetation in northwest- ern Alberta Can J Bot 31 448470

NICHOLS G E 1919 Raised bogs in eastern Maine Geogr Rev 7 159-167

OSVALD H 1925 Zur Vegetation der ozeanischen Hoch- moore in Nonvegen fuhrer fur die vierte IPE Sven Vaxtsoc Sallsk Handl 7 1-106 - 1933 Vegetation of the pacific coast bogs of North

America Acta Phytogeogr Suec 5 1-32 1949 Notes on the vegetation of British and Irish

mosses Acta Phytogeogr Suec 26 1-62 PAASIO I 1933 Uber die Vegetation der Hochmoore Finn-

lands Acta For Fenn 39 1-210 PRITCHARD N M and A J B ANDERSON 1971 Obser-

vations on the use of cluster analysis in botany with an ecological example J Ecol 59 727-747

RAMENSKY L G 1930 Zur Methodik der vergleichenden Bearbeitung und Ordung von Pflanzenlisten und anderen Objekten die durch mehrere verschiedenartig wir- kende Faktoren bestimmt werden Beitr Biol Pflanz 18 269-304

ROSE F 1953 A survey of the ecology of the British lowland bogs Roc Linn Soc Lond 164 186-21 1

ROWE J S 1972 Forest regions of Canada Dep Environ Can For Ser Publ 1300 Ottawa Canada

RUUHIJARVI R 1960 Uber die reeionale Einteilune der ~~- -~

~ordfinnischen Moore Ann ~ o t ~ o c ~ o o l ~ B o t Fenn Vanamo 31 1-360

SCHUSTER R M 1953 Boreal hepaticae a manual of the liverworts ofMinnesotaand adjacent regions Am Midl Nat 49 257-684

SJORS H 1948 Mvrvegetation in Bergslagen Acta Phv- - - togeogr Suec 2 i 1-299

1950 Regional studies in North Swedish mire vege- tation Bot Not 1950 173-222

1959 Bogs and fens in the Hudson Bay Lowlands Arctic 12(1) 1-19 - 1963 Bogs and fens on Attawapiskat River north-

ern Ontario Natl Mus Can Bull 186 45-133 SLACK N G 1971 Species diversity and community

structure in bryophytes PhD Thesis State Univ NY Albany New York

TOLONEN K 1967 Uber die Entwicklung der Moore im finnischen Nordkarelien Ann Bot Fenn 4 219416

TRANSEAU E N 1905-06 The bogs and bog flora of the Huron River Valley Bot Gaz40 251-37541844841 1742

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WALKER B H and R T COUPLAND 1968 An analysis of vegetation-environment relationships in Saskatch- ewan sloughs Can J Bot 46 509-522

WHITTAKER R H 1948 A vegetational analysis of the Great Smoky Mountains PhD Thesis University of Illinois Urbana Illinois

1956 Vegetation of the Great Smoky Mountains Ecol Monogr 26 1-80

1967 Gradient analysis Biol Rev 49 207-264 1970 Population structure of vegetation In Ge-

sellschaftsmorphologie Ber Int Symp Vegetations kunde Rinteln 1966 The Hague Edited by R Tuxen pp 39-62

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 29: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

358 CAN J BOT VOL 53 1975

open water In 1967 a class survey under the the University of Michigan Biological Station direction of Dr A Cooper reported remnants Pellston Michigan of a Carex lasiocarpa mat (mimeo-UMBS) al- though at present there is no C lasiocarpa in the ALDRICH J W 1943 Biological survey of the bogs and bog In our study we found stands of Dulichium swamps of northeastern Ohio Am Midl Nat 30 arundinaceum which is characteristic of the 346-402

AUCLAIR A N A BOUCHARD and J PAJACZKOWSKI Sphagnum teres - Carex lasiocarpa zone Ca2+ 1973 Plant com~osition and s ~ e c i e s relations on the concentrations and pH values taken in the prox- imity of Dulichium were 10 ppm and 51 respec- tivelv which indicates that the lake was of the

lt lt

alkaline type Also there is a large area occupied by the open mat of S capillaceum - C exilis - Vaccinium oxycoccos community type character- istic of alkaline lakes (see ordination stand No 37 and Fig 30) Even though the two most characteristic species C lasiocarpa and S teres are not now present it is still possible to classify this bog using characteristics of the other com- munity types together with pH and Ca2+ con- centration as of the alkaline type

It is dangerous to assume that successional patterns in bogs as one sees them in a spatial sequence indicate succession in time (sensu hydrarch sere of Clements) Although we do not believe hydrarch succession can explain all types of bog and fen vegetation in northern Michigan there is direct evidence from Weber Lake Bog - that succession of communities through time has occurred Earlier data have shown that a Carex lasiocarpa community was formerly pres- ent Its presence today is indicated only by relict populations of Dulichium arundinaceum and high pH and Ca2+ concentration and by the well- developed S capillaceum mat Follow-up studies of this bog are planned they may facilitate the understanding of the structure of bogs which no longer have open water Such long-range studies as well as the present ordination and direct gradient analysis approaches should increase our understanding of the structure and function of Sphagnum bog vegetation

Acknowledgments We thank Dr Howard Crum for his critical

suggestions and help in this research Drs G Evelyn Hutchinson Richard Andrus and George La Roi read the manuscript and we are grateful for their helpful suggestions We acknowledge the National Research Council of Canada for grant A-6390 to D H Vitt in support of this research Field studies were carried out while at

Huntington marsh Quebec can J Bot 51 1231-12471 BEALS E 1960 Forest bird communities in the Apostle

Islands of Wisconsin Wilson Bull 72 156-181 BEVIS F B 1960 Phytosociological study of Inverness

Mud Lake Bog Cheboygan County Michigan Pap Mich Acad Sci Arts Lett 45 61-75

BRAY J R and J T CURTIS 1957 An ordination of upland forest communities of southern Wisconsin Ecol Monogr 27 325-349

CATENHUISEN J 1950 Secondary successions on the peatlands of glacial Lake Wisconsin Trans Wis Acad Sci Arts Lett 40 29-48

CONWAY V 1949 The bogs of central Minnesota Ecol Monogr 19(2) 173-206

COOPER W S 1913 The climax forest of Isle Royale Lake Superior and its development I Bot Gaz 55 1-44 115-140

CRUM H A W C STEERE and L ANDERSON 1973 A new list of mosses of North America North of Mexico Bryologist 76 85-130

CURTIS J T and R P MCINTOSH 1951 An upland forest continuum in the prairie-forest border region of Wiscon- sin Ecology 32 476-496

DANSEREAU P and F SEGADES-VIANNA 1952 Ecologi- cal study of the peat bogs of eastern North America I Structure and evolution of vegetation Can J Bot 30 490-520

DIRSCHL H J and R T COUPLAND 1972 Vegetation patterns and site relationships in the Saskatchewan River delta Can J Bot 50 647-676

EUROLA S 1962 Uber die regionale Einteilung der Siidfinnschen Moore Ann Bot Soc Zool Bot Fenn Vanamo 33 1-243

FERNALD M L 1950 Grays manual of botany 8th ed Amer Book Co NY pp 1-1632

GATES F C 1912 The vegetation of the region in the vicinity ofDouglasLake CheboyganCounty Michigan 191 1 Rep Mich Acad Sci 14 46-106

1926 Evaporation in vegetation at different heights Am J Bot 13 167-178

1942 The bogs of northern Lower Michigan Ecol Monogr 12(3) 2 13-254

GAUCH H G and R H W H I ~ A K E R 1972 Comparison of ordination techniques Ecology 53 868-875

HALE M E and W L CULBERSON 1970 A fourth checklist of the lichens of the continental United States and Canada Bryologist 73 499-543

HEINSELMAN H L 1963 Forest sites bog processes and peatland types in the Glacial Lake Agassiz Region Min- nesota Ecol Monogr 33 327-374

15 String bogs and other patterned organic ter- rain near Seney Upper Michigan Ecology 46 185-188 - 1970 Landscape evolution peatland types and the

environment in the Lake Agassiz peatlands Natural Area Minnesota Ecol Monogr 40 235-261

JANSSEN C R 1967 A floristic study of forests and bog vegetation northwestern Minnesota Ecology 48 751-765

JEGLUM J K 1971 Plant indicators of pH and water level in peatlands at Candle Lake Saskatchewan Can J Bot 49 1661-1676

JESBERGER J A and J W SHEARD 1973 A quantitative

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VITT AND SLACK BOG VEGETATION ANALYSIS 359

study and multivariate analysis of corticolous lichen communities in the southern boreal forest of Saskatch- ewan Can J Bot 51 185-201

KATZ N J 1926 Sphagnitrn bogs in central Russia phytosociology ecology and succession J Ecol 14 177-202

KULCZYNSKI S 1949 TorfowiskaPolesia Mem LAcad Pol Sci Lett CI Sci Math Nat Ser B Sci Nat 15 1-356

LA ROI G H 1967 Ecological studies in the Boreal Spruce-Fir forests of the North American Taiga I Analvsis of the vascular flora Ecol Monogr 37 229-253

- LA Roi G H and M H L STRINGER 1975 Ecological

studies in the Boreal Spruce-Fir forests of the North American Taiga 11 Analysis of the bryophyte flora

MALMER N 1962 Studies on mire vegetation in the Ar- chaean area of southwestern Gotaland (south Sweden) I Vegetation and habitat conditions on the Akhult mire Opera Bot 7 1-322

Moss E H 1953 Marsh and bog vegetation in northwest- ern Alberta Can J Bot 31 448470

NICHOLS G E 1919 Raised bogs in eastern Maine Geogr Rev 7 159-167

OSVALD H 1925 Zur Vegetation der ozeanischen Hoch- moore in Nonvegen fuhrer fur die vierte IPE Sven Vaxtsoc Sallsk Handl 7 1-106 - 1933 Vegetation of the pacific coast bogs of North

America Acta Phytogeogr Suec 5 1-32 1949 Notes on the vegetation of British and Irish

mosses Acta Phytogeogr Suec 26 1-62 PAASIO I 1933 Uber die Vegetation der Hochmoore Finn-

lands Acta For Fenn 39 1-210 PRITCHARD N M and A J B ANDERSON 1971 Obser-

vations on the use of cluster analysis in botany with an ecological example J Ecol 59 727-747

RAMENSKY L G 1930 Zur Methodik der vergleichenden Bearbeitung und Ordung von Pflanzenlisten und anderen Objekten die durch mehrere verschiedenartig wir- kende Faktoren bestimmt werden Beitr Biol Pflanz 18 269-304

ROSE F 1953 A survey of the ecology of the British lowland bogs Roc Linn Soc Lond 164 186-21 1

ROWE J S 1972 Forest regions of Canada Dep Environ Can For Ser Publ 1300 Ottawa Canada

RUUHIJARVI R 1960 Uber die reeionale Einteilune der ~~- -~

~ordfinnischen Moore Ann ~ o t ~ o c ~ o o l ~ B o t Fenn Vanamo 31 1-360

SCHUSTER R M 1953 Boreal hepaticae a manual of the liverworts ofMinnesotaand adjacent regions Am Midl Nat 49 257-684

SJORS H 1948 Mvrvegetation in Bergslagen Acta Phv- - - togeogr Suec 2 i 1-299

1950 Regional studies in North Swedish mire vege- tation Bot Not 1950 173-222

1959 Bogs and fens in the Hudson Bay Lowlands Arctic 12(1) 1-19 - 1963 Bogs and fens on Attawapiskat River north-

ern Ontario Natl Mus Can Bull 186 45-133 SLACK N G 1971 Species diversity and community

structure in bryophytes PhD Thesis State Univ NY Albany New York

TOLONEN K 1967 Uber die Entwicklung der Moore im finnischen Nordkarelien Ann Bot Fenn 4 219416

TRANSEAU E N 1905-06 The bogs and bog flora of the Huron River Valley Bot Gaz40 251-37541844841 1742

Voss E G 1956 A history of floristics in the Douglas Lake Region (Emmet and Cheboygan Counties) Mich- igan with an account of rejected records J Sci Lab Denison Univ 44 16-75

WALKER B H and R T COUPLAND 1968 An analysis of vegetation-environment relationships in Saskatch- ewan sloughs Can J Bot 46 509-522

WHITTAKER R H 1948 A vegetational analysis of the Great Smoky Mountains PhD Thesis University of Illinois Urbana Illinois

1956 Vegetation of the Great Smoky Mountains Ecol Monogr 26 1-80

1967 Gradient analysis Biol Rev 49 207-264 1970 Population structure of vegetation In Ge-

sellschaftsmorphologie Ber Int Symp Vegetations kunde Rinteln 1966 The Hague Edited by R Tuxen pp 39-62

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This article has been cited by

1 Alex W Ireland Robert K Booth Sara C Hotchkiss Jennifer E Schmitz 2012 Drought as a Trigger for Rapid State Shifts inKettle Ecosystems Implications for Ecosystem Responses to Climate Change Wetlands 32 989-1000 [CrossRef]

2 R Andersen M Poulin D Borcard R Laiho J Laine H Vasander E-T Tuittila 2011 Environmental control and spatialstructures in peatland vegetation Journal of Vegetation Science 22101111jvs201122issue-5 878-890 [CrossRef]

3 Juan J Gaitaacuten Carlos R Loacutepez Donaldo E Bran 2011 Vegetation composition and its relationship with the environment inmallines of north Patagonia Argentina Wetlands Ecology and Management 19 121-130 [CrossRef]

4 Mare Nazaire Garrett E Crow 2008 A study of the vegetation and floristic diversity of two peatland complexes of post-settlement origin in Lake Umbagog National Wildlife Refuge Coos County New Hampshire Rhodora 110 296-344 [CrossRef]

5 Jill LBubierJL Bubier Tim RMooreTR Moore GarethCrosbyG Crosby 2006 Fine-scale vegetation distribution in a cooltemperate peatland Canadian Journal of Botany 846 910-923 [Abstract] [Full Text] [PDF] [PDF Plus]

6 DI Siegel PH Glaser J So DR Janecky 2006 The dynamic balance between organic acids and circumneutral groundwaterin a large boreal peat basin Journal of Hydrology 320 421-431 [CrossRef]

7 Jake F Weltzin Jason K Keller Scott D Bridgham J Pastor P B Allen J Chen 2005 Litter controls plant communitycomposition in a northern fen Oikos 110101111oik2005110issue-3 537-546 [CrossRef]

8 Heather E Whitehouse Suzanne E Bayley 2005 Vegetation patterns and biodiversity of peatland plant communities surroundingmid-boreal wetland ponds in Alberta Canada Canadian Journal of Botany 836 621-637 [Abstract] [PDF] [PDF Plus]

9 L D Gignac R Gauthier L Rochefort J Bubier 2004 Distribution and habitat niches of 37 peatland Cyperaceae species acrossa broad geographic range in Canada Canadian Journal of Botany 829 1292-1313 [Abstract] [PDF] [PDF Plus]

10 P Oacutedor T Standovaacuter 2002 Substrate specificity and community structure of bryophyte vegetation in a near-natural montanebeech forest Community Ecology 3 39-49 [CrossRef]

11 Michal Haacutejek Petr Hekera Petra Haacutejkovaacute 2002 Spring fen vegetation and water chemistry in the Western Carpathian flyschzone Folia Geobotanica 37 205-224 [CrossRef]

12 Brian R Hall Dudley J Raynal Donald J Leopold 2001 Environmental influences on plant species composition in ground-water seeps in the Catskill Mountains of New York Wetlands 21 125-134 [CrossRef]

13 Barbara L Bedford Mark R Walbridge Allison Aldous 1999 PATTERNS IN NUTRIENT AVAILABILITY AND PLANTDIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS Ecology 80 2151-2169 [CrossRef]

14 L Bragazza 1999 Spatial patterns of plant species in a poor mire on the Southern Alps (Italy) Plant Biosystems - An InternationalJournal Dealing with all Aspects of Plant Biology 133 83-92 [CrossRef]

15 Dennis S Anderson Ronald B Davis 1997 The vegetation and its environments in Maine peatlands Canadian Journal ofBotany 7510 1785-1805 [Abstract] [PDF] [PDF Plus]

16 R Diggelen WJ Molenaar AM Kooijman 1996 Vegetation succession in a floating mire in relation to management andhydrology Journal of Vegetation Science 7 809-820 [CrossRef]

17 Andrew S Fisher Gregory S Podniesinski Donald J Leopold 1996 Effects of drainage ditches on vegetation patterns inAbandoned agricultural peatlands in central New York Wetlands 16 397-409 [CrossRef]

18 B J Nicholson L D Gignac S E Bayley 1996 Peatland distibution along a north-south transect in the Mackenzie RiverBasin in relation to climatic and environmental gradients Vegetatio 126 119-133 [CrossRef]

19 Gay M Donaldson 1996 Oribatida (Acari) associated with three species of Sphagnum at Spruce Hole Bog New HampshireUSA Canadian Journal of Zoology 749 1706-1712 [Abstract] [PDF] [PDF Plus]

20 J Bradley Johnson 1996 Phytosociology and gradient analysis of a subalpine treed fen in Rocky Mountain National ParkColorado Canadian Journal of Botany 748 1203-1213 [Abstract] [PDF] [PDF Plus]

21 Joslashrn-Frode Nordbakken 1996 Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expansea numerical approach Nordic Journal of Botany 16 197-209 [CrossRef]

22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 30: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

VITT AND SLACK BOG VEGETATION ANALYSIS 359

study and multivariate analysis of corticolous lichen communities in the southern boreal forest of Saskatch- ewan Can J Bot 51 185-201

KATZ N J 1926 Sphagnitrn bogs in central Russia phytosociology ecology and succession J Ecol 14 177-202

KULCZYNSKI S 1949 TorfowiskaPolesia Mem LAcad Pol Sci Lett CI Sci Math Nat Ser B Sci Nat 15 1-356

LA ROI G H 1967 Ecological studies in the Boreal Spruce-Fir forests of the North American Taiga I Analvsis of the vascular flora Ecol Monogr 37 229-253

- LA Roi G H and M H L STRINGER 1975 Ecological

studies in the Boreal Spruce-Fir forests of the North American Taiga 11 Analysis of the bryophyte flora

MALMER N 1962 Studies on mire vegetation in the Ar- chaean area of southwestern Gotaland (south Sweden) I Vegetation and habitat conditions on the Akhult mire Opera Bot 7 1-322

Moss E H 1953 Marsh and bog vegetation in northwest- ern Alberta Can J Bot 31 448470

NICHOLS G E 1919 Raised bogs in eastern Maine Geogr Rev 7 159-167

OSVALD H 1925 Zur Vegetation der ozeanischen Hoch- moore in Nonvegen fuhrer fur die vierte IPE Sven Vaxtsoc Sallsk Handl 7 1-106 - 1933 Vegetation of the pacific coast bogs of North

America Acta Phytogeogr Suec 5 1-32 1949 Notes on the vegetation of British and Irish

mosses Acta Phytogeogr Suec 26 1-62 PAASIO I 1933 Uber die Vegetation der Hochmoore Finn-

lands Acta For Fenn 39 1-210 PRITCHARD N M and A J B ANDERSON 1971 Obser-

vations on the use of cluster analysis in botany with an ecological example J Ecol 59 727-747

RAMENSKY L G 1930 Zur Methodik der vergleichenden Bearbeitung und Ordung von Pflanzenlisten und anderen Objekten die durch mehrere verschiedenartig wir- kende Faktoren bestimmt werden Beitr Biol Pflanz 18 269-304

ROSE F 1953 A survey of the ecology of the British lowland bogs Roc Linn Soc Lond 164 186-21 1

ROWE J S 1972 Forest regions of Canada Dep Environ Can For Ser Publ 1300 Ottawa Canada

RUUHIJARVI R 1960 Uber die reeionale Einteilune der ~~- -~

~ordfinnischen Moore Ann ~ o t ~ o c ~ o o l ~ B o t Fenn Vanamo 31 1-360

SCHUSTER R M 1953 Boreal hepaticae a manual of the liverworts ofMinnesotaand adjacent regions Am Midl Nat 49 257-684

SJORS H 1948 Mvrvegetation in Bergslagen Acta Phv- - - togeogr Suec 2 i 1-299

1950 Regional studies in North Swedish mire vege- tation Bot Not 1950 173-222

1959 Bogs and fens in the Hudson Bay Lowlands Arctic 12(1) 1-19 - 1963 Bogs and fens on Attawapiskat River north-

ern Ontario Natl Mus Can Bull 186 45-133 SLACK N G 1971 Species diversity and community

structure in bryophytes PhD Thesis State Univ NY Albany New York

TOLONEN K 1967 Uber die Entwicklung der Moore im finnischen Nordkarelien Ann Bot Fenn 4 219416

TRANSEAU E N 1905-06 The bogs and bog flora of the Huron River Valley Bot Gaz40 251-37541844841 1742

Voss E G 1956 A history of floristics in the Douglas Lake Region (Emmet and Cheboygan Counties) Mich- igan with an account of rejected records J Sci Lab Denison Univ 44 16-75

WALKER B H and R T COUPLAND 1968 An analysis of vegetation-environment relationships in Saskatch- ewan sloughs Can J Bot 46 509-522

WHITTAKER R H 1948 A vegetational analysis of the Great Smoky Mountains PhD Thesis University of Illinois Urbana Illinois

1956 Vegetation of the Great Smoky Mountains Ecol Monogr 26 1-80

1967 Gradient analysis Biol Rev 49 207-264 1970 Population structure of vegetation In Ge-

sellschaftsmorphologie Ber Int Symp Vegetations kunde Rinteln 1966 The Hague Edited by R Tuxen pp 39-62

-

Can

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This article has been cited by

1 Alex W Ireland Robert K Booth Sara C Hotchkiss Jennifer E Schmitz 2012 Drought as a Trigger for Rapid State Shifts inKettle Ecosystems Implications for Ecosystem Responses to Climate Change Wetlands 32 989-1000 [CrossRef]

2 R Andersen M Poulin D Borcard R Laiho J Laine H Vasander E-T Tuittila 2011 Environmental control and spatialstructures in peatland vegetation Journal of Vegetation Science 22101111jvs201122issue-5 878-890 [CrossRef]

3 Juan J Gaitaacuten Carlos R Loacutepez Donaldo E Bran 2011 Vegetation composition and its relationship with the environment inmallines of north Patagonia Argentina Wetlands Ecology and Management 19 121-130 [CrossRef]

4 Mare Nazaire Garrett E Crow 2008 A study of the vegetation and floristic diversity of two peatland complexes of post-settlement origin in Lake Umbagog National Wildlife Refuge Coos County New Hampshire Rhodora 110 296-344 [CrossRef]

5 Jill LBubierJL Bubier Tim RMooreTR Moore GarethCrosbyG Crosby 2006 Fine-scale vegetation distribution in a cooltemperate peatland Canadian Journal of Botany 846 910-923 [Abstract] [Full Text] [PDF] [PDF Plus]

6 DI Siegel PH Glaser J So DR Janecky 2006 The dynamic balance between organic acids and circumneutral groundwaterin a large boreal peat basin Journal of Hydrology 320 421-431 [CrossRef]

7 Jake F Weltzin Jason K Keller Scott D Bridgham J Pastor P B Allen J Chen 2005 Litter controls plant communitycomposition in a northern fen Oikos 110101111oik2005110issue-3 537-546 [CrossRef]

8 Heather E Whitehouse Suzanne E Bayley 2005 Vegetation patterns and biodiversity of peatland plant communities surroundingmid-boreal wetland ponds in Alberta Canada Canadian Journal of Botany 836 621-637 [Abstract] [PDF] [PDF Plus]

9 L D Gignac R Gauthier L Rochefort J Bubier 2004 Distribution and habitat niches of 37 peatland Cyperaceae species acrossa broad geographic range in Canada Canadian Journal of Botany 829 1292-1313 [Abstract] [PDF] [PDF Plus]

10 P Oacutedor T Standovaacuter 2002 Substrate specificity and community structure of bryophyte vegetation in a near-natural montanebeech forest Community Ecology 3 39-49 [CrossRef]

11 Michal Haacutejek Petr Hekera Petra Haacutejkovaacute 2002 Spring fen vegetation and water chemistry in the Western Carpathian flyschzone Folia Geobotanica 37 205-224 [CrossRef]

12 Brian R Hall Dudley J Raynal Donald J Leopold 2001 Environmental influences on plant species composition in ground-water seeps in the Catskill Mountains of New York Wetlands 21 125-134 [CrossRef]

13 Barbara L Bedford Mark R Walbridge Allison Aldous 1999 PATTERNS IN NUTRIENT AVAILABILITY AND PLANTDIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS Ecology 80 2151-2169 [CrossRef]

14 L Bragazza 1999 Spatial patterns of plant species in a poor mire on the Southern Alps (Italy) Plant Biosystems - An InternationalJournal Dealing with all Aspects of Plant Biology 133 83-92 [CrossRef]

15 Dennis S Anderson Ronald B Davis 1997 The vegetation and its environments in Maine peatlands Canadian Journal ofBotany 7510 1785-1805 [Abstract] [PDF] [PDF Plus]

16 R Diggelen WJ Molenaar AM Kooijman 1996 Vegetation succession in a floating mire in relation to management andhydrology Journal of Vegetation Science 7 809-820 [CrossRef]

17 Andrew S Fisher Gregory S Podniesinski Donald J Leopold 1996 Effects of drainage ditches on vegetation patterns inAbandoned agricultural peatlands in central New York Wetlands 16 397-409 [CrossRef]

18 B J Nicholson L D Gignac S E Bayley 1996 Peatland distibution along a north-south transect in the Mackenzie RiverBasin in relation to climatic and environmental gradients Vegetatio 126 119-133 [CrossRef]

19 Gay M Donaldson 1996 Oribatida (Acari) associated with three species of Sphagnum at Spruce Hole Bog New HampshireUSA Canadian Journal of Zoology 749 1706-1712 [Abstract] [PDF] [PDF Plus]

20 J Bradley Johnson 1996 Phytosociology and gradient analysis of a subalpine treed fen in Rocky Mountain National ParkColorado Canadian Journal of Botany 748 1203-1213 [Abstract] [PDF] [PDF Plus]

21 Joslashrn-Frode Nordbakken 1996 Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expansea numerical approach Nordic Journal of Botany 16 197-209 [CrossRef]

22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 31: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

This article has been cited by

1 Alex W Ireland Robert K Booth Sara C Hotchkiss Jennifer E Schmitz 2012 Drought as a Trigger for Rapid State Shifts inKettle Ecosystems Implications for Ecosystem Responses to Climate Change Wetlands 32 989-1000 [CrossRef]

2 R Andersen M Poulin D Borcard R Laiho J Laine H Vasander E-T Tuittila 2011 Environmental control and spatialstructures in peatland vegetation Journal of Vegetation Science 22101111jvs201122issue-5 878-890 [CrossRef]

3 Juan J Gaitaacuten Carlos R Loacutepez Donaldo E Bran 2011 Vegetation composition and its relationship with the environment inmallines of north Patagonia Argentina Wetlands Ecology and Management 19 121-130 [CrossRef]

4 Mare Nazaire Garrett E Crow 2008 A study of the vegetation and floristic diversity of two peatland complexes of post-settlement origin in Lake Umbagog National Wildlife Refuge Coos County New Hampshire Rhodora 110 296-344 [CrossRef]

5 Jill LBubierJL Bubier Tim RMooreTR Moore GarethCrosbyG Crosby 2006 Fine-scale vegetation distribution in a cooltemperate peatland Canadian Journal of Botany 846 910-923 [Abstract] [Full Text] [PDF] [PDF Plus]

6 DI Siegel PH Glaser J So DR Janecky 2006 The dynamic balance between organic acids and circumneutral groundwaterin a large boreal peat basin Journal of Hydrology 320 421-431 [CrossRef]

7 Jake F Weltzin Jason K Keller Scott D Bridgham J Pastor P B Allen J Chen 2005 Litter controls plant communitycomposition in a northern fen Oikos 110101111oik2005110issue-3 537-546 [CrossRef]

8 Heather E Whitehouse Suzanne E Bayley 2005 Vegetation patterns and biodiversity of peatland plant communities surroundingmid-boreal wetland ponds in Alberta Canada Canadian Journal of Botany 836 621-637 [Abstract] [PDF] [PDF Plus]

9 L D Gignac R Gauthier L Rochefort J Bubier 2004 Distribution and habitat niches of 37 peatland Cyperaceae species acrossa broad geographic range in Canada Canadian Journal of Botany 829 1292-1313 [Abstract] [PDF] [PDF Plus]

10 P Oacutedor T Standovaacuter 2002 Substrate specificity and community structure of bryophyte vegetation in a near-natural montanebeech forest Community Ecology 3 39-49 [CrossRef]

11 Michal Haacutejek Petr Hekera Petra Haacutejkovaacute 2002 Spring fen vegetation and water chemistry in the Western Carpathian flyschzone Folia Geobotanica 37 205-224 [CrossRef]

12 Brian R Hall Dudley J Raynal Donald J Leopold 2001 Environmental influences on plant species composition in ground-water seeps in the Catskill Mountains of New York Wetlands 21 125-134 [CrossRef]

13 Barbara L Bedford Mark R Walbridge Allison Aldous 1999 PATTERNS IN NUTRIENT AVAILABILITY AND PLANTDIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS Ecology 80 2151-2169 [CrossRef]

14 L Bragazza 1999 Spatial patterns of plant species in a poor mire on the Southern Alps (Italy) Plant Biosystems - An InternationalJournal Dealing with all Aspects of Plant Biology 133 83-92 [CrossRef]

15 Dennis S Anderson Ronald B Davis 1997 The vegetation and its environments in Maine peatlands Canadian Journal ofBotany 7510 1785-1805 [Abstract] [PDF] [PDF Plus]

16 R Diggelen WJ Molenaar AM Kooijman 1996 Vegetation succession in a floating mire in relation to management andhydrology Journal of Vegetation Science 7 809-820 [CrossRef]

17 Andrew S Fisher Gregory S Podniesinski Donald J Leopold 1996 Effects of drainage ditches on vegetation patterns inAbandoned agricultural peatlands in central New York Wetlands 16 397-409 [CrossRef]

18 B J Nicholson L D Gignac S E Bayley 1996 Peatland distibution along a north-south transect in the Mackenzie RiverBasin in relation to climatic and environmental gradients Vegetatio 126 119-133 [CrossRef]

19 Gay M Donaldson 1996 Oribatida (Acari) associated with three species of Sphagnum at Spruce Hole Bog New HampshireUSA Canadian Journal of Zoology 749 1706-1712 [Abstract] [PDF] [PDF Plus]

20 J Bradley Johnson 1996 Phytosociology and gradient analysis of a subalpine treed fen in Rocky Mountain National ParkColorado Canadian Journal of Botany 748 1203-1213 [Abstract] [PDF] [PDF Plus]

21 Joslashrn-Frode Nordbakken 1996 Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expansea numerical approach Nordic Journal of Botany 16 197-209 [CrossRef]

22 Barbara J Nicholson 1995 The wetlands of Elk Island National Park Vegetation classification water chemistry andhydrotopographic relationships Wetlands 15 119-133 [CrossRef]

23 Katherine A Frego Terry J Carleton 1995 Microsite conditions and spatial pattern in a boreal bryophyte community CanadianJournal of Botany 734 544-551 [Abstract] [PDF] [PDF Plus]

24 N R Urban E S Verry S J Eisenreich 1995 Retention and mobility of cations in a small peatland trends and mechanismsWater Air amp Soil Pollution 79 201-224 [CrossRef]

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25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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Page 32: An analysis of the vegetation of Sphagnum-dominated kettle ... · VITT, D. H., et N. G. SLACK. 1975. An analysis of the vegetation of Spllagrcirm-dominated kettle-hole bogs in relation

25 L Dennis Gignac 1994 Peatland species preferences An overview of our current knowledge base Wetlands 14 216-222[CrossRef]

26 B J Nicholson D H Vitt 1994 Wetland development at Elk Island National Park Alberta Canada Journal of Paleolimnology12 19-34 [CrossRef]

27 NANCY G SLACK 1990 Bryophytes and ecological niche theory Botanical Journal of the Linnean Society 104101111boj1990104issue-1-3 187-213 [CrossRef]

28 Rune Halvorsen Oslashkland 1990 A phytoecological study of the mire Northern Kisselbergmosen SE Norway II Identification ofgradients by detrended (canonical) correspondence analysis Nordic Journal of Botany 10 79-108 [CrossRef]

29 Terry A Hedderson Guy R Brassard 1990 Microhabitat relationships of five co-occurring saxicolous mosses on cliffs and screeslopes in eastern Newfoundland Ecography 13101111eco199013issue-2 134-142 [CrossRef]

30 M B Falbo T E Weaks 1990 A Comparison ofEichhornia crassipes (Pontederiaceae) andSphagnum quinquefarium(Sphagnaceae) in treatment of acid mine water Economic Botany 44 40-49 [CrossRef]

31 Wai -Lin Chee Dale H Vitt 1989 The vegatation surface water chemistry and peat chemistry of moderate-rich fens in centralAlberta Canada Wetlands 9 227-261 [CrossRef]

32 William Shotyk 1988 Review of the inorganic geochemistry of peats and peatland waters Earth-Science Reviews 25 95-176[CrossRef]

33 Douglas A Wilcox Howard A Simonin 1988 The stratigraphy and development of a floating peatland Pinhook Bog IndianaWetlands 8 75-91 [CrossRef]

34 Peter J Blancher Donald K McNicol 1987 Peatland water chemistry in Central Ontario in relation to acid deposition WaterAir and Soil Pollution 35 217-232 [CrossRef]

35 BARRY G WARNER PETER J BARNETT 1986 Transport sorting and reworking of late Wisconsinan plant macrofossilsfrom Lake Erie Canada Boreas 15101111bor198615issue-4 323-329 [CrossRef]

36 Douglas A Wilcox 1984 The effects of NaCl deicing salts on Sphagnum recurvum P beauv Environmental and ExperimentalBotany 24 295-304 [CrossRef]

37 Peter Alpert Walter C Oechel 1984 Microdistribution and water loss resistances of selected bryophytes in an AlaskanEriophorum tussock tundra Ecography 7101111eco19847issue-2 111-118 [CrossRef]

38 Barry G Warner Richard J Hebda Brenda J Hann 1984 Postglacial paleoecological history of a cedar swamp manitoulinisland Ontario Canada Palaeogeography Palaeoclimatology Palaeoecology 45 301-345 [CrossRef]

39 PAUL F KARROW BARRY G WARNER 1984 A subsurface Middle Wisconsinan interstadial site at Waterloo OntarioCanada Boreas 13 67-85 [CrossRef]

40 Peter Alpert Walter C Oechel 1982 Bryophyte vegetation and ecology along a topographic gradient in montane tundra inAlaska Ecography 5101111eco19825issue-2 99-108 [CrossRef]

41 Bibliography 282-304 [CrossRef]42 M S Henebry J Cairns C R Schwintzer W H Yongue 1981 A comparison of vascular vegetation and protozoan communities

in some freshwater wetlands of Northern Lower Michigan Hydrobiologia 83 353-375 [CrossRef]43 A N D Auclair A Bouchard J Pajaczkowski 1976 Productivity relations in a Carex-dominated ecosystem Oecologia 26

9-31 [CrossRef]

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