Allen Et Al. (2009) -- Palaeoenvironmetal Context of Woolly Mammoth at Candover, UK

SGEOLOGICAL JOURNAL

Geol. J. 44: 414446 (2009)

Published online in Wiley InterScience

(www.interscience.wiley.com) DOI: 10.1002/gj.1161

*Correspondence to: J. R. M. Allen, School of Biological and Biomedical Sciences, DurhamUniversity, South Road, Durham DH1 3LE, UK.E-mail: [email protected] # 2009 John Wiley & Sons, Ltd.Birmingham, UK5Tigh-na-cleirich, Foss, by Pitlochry, Perthshire, UK

In 1986/1987 the remains of several mammoths,Mammuthus primigenius (Blumenbach), were discovered on the spoil heap ofan actively working gravel pit at Condover, Shropshire, England. The discovery of the remains posed two questions that could beaddressed by analyses of biological proxies. First, as none of the bones was found in situ it was necessary to confirm the stratumin which the remains occurred. Second, what was the environment in which these animals lived and died? A range of biologicalindicators was used to address these questions, including pollen, spore and algal, plant macrofossil, invertebrate, anuran andbiological mineral analyses. Multivariate statistical analyses of palynological and Pediastrum data, along with evidence from theColeopteran assemblages, support the attribution of the mammoth bones to a unit of dark grey clayey sandy silt, although theymay have lived at the time of the overlying green detritus mud. The palaeobiological data supports the correlation of thesesediments to the Devensian Late-glacial. The mammoths entered this basin at the start of the Late-glacial Interstadial (GreenlandInterstadial 1e) (ca. 14 8303930 cal. year BP; 12 300 110 14C year BP) and became mired in soft cohesive sediments.Palaeotemperature reconstructions, based on the Coleopteran assemblages, from the time when the mammoths actually becamemired, show that the climate was temperate with mean July temperatures between 15 and 198C and mean January temperaturesbetween 13 and 68C. Biological indicators from the sediments encasing the mammoths indicate that the landscapesurrounding the basin was treeless and dry, contrasting with rich vegetation within the basin itself that had possibly attracted themammoths to the site. Evidence of sedimentary disturbance suggests that the mammoths caused large-scale bioturbation of thedeposits making palaeoenvironmental interpretations difficult. Fossils of terrestrial blowflies, carcass and dung beetles show thatsome of the decaying corpses must have lain exposed on the land surface for sufficient time for the soft parts to have rotted awayand skin and bones to have become desiccated before many of them sank into the dark grey clayey sandy silt. Copyright# 2009John Wiley & Sons, Ltd.

Received 12 February 2009; accepted 23 April 2009

KEY WORDS Devensian Late-glacial; palaeoclimate; pollen; Coleoptera; plant macrofossils; kettle-hole; Pediastrum

Supporting information may be found in the online version of this article (Supplementary Tables S1S10).

1. INTRODUCTION

In 1986 a spectacular set of remains of several Mammuthus primigenius (Blumenbach) (woolly mammoth) were

found at Norton Farm Pit, Condover, Shropshire, UK in the spoil heaps of overburden removed during quarrying

operations prior to extraction of the gravel (Coope and Lister 1987; Lister 2009). This sediment was derived from aDepartment of Archaeology, University of York, The Kings Manor, York, UKchool of Geography Earth and Environmental Sciences, University of Birmingham, Edgbaston,Palaeoenvironmental context of the Late-glacial woolly mammoth(Mammuthus primigenius) discoveries at Condover, Shropshire, UK

J. R. M. ALLEN1,2*, J. D. SCOURSE 2, A. R. HALL 3 and G. R. COOPE 4,5

1School of Biological and Biomedical Sciences, Durham University, Durham, UK2School of Ocean Sciences, College of Natural Sciences, Bangor University, Anglesey, UK

3

4

largely infilled kettle-hole (Scourse et al. 2009). Over the next few years (19861990) the site was the subject of a

series of investigations (Lister 1993). The regional setting of the site (Latitude 5283801500N; Longitude 284504600W;UK NGR SJ494073, 80m above sea level) has been described by Worsley (2005). Detailed investigations of the

palaeoenvironmental context of condover mammoth site, uk 4152.1.1. Field sampling

The samples used for pollen and Pediastrum analyses are shown in Supplementary Table S1 and the locations of the

various profiles from which the samples were taken are shown on the map of the site (Figure 1).

In 1987 Profile C (Figure 1; Face 1) was chosen as the reference section for the entire sequence infilling the

kettle-hole basin. This was because

(i) many of the identified sedimentary units (Scourse et al. 2009) could be observed in stratigraphic section in this

single profile and were accessible for sampling; and

(ii) this was where the stratigraphic unit hypothesized, during the JuneJuly 1987 sampling season, to be the

mammoth stratum (Unit C1), was thickest and most easily related to the other sedimentary units.

In order to maximize the sampled thicknesses of Units C1, D, E and F the sample sequence was divided into two

parts (Figure 2); a total of 27 samples was taken through Units C1, D, E and F at Profile C.

In addition to the samples from Profile C, a single sample of the overlying green detritus mud (Unit C3) was

collected in 1987 from Profile D (Figure 1; Face 1).

To relate the unstratified mammoth bones to the reference profile and to the other samples, three sediment

samples scraped from mammoth bones newly recovered from below the water level of the gravel pit during the

JuneJuly 1987 excavation were also prepared for pollen analysis. One of the samples was scraped from a juvenile

vertebra discovered close to the site of Borehole g (Figure 1); the other two were scraped from the juvenile jaw bones.

In January 1988, the water level in the pit was lowered by pumping and this allowed further sampling of Unit C1,

this time at Profile Z, closer to the location from where the bones had been removed during quarrying activities, to

Copyright # 2009 John Wiley & Sons, Ltd. Geol. J. 44: 414446 (2009)

DOI: 10.1002/gjas pollen (with accessory spores and algae) are one of the most useful approaches and can provide evidence of the

regional vegetation of the locality, thus enabling broad stratigraphical correlations to be made. These can be

supplemented by more local information gained from macrofossil remains of plants, principally fruits and seeds,

but also leaves and wood. Evidence from faunal remains such as Coleoptera, Trichoptera, Chironomidae and

Anurans can also make unique contributions to the palaeoenvironmental interpretation.

2. MATERIALS AND METHODS

2.1. Pollen and Pediastrumand gering to them to the radiocarbon-based age of the sediments (Scourse et al. 2009)), mineralogy, geochemistry

rainsize analysis. Because the amount of sediment available for analysis is often very small, microfossils suchbones

adhnumber of techniques can be used to provenance unstratified sediment such as that adhering to the mammoth

. This includes radiocarbon dating (i.e. by matching the radiocarbon-based age of the bones and the sedimentmammoth bones (Lister 2009), which came from an adult male and several juvenile animals, and their local

stratigraphic context (Scourse et al. 2009) have also been undertaken. Biological investigations of the sediments

including pollen, spore and algal, plant macrofossil, insect and other invertebrate, anuran (frog and toad) and fish, as

well as biological minerals, are reported here.

Samples for biological analyses were taken on a number of occasions between 1986 and 1990, and from a

number of sedimentary units, with two main objectives. First, as none of the mammoth bones was discovered

in situ, a prime aim of the investigations was to identify as accurately as possible the sedimentary sequence infilling

the kettle-hole basin, so that sediment adhering to the bones could be compared with the in situ sedimentary units

(Scourse et al. 2009) and the source of the bones identified. The second aim was to provide evidence on the nature

of the changing environment in and around the Norton Farm Pit site before, during, and after the period when the

mammoths entered the basin.

A

416 j. r. m. allen ET AL.the southsouthwest of Profiles C and D (Figure 1; Face 2). Avertical series of ten samples was taken through Unit

C1 at Profile Z; of these, only three, all situated towards the base of Unit C1, proved to contain palynomorph

concentrations in excess of 4000 grains g1 wet weight and were thus suitable for analysis. In addition, two sampleswere taken from Unit C1 at Profile Y in Face 2, further to the north and closer to Face 1 than Profile Z (Figure 1).

During May 1990 a particularly thick sequence of Units E and F was exposed by quarrying activities in the

vicinity of Profile B (Figure 1). In order to confirm a Holocene age for the black humified peat of Unit F a further

series of 28 samples was taken; in this exposure Unit F was 2.0m thick, Unit E 70 cm thick; the contact between

Unit E and Unit D also was covered in the sampling.

2.1.2. Laboratory methods

Samples for pollen analyses were initially prepared using standard treatment, including hydrofluoric (HF) acid

(Faegri and Iversen 1989), but the results were extremely disappointing. The pollen concentrations were low, and

the grains very badly corroded. An alternative method was subsequently used replacing HF digestion with a

Figure 1. Site map showing location of Profiles at Norton Farm Pit 19861990.


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palaeoenvironmental context of condover mammoth site, uk 417physical method for the removal of inorganic material. After disaggregation samples were washed over a 10mmmonofilament nylon mesh (Cwynar et al. 1979; Lowe and Walker 1986) prior to heavy liquid separation using

saturated zinc chloride (ZnCl2: density 2.0 g cm3; (Bjorck et al. 1978; Watkins et al. 2007)). These preparationswere far superior, in terms of both the concentration and preservation of the palynomorphs, as has previously been

noted for highly minerogenic samples at other sites (Scourse et al. 1992). The addition of Lycopodium (clubmoss)

spore tablets (Benninghoff 1962; Matthews 1969; Bonny 1972) to weighed subsamples of the sediment prior to

pollen extraction enabled the calculation of pollen concentrations. The pollen extracts were mounted in silicone oil

and examined using a compound light microscope at a routine magnification of400 with 1000 (oil immersion)magnification being used for the examination of critical features. For each level at least 300 identifiable land pollen

and spores were counted. Unidentifiable pollen was recorded as crumpled, broken, corroded, degraded

(amorphous) or concealed (Cushing 1967). Plant names follow Clapham et al. (1962), and the pollen and spore

types follow the schemes of Faegri and Iversen (1989) and Birks (1993); the conventions used to indicate the

certainty of identification follow Birks (1993).

The use of density separation rather than HF also enhances the recovery of Pediastrum (class: Chlorophyceae,

order Chlorococcales). The colonies of this alga form from aggregating zoospores and the species differ in the

number and form of prongs on the peripheral cells, the amount of space between the cells, and the wall sculpture

(Millington et al. 1981). Pediastrum colonies were particularly rich in these samples and it was clear during routine

Figure 2. Profile C showing relative locations of the pollen sampling sequences. Unit Fblack humified peat; Unit Egrey clay; Unit Dbrown sedge peat; Unit C1dark grey clayey sandy silt; Unit B2clast-supported gravel; Unit B1pink laminated clays and silts.


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pollen counting that a number of very distinct forms could be identified. The preservation was generally good,

although in some cases the wall sculpturing and the prongs were either eroded or broken, and there were also some

broken colonies of irregular shape. The identification of Pediastrum taxa found in these sediments follows

418 j. r. m. allen ET AL.adopted, from 1 (one or a few individuals or fragments) to 4 (an abundant component of the sample).

2.3. Insects

The samples used for insect analyses are shown in Supplementary Table S3.

The samples of sediment collected at the site were placed directly into polythene bags and kept in their field-

damp conditions until ready for laboratory processing. Standard techniques were adopted to recover insect fossils

(e.g. Coope 1968; Elias 1994). The sample was disaggregated by applying a gentle stream of water; the resultant

mixture was washed over a 300mm aperture sieve; this is sufficiently fine to retain most insect remains. The>300mm fraction was washed with water and mixed with paraffin oil. Water was then added to permit a clearseparation between the two fractions. Floating material was decanted into a 300mm sieve, washed in dilutedetergent and water to remove the oil and then in alcohol to remove the water. The insect fossils were sorted under

alcohol using a binocular microscope and stored in tubes of alcohol to prevent attack by fungi.

Identification of the fossil insect fragments was by direct comparison with specimens of modern animals. No

significant differences were found between the fossils and the modern material. By far the most common of the

recognizable fossils were of Coleoptera. Although remains of Diptera, Hymenoptera and other orders of insect were

also present, only the Trichoptera and Chironomidae have been studied in detail.


DOI: 10.1002/gjlevels was therefore used.

2.2. Plant macrofossils

The samples used for macrofossil analyses are shown in Supplementary Table S2.

Subsamples of raw sediment were broken down in water in a bucket and sieved (300mm mesh) in a stream ofwater to remove the fine sand, silt, clay and unidentifiable organic debris. In deposits such as these, where the

content of plant remains is generally quite small, it is most practicable to concentrate the organic fraction by means

of a washover, in which the disaggregated sediment is swirled in water in the bucket and the less dense material

decanted off before it settles out. The sieved plant remains were sorted under a binocular microscope and put aside

for identification, which relies on modern reference material and published accounts of modern and fossil

specimens such as Beijerinck (1947) and Katz et al. (1965). For this site a four-point scale of abundance wasJankovska and Komarek (1982) and Pascher (1913). The species/varieties identified include Pediastrum boryanum

typicum, P. boryanum longicorne, P. boryanum brevicorne, P. duplex, P. kawraiskyi, P. sturmii and P. tetras. An

indeterminable group included all broken and degraded colonies. Some of the P. boryanum brevicornemay include

small numbers of P. integrum. The frequencies are expressed as percentages of the total number of Pediastrum

counted. As different Pediastrum species have different responses to nutrients they can be used as indicators of the

trophic status of the water body. In general Pediastrum numbers increase as nutrients increase and changes in

species composition occur in connection with eutrophication or oligotrophication (Cronberg 1982).

The pollen diagram for Profile C was zoned using ZONATION (Gordon and Birks 1972) using 14 taxa (Betula

(excluding B. nana), Salix (excluding S. herbacea), Betula nana, Salix herbacea, Juniperus communis,Gramineae,Cyperaceae, Solidago type (Compositae), Artemisia, Filipendula, Myriophyllum, Nymphaea, Potamogeton andPediastrum (total)). This Fortran IV package combines three zonation procedures, CONSLINK(dendrogram for constrained single link analysis), SPLINF (dendrogram for binary division using aninformation content criterion) and SPLITSQ (dendrogram for binary division using sum-of-squarescriterion). These procedures group similar samples together, and split or separate samples with fewsimilarities. The three levels within Unit C1 at the base of the upper part of the diagram were excluded fromthis zonation procedure because they are possibly duplicated by samples from the lower part; a total of 24

2.4. Other environmental indicators

Anura, Trichoptera, Chironomidae, Ostracoda and biological mineralogical analyses all provided additional

palaeoenvironmental context of condover mammoth site, uk 419evidence relating to the environment around the time of the demise of the mammoths; four brief reports are

included in the results section.

3. RESULTS

3.1. Pollen and Pediastrum

The pollen diagram for Profile C is shown in Figure 3. Sample depths are expressed as depths below the top of the

profile. The results of the ZONATION analysis applied to Profile C are shown in Figure 4; this supports the adoption

of pollen zones which coincide with the major lithostratigraphic boundaries. Zone NF1 coincides with sedimentary

Unit C1, NF2 with Unit D, NF3 with Unit E and NF4 with the base of Unit F (Figure 2). The Profile C pollen

diagram is divided into two sections, the separation being marked by the non-continuity of the pollen curves. The

sections in which the curves are joined represent continuous stratigraphic sequences. The lower part of the diagram

relates exclusively to Unit C1, whereas the upper part covers the stratigraphic sequence from the top of Unit C1

through Units D (brown sedge peat) and E (grey clay) to the base of F (black humified peat) (Figure 2). It is probable

that the upper part of the lower sequence overlaps the three samples of Unit C1 at the base of the upper sequence.

Samples taken from the very base of the upper sequence (Figure 3), and probably from Unit B1 (pink laminated

clays and silts), proved to be devoid of pollen and are not included on the pollen diagram.

3.1.1. Profile C: pollen assemblage zone NF1 (Unit C1): GramineaeCyperaceaeArtemisia

This pollen assemblage zone (paz) consists of ten samples taken from Unit C1 in Profile C (seven from the lower

section and three from the upper section). Pollen concentrations fluctuate widely from just under 6000 grains g1

wet weight in the basal level to over 109 000 grains g1 in the highest level; this increase is masked by saw-toothvariations in the intervening levels.

The dominant taxa in this zone are Gramineae, Cyperaceae, and to a lesser extent, Artemisia. In the lower

part of the diagram Gramineae rises from just under 2% in the basal level to ca. 69% in the highest level againstwhich Cyperaceae displays a reciprocal relationship, falling from ca. 87% in the basal level to a little over 8%in the highest. Artemisia uctuates between 1 and 11%, and occurs alongside a diverse range of herbs whichare all represented at low values (

Figure

3.Pollen

percentagediagram

forProfile

C.Taxathat

never

exceed

1%

areincluded

asother

tree/shrub/herbaceous.Taxabetween1and2.5%

areshownas

dots.Pollen

sum

(SP)TLP(TotalLandPollen);CCryptogam

s;AAquatics;Iindeterminables;PedPediastrum;PPPre-Pleistocenespores.

Copyright # 2009 John Wiley & Sons, Ltd. Geol. J 44: 414446 (2009)

DOI: 10.1002/gj

420 j. r. m. allen ET AL..

curve rises to a peak of over 46% in the centre of the zone; Salix undiff. ranges between 5 and 12% and Juniperus

between 1 and 11%. Juniperus and Salix cf. S. herbacea, which has a peak of over 9% in the basal level, decline

from maxima early in the zone as Betula undiff. (mostly tree birch) increases. As the latter taxon declines towards

the end of the zone so Betula cf. B. nana increases, reaching a peak of almost 18% towards the top. As these changes

occur within a highly organic sedge peat of telmatic character they probably reflect actual changes in the local

vegetation rather than any taphonomic artifact of sedimentation or preservation.

Figure 4. Profile CResults of Zonation.

palaeoenvironmental context of condover mammoth site, uk 421Figure 5. Pediastrum percentage diagram for Profile C.


DOI: 10.1002/gj

Accompanying these changes in the tree and shrub vegetation are changes in the herbaceous community. Other

than grasses and sedges, the most consistent herb taxa in paz NF1 are Artemisia, Compositae subfamily

Liguliflorae, Arenaria type, Caryophyllaceae undiff., Chenopodiaceae and Trifolium. These are replaced in paz

422 j. r. m. allen ET AL.NF2 by Rosaceae undiff., Helianthemum, Liliaceae, Ranunculus repens type, Rubiaceae and Angelica type.

Changes also occur in the aquatic taxa. Pediastrum declines as macrophyte aquatic vegetation becomes

important. Pediastrum fluctuates between 0 and 4% in the lower part of the zone, but rises to 35% towards the top.

As in paz NF1 P. boryanum typicum dominates the Pediastrum assemblages, but P. duplex, which was entirely

absent in NF1, rises to exceed 40% of the Pediastrum spectra in the centre of the zone. Macrophyte aquatic taxa in

NF2 include Myriophyllum spp., predominantly M. spicatum (almost 12% at the base of the zone), but also

M. verticillatum and M. alterniflorum. Peaks of Potamogeton occur at the base (almost 2%) and top (26%) of the

zone, framing a continuous record of Nymphaea which reaches a peak of over 8% in the centre of the zone. Other

obligate aquatic taxa include Nuphar, Menyanthes trifoliata and Typha latifolia. NF2 is also rich in the usually

helophytic pteridophyte Equisetum which reaches almost 15% in the central part of the zone.

The low totals for indeterminate pollen, and the complete absence of dinoflagellate cysts and pre-Pleistocene

taxa, suggest minimal reworking in this zone.

3.1.3. Profile C: pollen assemblage zone NF3 (Unit E): BetulaGramineaeCyperaceaeArtemisiaPaz NF3 consists of seven samples taken from Unit E (grey clay) in Profile C. Pollen concentrations show a decline

from the high levels attained in NF2, rising from a minima of under 3000 grains g1 wet weight at the base of the zoneto over 70000 grains g1 at the top. Reworking resumes in NF3 with increases in the occurrence of pre-Pleistocene andindeterminable taxa. Though Betula undiff.,B. cf. nana, Salix undiff. and S. herbacea decline in comparison with NF2,

they nevertheless maintain a presence throughout NF3. Apart from two isolated grains, Juniperus is absent in NF3.

Gramineae and Cyperaceae increase in frequency, Gramineae ranging between 41 and 20%, and Cyperaceae between

23 and 16%. Artemisia is well represented, rising to over 20% in the central part of the zone. The helophytic herb taxon

Ranunculus trichophyllus type rises to a peak of over 25% at the top of the zone. Other significant herb taxa include

Rumex acetosa type, for which there is a continuous curve, Cirsium/Carduus, Compositae subfamily Liguliflorae,

Arenaria type, Caryophyllaceae undiff., Potentilla type and Thalictrum.

The macrophyte aquatic taxa present in NF2 are almost completely absent in NF3, with the exceptions of

sporadic occurrences ofMenyanthes trifoliata and Typha latifolia which may be re-worked from NF2. Pediastrum

increases in NF3 to reach a peak of over 71% in the top level, its highest value in the entire diagram. Once again

P. boryanum typicum dominates (up to 50% of the Pediastrum assemblage), P. boryanum longicorne reaches 25%

and there is a diverse range of other taxa, including P. boryanum brevicorne, P. duplex, P. kawraiskyi, P. sturmii and

P. tetras.

3.1.4. Profile C: pollen assemblage zone NF4 (base of Unit F): BetulaSalixJuniperusGramineaePaz NF4 is represented by the single level from Unit F (black humified peat) at the top of Profile C (Figure 3). This

level has a pollen concentration of over 68 000 grains g1 wet weight and shows some significant changes fromzone NF3 below. These include an increase in Betula undiff. to almost 22%, in Salix undiff. to almost 17%, in

Salix cf. herbacea to almost 6%, and reductions in Artemisia, and Ranunculus trichophyllus type. Juniperus rises to

over 1%, and Equisetum reaches over 6%. Pediastrum declines from over 71% in the top level of NF3 to just over

1% in NF4.

3.1.5. Palynology of Unit C3 (green detritus mud)

Unit C3 (green detritus mud), which did not occur at Profile C, lies stratigraphically between Unit C1 (pollen

assemblage zone NF1) and Unit D (pollen assemblage zone NF2 (Scourse et al. 2009). The sample of Unit C3,

collected in 1987 from Profile D (Face Y; Figure 1) yielded a pollen spectrum dominated by Cyperaceae (43.7%),

Gramineae (37%) and Ranunculus repens type (7.7%), with traces of Betula, Artemisia, Armeria/Limonium (type

A), and Thalictrum (Figure 6). The pollen concentration exceeds 35 000 grains g1 wet weight. Pediastrum,


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Figure

6.Pollen

percentagediagram

forbonescrapingandother

samplesfrom

UnitsC1andC3.Taxathatnever

exceed

1%

areincluded

asother

tree/shrub/herbaceous.Taxabetween

1and2.5%

areshownas

solidcircles.Pollen

sum(SP)TLP(TotalLandPollen);CCryptogam

s;AAquatics;Iindeterminables;PedPediastrum;PPPre-Pleistocenespores.

opyright # 2009 John Wiley & Sons, Ltd. eol. J. 44: 414446 (2009)

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palaeoenvironmental context of condover mammoth site, uk 423C G

dominated by P. boryanum typicum with subdominant P. boryanum brevicorne and P. boryanum longicorne,

reaches almost 49% (Figures 3 and 5).

3.1.6. Palynology of sediment scraped from mammoth bones

Three sediment samples scraped from juvenile mammoth bones were analysed. The sample from a vertebra

discovered close to the site of Borehole g (Figure 1) is the uppermost sample in Figures 6 and 7. The other two

samples were from scrapes of jaw bones. The two jaw samples are very similar to each other, with pollen

concentrations of 8000 grains g1 wet weight and almost 29 000 grains g1. The pollen assemblages (Figure 6) aredominated by Cyperaceae (3844%), with subdominant Gramineae (1930%), Pinus (67%), Salix (45%) and

Trifolium cf. T. repens (610%). The assemblage in the vertebra sample (pollen concentration, 19 000 grains g1

wet weight) shows some differences. Cyperaceae is even more dominant (almost 80%) and Gramineae less so

(under 10%), with Pinus (5%) and Salix (2.5%); Trifolium cf. T. repens is absent. In all three samples levels of pre-

Pleistocene and indeterminable palynomorphs are high, and reworked dinoflagellate cysts occur in the vertebra

sample. Pediastrum totals are also high in all three samples (4175%; SPPediastrum), dominated byP. boryanum typicum with subdominant P. boryanum longicorne and P. sturmii (Figure 7).

3.1.7. Palynology of Unit C1 (dark grey clayey, sandy silt) from Profiles Z and Y, Face 2

The three samples from Unit C1 at Profile Z yielded assemblages extremely similar to zone NF1 from Profile C,

also from Unit C1. Cyperaceae dominates the spectra (up to 84%), with subdominant Gramineae (up to 31%) and

Pinus (up to almost 7%). As in NF1 there are isolated occurrences of thermophilous tree and shrub taxa, including

Alnus and Corylus/Myrica, and there are significant numbers of pre-Pleistocene, indeterminable and dinoflagellate

taxa. The pollen concentrations are low, ranging between 6000 and 13 000 grains g1 wet weight. Pediastrum

424 j. r. m. allen ET AL.Figure 7. Pediastrum percentage diagram for bone scraping and other samples from Units C1 and C3.


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concentrations rise from less than 2% in the basal sample to over 66% (SPPediastrum) in the top sample,dominated by P. boryanum typicum with P. boryanum brevicorne (Figure 7).

A further two samples were taken in 1988 from Unit C1 at Profile Y in Face 2, further to the north and closer to

palaeoenvironmental context of condover mammoth site, uk 425Face 1 than Profile Z (Figure 1). The pollen concentrations and assemblages here proved to be extremely similar to

both the samples from Profile Z (Figure 6) and zone NF1 from Profile C (Figure 3). The lower sample has a pollen

concentration of nearly 9000 grains g1 wet weight, rising to almost 23 000 in the upper sample. The pollenassemblages in these two samples are dominated by Cyperaceae (5174%), with subdominant Gramineae (15

34%) and Salix (24%). High levels of reworking are indicated by significant quantities of pre-Pleistocene and

indeterminable taxa. Pediastrum rises from just over 11% (SPPediastrum) in the lower sample to almost 65% inthe upper; these consist of dominant P. boryanum typicum with P. boryanum longicorne and P. boryanum

brevicorne (Figure 7).

3.1.8. Profile B pollen diagram

The base of this pollen diagram (Figure 8) overlaps with the top of the diagram from Profile C (Figure 3), and the

upper part, by comparison with other nearby well-dated pollen diagrams (e.g. Beales 1980), covers much of the

early Holocene. Important features in this respect include the immigration of Corylus, and later, Alnus. This

sequence is, however, difficult to interpret in detail because it is dominated by pollen taxa from very local sources;

many of the observed changes are therefore probably caused by small changes in the hydrology/water level of the

basin, as might be expected in a telmatic peat. The large fluctuations in the Cyperaceae curve, and the irregularity of

the curves for the major tree taxa, can probably be explained in this way. However, these tree taxa are very poorly

represented. In particular, the very low frequencies for the mixed woodland taxa Corylus, Ulmus and Quercus are

puzzling for most other pollen records covering the early Holocene from this area (e.g. Beales 1980) record high

frequencies for these taxa. Even accounting for the local pollen catchment, the record appears to indicate minimal

mixed woodland development in the early to mid-Holocene.

The diagram can be subdivided into five pollen assemblage zones (Figure 8). Zones NF2 and NF3 correlate with

these same zones in Profile C (Figure 3), and correspond to Units D and E respectively. Zone NF4, the base of Unit

F, correlates with the uppermost sample from Profile C (Figure 3). Zone NF5 is defined by the rise in Corylus/

Myrica, but is dominated by Betula, Salix and Cyperaceae, with subdominant Pinus and Gramineae. Zone NF6 is

defined by the rise in Alnus and is represented by only one level at the top of the profile.

3.1.9. Multivariate statistical analysis of the pollen spectra

In order to assess objectively the similarity between the pollen spectra from the bone scrapes and the spectra from

the stratigraphic samples the data were analysed using detrended correspondence analysis (DECORANA; Hill

1979). The same fourteen pollen taxa as used for the zoning of the main pollen diagram were used in the analysis.

The results are shown in Figure 9. The plot shows that the analysis separates the four pollen zones identified from

Profile C and that the pollen spectra from the bone scrapes are more similar to those from paz NF1 (Unit C1) than

any of the other zones. This supports the working hypothesis generated in the field, and supported by the

radiocarbon dating (Scourse et al. 2009), that most of the bones came from Unit C1. However one of the jaw bones

might derive from Unit C3 (Figure 9).

The elements which characterize and unify the pollen assemblages from Unit C1 include high Cyperaceae and

Gramineae with subdominant Pinus, Artemisia and Trifolium cf. T. repens. Indeterminate, pre-Pleistocene and

dinoflagellate levels are high, as are frequencies of Pediastrum; P. duplex is either entirely absent or present in

extremely low concentrations. Overall, pollen concentrations are low, and in some cases so low as to make the

samples effectively barren.

Comparison of the changes in the assemblages from NF1, Profile C (Figure 3) with the changes within Unit C1

from Profiles Z and Y (Figure 6) shows some common trends. Gramineae rises from the base to reach maximum

levels towards the top, whereas Cyperaceae shows a reciprocal pattern of decline upwards. Moreover, this is

matched by an increase in the total Pediastrum concentrations upwards. These detailed changes enable the bone

scrape samples to be more accurately placed within Unit C1. The juvenile vertebra sample (Figure 6) is rich in


DOI: 10.1002/gj

Figure

8.Pollen

percentagediagramforProfileB.Taxathatneverexceed

1%areincluded

asothertree/shrub/herbaceous.Taxabetween1and2.5%areshownas

solidcircles.Pollen

sum

(SP)TLP(TotalLandPollen);CCryptogam

s;AAquatics;Iindeterminables;PedPediastrum.

Copyright # 2009 John Wiley & Sons, Ltd. Geol. 44: 414446 (2009

DOI: 10.1002/g

426 j. r. m. allen ET AL.J. )

j

palaeoenvironmental context of condover mammoth site, uk 427Cyperaceae, relatively low in Gramineae with moderate Pediastrum concentrations, whereas the juvenile jaw

samples are less rich in Cyperaceae, but with more Gramineae and moderate to high levels of Pediastrum. This

suggests that the juvenile vertebra bone derives from a lower stratigraphic level within the sequence than either of

the jaw bones. This conclusion is supported by the correspondence analysis (Figure 9), one of the jaw bones

probably deriving from Unit C3.

The correspondence analysis demonstrates similarity of pollen assemblages in zones NF3 and NF1. Zone NF3

can be clearly separated, however, on the basis of the presence of Betula (tree and B. nana), increased significance

Figure 9. DECORANA Factor 1 and Factor 2. Samples are 124 the samples from Profile C ((117 are from the upper section (sample 1 fromUnit F, 28 fromUnit E and 917 fromUnit D) and 1824 from the lower section Unit C1), sample 25 is from the juvenile jaw scrape, samples 26

and 27 from vertebra scrapes, 28 is from Unit C3, samples 2933 are from Unit C1 samples in Profiles Y and Z.of Salix (including S. herbacea) and Artemisia, Ranunculus trichophyllus type and Rumex acetosa type.

3.2. Plant macrofossils

Nomenclature and taxonomic order for vascular plants follow Tutin et al. (19641990) and those for mosses follow

Smith (1978). Plant macrofossil abundance scores are shown in Supplementary Tables S4 and S5. Together with

identifiable plant remains, a few other components of the samples were noted, and these are also presented in the tables.

The assemblages from samples E, F, G and 58 (Supplementary Table S4) have some taxa in common, but they

cannot be compared too closely since sample size was so variable (samples E and 58 were large; samples F and G

were small). Sample F is noticeably richer than the others, and contains abundant remains of birch (Betula),

suggesting it has more in common with Unit D than C1. However, the possible origin of at least some of the bones

from Unit C3 (green detritus mud) on the basis of pollen spectra (see above), suggests that this particular sample

may have consisted of Unit C3 rather than C1. This might explain its somewhat richer assemblage, as C3 was much

more organic than C1; the stratigraphic position of C3 and its associated AMS 14C date (Scourse et al. 2009) would

be consistent with the start of the Betula rise (Figure 3). The sample from the top of C1 prepared for AMS14C dating (Scourse et al. 2009) was, however, also rich in Betula macrofossils.

There is greater consistency between one sample and the next in the series from Profile Z (Face 2)

(Supplementary Table S5), with a gradual decline in the overall diversity of taxa upwards through this sequence,

corresponding to an increase in the sand content and a decrease in the organic component of the samples


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represented by samples B8B1. Between the series from Profile Z, and the sample from Profile C and the bone

scrapes, there are some striking differences; the samples from Profile Z are much less rich in taxa, lacking most of

the aquatics (but the one pondweed, Potamogeton rutilus, found at Profile Z is not recorded from Profile C).

Hyd

vegetation (Balfour-Brown 1950, p.235). Colymbetes dolabratus is adapted to very cold water and is now totally

428 j. r. m. allen ET AL.Copyright # 2009 John Wiley & Sons, Ltd. Geol. J. 44: 414446 (2009)

DOI: 10.1002/gjgriseostriatus inhabits pools of clear water over a sandy or stony bottom, not extensively overgrown by vegetation.

It is a species of cold water, and in Britain it tends to be found in peaty pools at high altitude: in northern European

latitudes it is found at lower elevations. Potamonectes depressus is both a flowing and still-water species. At

Condover, it was represented by a very pale, largely yellow form that is often found in ponds with a clear sandy

bottom. Agabus bipustulatus is found in pools and puddles and is not a stream species, except where slowly moving

or almost stationary habitats occur. Agabus arcticus is a cold-water species. Agabus congener is here a

representative of a species complex of largely northern forms that includes A. lapponicus, A. thomsoni and others

from the eastern palaearctic region. This is a species group that prefers cold-water ponds that are not overgrown by

aquatic plants. Rhantus notatus shows a strong preference for ponds with a silty bottom with some aquaticHalip

dytisple E. Most of the other water beetles live in slowly moving or stationary water. Both larvae and adults of

lus are relatively poor swimmers and so live only in standing or very slowly moving water. In contrast, the

cids are powerful swimming carnivores that can live in both flowing and standing water. PotamonectesTh

samrmation about the aquatic environment. These include Haliplidae, Dytiscidae, Gyrinidae, Hydraenidae,

rophilidae, Dryopidae and some of the Chrysomelidae and Curculonidae.

ree species of Dryopidae are characteristic of rapidly flowing, well-aerated water. They occurred only in3.3.1

info.1. Local environments indicated by the Coleopteran assemblage. Several families of Coleoptera provideprefeplementary Table S3). A list of the taxa recorded is given in Supplementary Table S6 and the environmental

rences of these taxa are outlined below.3.3. Coleoptera

Coleoptera lists are shown in Supplementary Table S6 (samples AG) and Supplementary Table S7 (samples from

Profile Z). The nomenclature and taxonomic order follows Lucht (1987), with species that do not occur today in

central Europe being inserted at the most suitable taxonomic level. The numbers indicate the minimum number of

individuals recovered from each sample; 134 taxa of Coleoptera were identified, of which 94 have been determined

to the level of species or species group. Eleven of the latter are now extinct in the British Isles. These exotic species

are indicated by in the Tables.The sequence of samples shown in Supplementary Table S6 is subdivided into four sections.

(1) Samples A, B, D and E represent the dark grey clayey sandy silt (Unit C1), either as the direct matrix of the

bones or associated with them on the spoil heap.

(2) Sample C was collected in situ from the floor of the pit and well separated from the bone-bearing locality, but

included almost all the species found in the rest of this sub-group including many of the exotica. Sample C is

correlated with Unit C1.

(3) Sample F was obtained from the sinus cavities and lower jaw dental alveoli and is clearly different from the rest

of the samples.

(4) Sample G was taken from the green detritus mud (Unit C3) that overlay the dark grey clayey sandy silt (Unit

C1) and contains fauna so different from that of Unit C1 that it is here considered on its own.

Supplementary Table S7 shows the beetle fauna from the sequence of samples from Profile Z Face 2. It is dealt

with separately because of its stratigraphical integrity and because this series of samples shows progressive change

in the local environment.

3.3.1. Samples A, B, D and E

These four samples were of the dark grey clayey sandy silt (Unit C1) adhering to the mammoth bones

(Sup

absent from the British Isles; its nearest occurrences today are in the alpine and subalpine zones of the Norwegian

mountains where it lives in small ponds and peaty pools. Dytiscus circumcinctus and Acilius, in this case probably

A. sulcatus, are both pond species and, being rather large beetles, they tend to inhabit rather larger bodies of water

palaeoenvironmental context of condover mammoth site, uk 429than most of the other dytiscid species in this fossil assemblage.

Gyrinus aeratus andG. marinus are difficult to separate and it is possible that both species occurred at Condover.

The hydraenid species are almost all pond or puddle species. Most of the species here occur in grassy ponds with the

exception ofHelophorus obscurellus which is one of the least aquatic species of its genus. Although little is known

about its precise ecological requirements, it is normally found under stones and vegetation in sandy places in the

tundra of northern Asia and on the high cold steppes of the mountains further south as far as the Tibetan plateau

(Angus 1992; p.29). By reference to its nearest relatives (e.g. H. nubilus), it is highly likely that its larvae are

phytophagous, feeding on grass shoots. H. sibiricus is a northern species found today from Siberia to Alaska, in

northern Canada and Fennoscandia, and in mountains further south in Scandinavia. It is found in grassy pools and at

the edges of streams and rivers (Angus 1992; p.41). Helophorus splendidus is one of the most cold-adapted species

of this genus. It is known from the tundra across arctic Siberia to the north coast of Canada.

Phytophagous aquatic Coleoptera provide information on the plant species present. Macroplea appendiculata

lives under water on various species of Potamogeton (pondweeds), Myriophyllium (milfoils), aquatic species of

Ranunculus and Nymphaea (water lilies). Donacia versicolorea feeds exclusively on Potamogeton natans (Koch

1992; p.52). The aquatic weevil Bagous lives on submerged water plants and its larvae feed on their stems and

leaves. It is a very poor swimmer and is thus confined to standing waters. Hydrobius fuscipes is a detritus pond

species tolerant of some brackishness in the water.

Hints of brackish habitats are provided by Berosus spinosuswhich is strongly halophilous being found in shallow

pools on salt marshes. Other species that are at least salt tolerant includeOchthebius lenensis and Cercyon marinus.

Bembidion aeneum is by far the most abundant species of carabid beetle in this assemblage. It is a very common

member of fully glacial and Late-glacial faunas in Europe and also tolerates brackish conditions (Lindroth 1985, p.173).

3.3.1.2. Terrestrial habitats indicated by the Coleoptera. Many species of the Hydrophilidae are not water

beetles; many species of Cercyon live in rotting plant detritus and sometimes in dung. Coelostoma orbiculare

occurs in marshy places where the margins of a pond are choked with dead or dying vegetation.

The carnivorous or scavenging ground beetles, the Carabidae, are represented by twenty species. Notiophilus

aquaticus is a xerophilous species that lives in dry open ground often on what would appear to be completely sterile

gravel and in montane environments it is associated with ericaceous heath (Lindroth 1985, p.78). Blethisa

multipunctata is a hygrophilous species often associated with Agonum versutum (Lindroth 1985) living in marshy

fens with Carex and Eriophorum but not where Sphagnum is predominant in the vegetation. The species have a

preference for sun-exposed ground. Elaphrus cupreus is a hygrophilous species that prefers eutrophic fens.

Elaphrus riparius prefers sparsely vegetated sun-exposed habitats on both clay and sandy soil.Dyschirius tristis is a

subterranean species that excavates burrows in firm clay soil beside water where the soil is bare in places otherwise

densely vegetated withCarex. This species prefers, but is not confined to, saline habitats. Trechus secalis is found in

both open country and woodland, under plant debris in meadows on clay-rich soils. Seven species of Bembidion

occur in this assemblage; these are highly active scavengers or predators living in damp habitats where they are

often dependent on particular types of soil, except Bembidion nigricorne which is xerophilous and lives on dry,

sandy, Calluna heaths in sparsely vegetated places. Bembidion bipunctatum occurs near water on a variety of soils

from stony gravel to silty sites with Carex or grasses. Bembidion obliquum occurs on soft moist ground with a

growth of Carex, grasses and Equisetum on clayish or peaty soils with sun-exposed patches. Bembidion

quadrimaculatum is a heliophilous species, living on clay-rich or sand clay soils with thin vegetation on the drier

banks of fresh water. Bembidion doris is a very hygrophilous species in wet highly vegetated places characterized

by, for example, Carex and Eriophorum. Poecilus cupreus inhabits moist meadow-like habitats with a dense

vegetation of sedges and grasses, and preferably clay soil. Pterostichus minor is found in wet habitats particularly

beside eutrophic ponds. Pterostichus adstrictus lives in open grassy places especially on gravelly soil that has a

mixture of clay and humus. Calathus melanocephalus is a very common species in open country, in dry grassy


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places, especially on sandy soils. Agonum versutum is also a species of open country, often where there is a

luxuriant growth of Carex and where the substrate is soft and muddy. Amara alpina is one of the most characteristic

carabid species of montane and tundra environments, ranging from shrub tundra to almost sterile ground with only

430 j. r. m. allen ET AL.isolated patches of vegetation.

Species of Staphylinidae are mostly carnivores, preying on soil arthropods and worms. Many of the species in

this fossil assemblage live under plant debris in damp conditions, including Eucnecosum brachypterum, Acidota

crenata, Lesteva longelytrata, Geodromicus nigrita, Oxytelus rugosus, O. nitidulus and O. laqueatus. Platystethus

cornutus, the most abundant species here, is fossorial, digging tunnels in sandy banks. Bledius is not a carnivore but

feeds on algae in the superficial layers of sandy and silty soils Tachinus fimetarius prefers open, rather dry sandy

places where it lives under vegetable debris or dung.

Most species of byrrhids are strictly moss-feeders, both as adults and as larvae. Byrrhus, Cytilus,Curimopsis and

Simplocaria form a group that live in microhabitats with loose, well-drained granular soils which are often skeletal

and usually include a high proportion of sand, gravel, cobbles or broken rock. Typically the mosses Ceratodon

purpureus and Polytrichum juniperinum dominate these habitats in North America which also often contain many

weedy and pioneer plant species (Johnson 1987, p.126).

Other plant-eating beetles that feed exclusively on macrophyte vegetation belong to the Chrysomelidae and

Curculionidae. One of the most abundant is the weevil Sitona sp. whose larvae live exclusively on the roots of

Papilionaceae. The relatively high numbers of this weevil indicate pioneer vegetation invading an area of raw soils

with a poorly developed nutritional status. Several other species of beetle indicate the preponderance of weedy

species in the neighbourhood. Galeruca tanaceti lives on a variety of Compositae, but above all on Tanacetum

vulgare (tansy) and Achillea millefolium (yarrow). Adoxius obscurus feeds on Epilobium (willow herbs), its larvae

feeding on the roots of the same plant.Miarus campanulae is confined to Campanulaceae in rather dry places where

its larvae develop in galls on the seed heads. Hypera elongata is chiefly found onMyosoton, in particular the water

chickweed M. aquaticum, but it is also known on species of Stellaria (chickweeds and stitchworts) and on

Cerastium arvense (field mouse-ear). Otiorhynchus ovatus is a xerophilous weevil that feeds on a wide variety of

herbaceous plants. Barynotus squamosus is also polyphagous on low plants but in rather damper habitats. The only

evidence of shrubs is provided by three species,Melasoma collaris, Phyllodecta and Rhynchaenus folorium which

feed primarily on Salix. Of these only the latter species is at all common and it is an extremely small weevil that

likely fed on one of the dwarf willows. Chaetocnema hortensis feeds on grasses and is somewhat salt tolerant.

Species of Coccinellidae are predatory on aphids. Thus, although no aphids were recovered from the sediment,

no doubt because their frail cuticle failed to be preserved, it can be inferred with certainty that many plants were

infested with aphids. Hippodamia arctica is a species of the far north of Europe and is currently extinct in the

British Isles. It is found on small Salix bushes, Betula nana and Arctostaphylos alpinus (Strand 1946).

Specialist dung beetles are important in the context of these mammoth remains since they would be expected to

occur in great profusion had the mammoths been present at the time of sedimentation (e.g. Coope et al. 1961 and

Bosemier et al. 2003). Two species were present, Aphodius prodromus (Brahm.) and A. distinctus (Mull.), mostly

the former species but, since they are not always possible to separate from one another they are included in the

Tables as Aphodius spp. Both these species are obligate coprophages rarely found in rotting vegetable matter.

However, only in sample E is Aphodius prodromus at all common.

3.3.2. Sample C

A list of the Coleoptera recovered from sample C is given in Supplementary Table S6. The sample was obtained

from sediment in situ from sedimentary Unit C1. The fauna was almost identical to the assemblages from samples

A, B, D and E and thus suggests that Unit C1 was the mammoth stratum, as suggested also by the palynology and

the radiocarbon data (Scourse et al. 2009).

3.3.3. Samples from Profile ZFace 2; Unit C1

At this locality the dark grey clayey sandy silt (Unit C1) was arched into a sharp anticline (Scourse et al. 2009).

Detailed sampling of this stratum was undertaken because the sediment was similar to the matrix of the mammoth


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bones and it was one of the few opportunities to examine the stratigraphy in situ. The assemblage of Coleoptera

from the sequence of samples taken through this exposure of Unit C1 is shown in Supplementary Table S7 with

sample 010 cm taken from the top of the stratum and 9095 cm from the base. This suite of Coleopteran species

palaeoenvironmental context of condover mammoth site, uk 431leaves no doubt that this outcrop of Unit C1 correlates with those samples from Unit C1 elsewhere and from the

sediment adhering to the mammoth bones, though no mammoth bones were found at this locality. Unfortunately,

the Coleopteran assemblages provided no evidence of any environmental changes though, again, the fauna includes

both cold-adapted species as well as relatively temperate ones.

Out of a total of 35 species recorded here, only six were not found in samples AE. Furthermore, the species that

are common to both assemblages include the exotic forms that are now absent from the British Isles. These include

Helphorus splendidus, whose occurrence in the British Late-glacial appears to be restricted to the period of cold

climate just prior to 15 440 cal. year BP (13 000 14C years BP) (e.g. Coope and Brophy 1972) i.e. to Greenland

Stadial 2 (Walker et al. 1999; Lowe et al. 2008).

Clivina fossor is a subterranean species that burrows in rather damp clay-rich soil in open country. Bembidion

virens is characteristic of sterile, gravelly or stony stream banks or similar lake margins and is always found near

water in company with Bembidion bipunctatum (Lindroth 1985, p.182). Bembidion quadripustulatum is a relatively

southern species compared with most other species in this assemblage, in Britain only reaching as far north as

Derby and in Scandinavia only occurring as accidental individuals on the south coast of Scania. It is found on damp

sun-exposed clay or sandy mud. Bembidion minimum is a halophilic species that lives on salt marshes on moist clay

where the vegetation is sparse and leaves bare patches where the beetle is often found in direct sunlight.

3.3.4. Sample F

This beetle assemblage, obtained from the mandibular canal in the lower jaw and from the sinus cavities in the

juvenile skull when these were being prepared in the laboratory, is very different from those described above

(Supplementary Table S6). The total weight of sediment involved was less than 1 kg yet the amount of insect

material recovered was prodigious. By far the largest quantity of insect fragments were the broken pupal cases of

the fly Phormia terraenovae, the larvae of which must have infested the body of the young mammoth (see

Erzinclioglu, in Lister 2009). Almost all the beetle fragments were of Aphodius prodromus, indicating that dung

was abundantly available at this time and, by the way the highly organic material had been forced into the jaw

cavities, it seems likely that the skull bones could have been trampled into the layer of dung. A smaller amount of

similar material was also recovered from the sinus cavities of the juvenile skull fragment. In this context it is notable

that the fly puparia were concentrated at the back of these cavities whilst the sediment rich in Aphodius remains was

more superficial, suggesting that the fly puparia were in place first before the dung beetles were forced into the

cavities. The large number of individuals of Aphodius (n 128) from this small sample is undoubtedly anunderestimation of their abundance (heads were the only skeletal element counted). This sample demonstrates just

how common the dung beetles must have been at the time when the mammoths were present at the Condover site

and contrasts markedly with their poor representation in the samples of dark grey clayey sandy silt matrix encasing

the majority of the mammoth bones.

Necrobia violacea is a corpse beetle that feeds on the larvae of flies and other insects in dried-out carcasses that

have been reduced to skin and bone. The occurrence of large numbers of Aphodius and the presence of Necrobia

shows that at this time, at least, both the dung and the bodies of juvenile mammoth must have lain about on the

surface for sufficient time for the carcass to be largely consumed by blow fly maggots and subsequently to have

become desiccated. This may account for the fact that some of the bones of the juvenile skull appeared to be rather

more decomposed than the rest of the skeleton, suggesting that they had been subjected to weathering prior to their

incorporation into the sediment.

The assemblage from sample E bears some similarities to that from sample F. Aphodius is moderately abundant

in sample E in marked contrast to samples A, B, C and D, also from the dark grey clayey sandy silt of the mammoth

bone matrix, and from Unit C1. Necrobia violacea was also present in both samples E and F, suggesting that these

comprised an admixture of the dark grey clayey sandy silt with some of the dung-rich material from the surface


DOI: 10.1002/gj

represented by sample F. Some evidence of this physical mixing of the grey silty clay with more organic material

was visible in the field, in the form of scattered clasts of darker mud scattered through the grey matrix.

432 j. r. m. allen ET AL.3.3.5. Sample G

There are a number of important differences between the assemblage from samples from sedimentary Unit C1

(samples AE; Supplementary Table S6) and Sample G taken from the green detritus mud (Unit C3) directly

overlying it. There is a total absence in sample G of the suite of cold-adapted species that are a distinctive

component of the beetle assemblages in the dark grey clayey sandy silt of Unit C1. These absentees include Amara

alpina, Potamonectis griseostriatus, Agabus arcticus, Ochthebius lenensis, Helophorus obscurellus, Helophorus

sibiricus, Helophorus splendidus, Eucnecosum brachypterum, Simplocaria metallica, Curimopsis cyclolepidia and

Hippodamia arctica. The absence of some of these species could be attributed to sampling inadequacy, but the total

lack of them all is not so easily dismissed, especially since some of them were relatively abundant in samples AE.

In their place the fauna from sample G includes a number of relatively southern species such as Agonum

sexpunctatum, Coelambus impressopunctatus, Hygrotus inaequalis, Potamonectis assimilis, Plateumaris affinis

and Notaris scirpi, none of which are obligate northern species. A significant feature of this sample that

distinguishes it from the other samples is the large numbers of bones and spines of Gasterosteus aculeatus, the

three-spined stickleback, that were recovered from it as a by-product of the extraction of the insect fossils. No fish

remains of any sort were found in samples AE, in spite of the use of exactly the same extraction procedure. These

differences between the faunas of Unit C1 and Unit C3 indicate that the local environment and the regional climate

were also very different and would suggest that the two units are not just lateral variations of one another.

The occurrence in this sample of moderate numbers of elmid beetles indicates that here, as in sample E, there

must have been running water available. Potamonectes assimilis lives in streams or pools with a sandy bottom

where there is little vegetation (Nilsson and Holmen 1995). In contrast, Coelambus impressopunctatus and

Hygrotus inaequalis are species of well-vegetated ponds or very slowly moving water. The relatively large number

of individuals of Gyrinus indicates that the water surface was not completely covered with vegetation.

Bembidion nigricorne is a species of dry sand heathland, or moderately humid peaty soils where it occurs in

sparsely vegetated places. Agonum sexpunctatum is a strongly heliophilous species of open country on moist soils

that are exposed to the sunlight and sparsely covered with mosses or grasses.

3.3.5.1. Climatic interpretation of the beetle assemblages. In a fauna that is dominated by cold-adapted species,

the Coleopteran assemblages from Unit C1 and samples AE contain an admixed element of temperate forms that

are incompatible in terms of their present day geographical ranges; some high arctic species occur in the same

samples as southern English ones. The cold element in the beetle assemblage includes Bembidion virens,

Helophorus obscurellus, Amara alpina, Helophorus sibiricus, Potamonectes griseostriatus, Helophorus

splendidus, Agabus arcticus, Eucneiosum brachypterum/norvegicum, Agabus congener, Limplocaria metallica,

Colymbetes dolabratus, Curimopsis cyclolepidia, Ochthebies lenensis and Hippodamia arctica. The warm

element includes Bembidion minimum, Hygrotus inaequalis, Bembidion quadripustulatum, Chaetarthria

seminulum, Bembidion nigricorne, Berosus spinosus, Pterostichus minor and Oulimnius tuberculatus.

This mixture cannot be explained by invoking extreme variation in local microclimates, especially since the

topography of the Condover district is so flat that there is very little altitudinal contrast within the catchment area

from which the fossil insect assemblage was drawn. Theoretically no-analogue assemblages (i.e. assemblages of

species which do not live together today) are possible, because of lag effects at a time of rapidly changing climate

and the possibility that modern ranges are partly determined by competitive exclusion and not just individual

species physiological boundaries. However, this is thought an unlikely explanation for the mixed assemblage

because the known pattern of Late-glacial faunal change from other British sites indicates clear transitions from

cold to warm associations and not disharmonious faunas. For instance, at Glanllynnau (Coope and Brophy

1972), the northern and southern species were sequentially separated and clearly came from periods of quite

different climatic conditions. Thus, the mixture at Condover cannot be ascribed to a no-analogue situation. It is


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suggested that this mixture of species comprises animals that lived at different times in contrasting climates,

physically brought together into the same sediment.

As the beetle assemblage from the green detritus mud of sample G (Unit C3) showed no evidence of any mixing,

it was possible to apply the Mutual Climatic Range (MCR) technique (Atkinson et al. 1987) to derive atmospheric

palaeotemperatures. The mean July temperature is reconstructed to be 15198C and the mean January temperatureto13 to68C; this implies that the mean monthly temperatures lies somewhere between these values and not thatthe temperatures varied between these limits.

3.4. Other environmental proxies

3.4.1. Anuran remains

(J. Alan Holman (deceased), Michigan State University Museum, East Lansing, Michigan, 48825-1045 USA.

Personal communication 1990)

Late-glacial amphibians have been very rarely reported from Britain (Stuart 1982) and have rarely been

specifically identified. Thus, the recovery of anuran (frogs and toads) bones that may be assigned to species from

the Condover mammoth site is of interest. The anuran bones were found in the green detritus mud (Unit C3) where

they were in association with fish remains, mainly Gasterosteids (sticklebacks). This anuran-bearing stratum was

directly above Unit C1, the hypothesized source of the mammoth remains.

The finds were a left femur (Figure 10A), a metatarsal (Figure 10B), and an omosternum (Figure 10D). The frog

bones are well preserved and mainly black in colour, with a very slight reddish tint. Measurements of the remains

are given in Supplementary Table S8.

All of the anuran bones have been assigned to the living species Rana temporaria Linnaeus, 1758, which occurs

today throughout Europe east to the Urals, but excluding most of Iberia, much of Italy, and the southern Balkans

palaeoenvironmental context of condover mammoth site, uk 433(Arnold and Burton 1978; Frost 1985).

The femur: Fossil anuran femora have rarely been identified to either the generic or the specific level. This is

probably because it is difficult to distinguish left and right fossil specimens, and thus difficult to compare specific

structures on the femora of fossil and modern skeletons. Left and right ranid femora are usually distinguished on the

basis of a groove that occurs on the lateral side of the posteroventral surface of the bone (Figure 10A). At the generic

level the Condover femur may be identified as Rana as follows. It is distinguished from Bufo (Bufonidae) by having

Figure 10. (A) left femur (ventral view) of Rana temporaria from the Condover mammoth site; (B) posterior two-thirds of right femur (ventralview) of modern Rana esculenta; (C) metatarsal of Rana temporaria from the Condover mammoth site; (D) omosternum of Rana temporaria

from the Condover mammoth site. All of the lines represent 1mm.


DOI: 10.1002/gj

a more gracile femur and by lacking a posteroventral crest. It is distinguished from Alytes, Bombina, and

Discoglossus (Discoglossidae) by lacking a distinctly S-shaped shaft, having less swollen anterior and posterior

ends, and lacking a posteroventral crest. From Hyla (Hylidae) it is distinguished by having a less gracile femur and

434 j. r. m. allen ET AL.in lacking a posteroventral crest. Finally, it is distinguished from Pelobates and Pelodytes (Pelobatidae) by having a

more gracile and much less strongly bowed femur.

At the specific level the Condover femur may be assigned to Rana temporaria on the basis of the deep lateral

groove that occurs on the posteroventral surface of the femur. This portion of the femur is smoother and more

rounded in Rana esculenta (Figure 10B) and Rana ridibunda. Moreover, femora of other European Rana appear to

be generally less gracile than those of Rana temporaria, especially British specimens.

The metatarsal: The fossil metatarsal (Figure 10C) represents a larger frog than the one represented by the femur,

thus at least two individual frogs are indicated by the fossil material. Although the bone appears to be indistinguishable

from modern Rana temporaria, it was not possible to tell whether it was from the left or the right foot.

The omosternum: The presence of a bony rather than a cartilaginous omosternum is an important diagnostic

character of the family Ranidae and the genus Rana (Duellman and Trueb 1986, p. 542). The family Bufonidae

usually lacks an evidence of an omosternum (as in Bufo bufo), and when it is present, it is cartilaginous and

normally would not fossilize. The other European anuran families (Discoglossidae, Hylidae, Pelobatidae) have

cartilaginous omosterna. The bony omosternum of Rana is normally subtriangular (Figure 10D) and has a wide

posterior articular portion and a constricted anterior portion. The fossil bone was indistinguishable from those of

modernRana temporaria. The omosternum also represents amuch larger specimen than the one represented by the femur.

Rana temporaria has previously been reported from the Middle Devensian site at Upton Warren, Worcestershire

(Coope et al. 1961). At this site, silt bands within the sands and gravels yielded pollen spectra that indicated

treeless, herb-dominated vegetation. One of the silt bands yielded a radiocarbon date of 42 100 800 14C year BP.As at Condover, the Rana temporaria bones at Upton Warren were associated with those of the three-spined

stickleback (Gasterosteus aculeatus). The only other British Devensian herpetological records are Rana sp. and/or

Bufo sp. from the early Devensian site at Coston near Barnham Broom, Norfolk (Allison et al. 1952; Stuart 1982).

It seems noteworthy that the two Devensian herpetological species that have thus far been identified from Britain,

Rana temporaria and Lacerta vivipara, are forms that occur well within the Arctic Circle in Europe today (Arnold

andBurton 1978;maps 36 and 70).Moreover, of the 126 herpetological species reported in Europe byArnold andBurton,

Rana temporaria and Lacerta vivipara are by far the most widely distributed herpetological species in far northern areas.

Today, Rana temporaria is very widespread in Britain and may be found in a variety of damp situations where

adequate herbage exists. Nevertheless, it is most often found near ponds (Smith 1964), and may stay in the water for

a month or more after the breeding season (Frazer 1983). It would seem likely that the fossil frogs lived in or near

the basin in which the mammoths became entrapped. The fact that the delicate anuran bones are well preserved

indicates that there was little or no post-mortem transport.

3.4.2. Trichoptera and Chironomidae

(N. E. Williams (retired), then Division of Life Sciences, Scarborough Campus, University of Toronto, 1265

Military Trail, Scarborough, Ontario. M1C 1A4 Canada. Personal communication, December 1991)

Two samples, one from Unit C1 (dark grey clayey sandy silt) and the other from Unit C3 (green detritus mud)

were examined. Trichopteran (caddisfly) and chironomid sclerites were extracted from the sediments, using the

same techniques as for the Coleoptera, and were mounted on glass slides in Berlese fluid. Totals of six caddisfly, and

twelve chironomid taxa were identified (Supplementary Table S9).

All of the aquatic insect taxa identified from Unit C1 are found in Britain today. There were no aquatic taxa

restricted to arctic, subarctic or boreal regions. Limnephilus stigma/nigriceps, L. vittatus and Agrypnia obsoleta are

all now widely distributed in northern and central Europe and European Russia, although L. vittatus does not reach

the northernmost parts of Scandinavia (Botosaneanu and Malicky 1978). The sand-cased L. vittalus is associated

with sandy and silty substrates of lakes, ponds and temporary pools, while both L. stigma and L. nigriceps occur

among dense emergent marginal vegetation in still and running waters, using plant materials for food and case

construction (Wallace et al. 1991). The predaceous Agrypnia obsoleta occupies all types of freshwaters but requires


DOI: 10.1002/gj

plant materials for case construction. In Britain it occurs mainly in upland areas of Scotland, northern England and

Wales, but it is more widespread in Ireland.

Two additional caddisfly species were present in the green detritus mud (Unit C3) sample. Both Lepidostoma

palaeoenvironmental context of condover mammoth site, uk 435hirtum andOecetis ochracea are now common and widespread throughout Britain, central and northern Europe and

European Russia (Botosaneanu and Malicky 1978).

The chironomids provide evidence for conditions at the time of deposition of Unit C3. Chironomini, excluding

Chironomus, Sergentia and Stictochironomus, are included by Walker et al. (1991) amongst the primarily

temperate, littoral taxa with distributions that do not extend beyond the tree line. They state that although most of

these thermophilous Chironomidae never individually compose a large proportion of the chironomid fauna of a

lake, their overall relative abundance declines rapidly with increasing latitude. In their study of Labrador lakes, all

sites where temperate littoral chironomids represented more than 10% of the total chironomid specimens counted

were south of 558N and mostly in the boreal spruce-fir forest vegetation zone; in tundra lakes this group comprised05% of chironomid specimens. In the Unit C3 sample head capsules of members of this group associated with both

sediments and littoral plants were common. Of the remaining taxa, the detritivorous Chironomus sp. was abundant.

This genus has a worldwide distribution, but tends to be extremely abundant in very productive temperate lakes.

The considerable change in the chironomid assemblage between Unit C1 and Unit C3 would seem to indicate

warmer conditions during deposition of the upper sediments or a lag in arrival of thermophilous species caused by

very rapid warming. The latter is a definite possibility from the point of view of aquatic species since there were no

obligate northerners present in either sample.

3.4.3. Ostracoda

(Eric Robinson (retired), Department of Earth Sciences, University College London, now at Riverside Farm,

Whitehall, Watchet, Somerset, TA23 0BB, Personal communication, February 1988)

Investigations using standard techniques (e.g. Chapter 5 in Griffiths and Holmes 2000) were made of samples of

sediments from Unit C1 Profile Z (1988). These revealed a restricted species diversity of five species, of which one,

Limnocythere blankenbergensis (Diebel 1968), dominated, forming a species life assemblage comprising 85% of

the total assemblage. Originally described from Late Weichselian lake sediments in North Germany (Diebel

1968), it has not been found alive and this is its first record from Britain. It forms a dominant component of the

Late-glacial (Older Dryas) ostracod assemblage at Duvensee, North Germany (Gunther 1986) where, along

with Limnocythere inopinata (Baird, 1843) and Candona candida (Mueller, 1776), two species also found at

Condover, it has been interpreted as a characteristic shallow limnic, pioneer species capable of tolerating low

temperatures. Candona candida and Ilyocypris bradyi Sars, another of the Condover assemblage, are also

stenothermal, cold-water species (Klie 1938) often characteristic of pioneer assemblages of the Late-glacial

(Absolon 1973). The structure and context of the Condover ostracod assemblage from Unit C1 is therefore entirely

consistent with the age and environmental context of similar assemblages from the Weichselian Late-glacial of

North Germany.

3.4.4. Mineral magnetic measurements and the occurrence of greigite

(J. P. Smith (retired), Division of Environmental Science, School of Applied Sciences, University of

Wolverhampton. Personal communication, 1992)

Because of the disturbed nature of the Condover site, it seemed likely that simple, rapid and non-destructive

mineral magnetic measurements, that can be made on small samples of sediment, might be an aid to reconstructing

the general stratigraphy of the site, in providing a stratigraphic context for the excavated mammoth bones (cf.

Walden et al. 1992), and for gaining insights into the provenance of the basin infill.

Magnetic minerals can be both from primary (minerals formed in a rock at its time of formation) or secondary

(formed at the time or after deposition of the sediment by processes such as pedogenesis, fire or diagenesis/

authigenesis) (Oldfield and Thompson 1986). In sedimentary sequences such as that at Condover it might be

expected that the magnetic mineral assemblages would be dominated by the clastic detrital input to the basin.

Although the magnetic signature of sediment responds to the mixture of magnetic grain species, their size, shape


DOI: 10.1002/gj

and stoichiometry, in this environmental setting the likely contained magnetic minerals would be various

magnetites and haematites.

The values obtained for magnetic parameters can be used to characterize the contained magnetic mineral

436 j. r. m. allen ET AL.assemblage. Supplementary Table S10 gives the data for three magnetic parameters for the sediment samples.

Magnetic susceptibility (x) is a measure of the concentration of ferrimagnetic minerals present, while the other two

parameters (IRM100mT / IRMSat and IRMSat/x) represent the magnetic mineral mixture and magnetic grain sizes

present. The samemeasurements for sediment samples scraped from themammoth bones and for sediment believed to

be from the same stratum but not in immediate contact with the bone are also shown in Supplementary Table S10.

The parameter values for samples taken from the exposed faces are typical of Triassic substrates and Triassic

derived drift (Smith et al. 1990) and reflect differences in the mixture of detrital magnetites and haematites in the

deposits. However, the sediment in contact with the bone has a magnetic susceptibility value an order greater that of

the sediments in the basin and an IRMSat/x ratio much higher than that most other sediments. The only similarity is

with the sediment believed to come from the same stratum.

3.4.4.1. Magnetic extracts and XRD. The magnetic characteristics of the sediment attached to the bone

suggested a high concentration of a multidomain ferrimagnet, which was initially thought to be magnetite. A

magnetic extract was made from the sediment and the material analysed by XRD. The magnetic mineral was

identified as greigite, a ferrimagnetic spinel with the chemical formula Fe3S4.

Greigite is a metastable iron sulphide formed in aquatic environments normally as a member of a series between

amorphous FeS and pyrite. It is common in marine sediments but has also been recognized in freshwater systems

(Hilton 1990). The mineral is not stable in aerobic conditions and was not therefore part of the clastic input to the

deposit, but must have formed authigenically, as a result of the particular geochemical environment existing in the

deposit at the time of deposition or, diagenetically, soon after burial by overlying deposits. Adapting the findings of

Hilton (1990) and others, who haveworked on the occurrence of greigite in limnic systems, the requirements for the

formation of greigite appear to be a source of labile iron, reducing conditions and a source of organic sulphur.

Such a geochemical environment must also be one that is consistent with the preservation of the mammoth bones

and other organic remains. The excess of greigite adjacent to the bones presumably reflects the availability of

organic sulphur. Some of the unusual effects reported by other scientists working on the remains (e.g. magnetic

insect remains, oxidized coatings on insect remains after separation and drying, and the corrosion of pollen grains

during chemical treatment) all reflect the coating of organic surfaces by the authigenic greigite. The greigite clearly

is a manifestation of the extreme geochemical environment responsible for the high quality of the preservation of

organic remains at the site. Presumably, the very high organic loading drove geochemical reactions towards the kind

of reducing conditions necessary for preservation.

Detailed comparisons of the magnetic signature of the sediments from the same stratum but not in immediate

contact with the bone gave similar characteristics, but the lower magnetic susceptibility suggested a much lower

concentration of the mineral. It therefore seems likely that the greigite was contained in the stratum containing the

mammoth bones and was selectively concentrated in and around the bones themselves.

4. ENVIRONMENTAL INFERENCES

Inferences based upon the range of environmental indicators are presented for the sedimentary units investigated.

The ages of these units are derived from a series of radiocarbon dates, calibrated using OxCal v 4.0 and calibration

data set IntCal04, reported in full in Scourse et al. (2009). The Devensian Late-glacial and early Holocene

environmental history of the basin at Norton Farm, Condover, is summarized in Table 1.

4.1. Units C1 and C3: Before 13 82014 830 cal. year BP (12 240W 120 14C year BP)

These units, which were the source of the mammoth bones, were the most intensively studied with data available for

all the proxies investigated.


DOI: 10.1002/gj

Table

1.Summaryofenvironmentalhistory

Water

inBasin:

trophic

status

Aquatic/helophyticenvironment

within

basin

Dry

landvegetationoutsidebasin

After

1079010250cal.yearBP

(9350100

14Cyears

BP)

Noinform

ation

Successivecolonizationbyjuniper

(Juniperus),willow

(Salix),birch

(Betula),hazel

(Corylus),pine

(Pinus)

andalder

(Alnus)

ca.11500to


(ca.10000to

9350100

14CyearBP

HIATUSATPROFILEC

1286012380to

ca.11500cal.yearBP

(10640100to

ca.10000

14CyearBP)

Oligotrophic,nutrient

poor,highO2

Fringingcommunitiesofsedge

(Carex)

withhelophyticRanunculus.

Grass-dominated,butwith

Artem

isia,Rumex

andThalictrum

1370013250to


(11590110to

10640100

14CyearBP)

Moderatelyeutrophic,

calcareous

Richabundance

ofboth

submerged,

floating-leaved

andsubmergent

waterplants,includingwater-m

ilfoil

(Myriophyllum),marestail

(Hippurisvulgaris)

andpondweeds

(Potamogeton).Diverse/abundant

helophyticassemblage

Open

birch

woodland,withtree

anddwarfform

s,juniper,

poplar

probably

aspen

(Populus)

andwillow

1483013930to


(12300100to

11590110

14CyearBP)

HIATUSATPROFILEC

MAMMOTHSENTERBASIN

BIOTURBATIO

N

Before

andaround1483013930cal.yearBP

(12300110

14CyearBP)

Eutrophic,nutrientrich,

low

O2,calcareous

Characeae,

(calcareousalgae,

stonew

orts)

andpondweeds.Fringing

communitiesofsedge,

club-rush

(Scirpuslacustris)

andmarsh

cinquefoil

(Potentillapalustris).Thepresence

variousspeciesofPotamogeton,

Myriophyllium,Nym

phaea

andaquatic

speciesofRanunculusareallindicated

bytheaquatic

plant-eatingColeoptera.

TheColeoptera

also

indicateamosaic

ofslow

movingandstagnantwater

and

fastflowingwell-aeratedwater.Frogs

(Ranatemporaria)werepresentin

the

vicinityandsticklebacks(G

asterosteus

aculeatus)

werepresentin

thewater

body.Thechironomidsandcaddisflies

indicatetemperateconditions

Grass-dominated,butwith

weedtaxa

including

Chenopodiaceae,Rumex

andthrift

(Arm

eria

maritima).Somedwarf

willow.Notrees,butopen

birch

scrublandwithdwarfbirch

(Betula

nana)towards14400cal.

yearBP(12300

14Cyears

BP).

Agenerally

barren,sparsely

vegetated,landscapeisindicated

bytheColoepteranassemblage.

Theweevilsindicatepioneer

vegetation.Ladybirdswerepresent

indicatingthepresence

ofaphids,

anddungbeetles

indicatethe

presence

ofsomelarge

dung-producinganim

als

opyright # 2009 John Wiley & Sons, Ltd. Geol. J. 44: 414446 (2009)

DOI: 10.1002/gj

palaeoenvironmental context of condover mammoth site, uk 437C

The plant macrofossil data from Unit C1 clearly indicate the local presence of a number of taxa recorded in the

pollen spectra, including Salix sp(p)., Betula sp(p)., Chenopodiaceae, Potentilla sp., Myriophyllum verticillatum,

Armeria maritima, and species of Gramineae and Cyperaceae, including Juncus sp(p)., Scirpus lacustris,

438 j. r. m. allen ET AL.Eleocharis palustris and Carex spp. The decrease in abundance of plant macrofossils upwards through the series of

samples from Profile Z parallels the decline in pollen concentrations from the same profile. As with the

macrofossils, this profile was notably less rich (in terms of pollen concentrations) than Profile C. This may be

explained in terms of Profile Z lying closer to the centre of the fluvial channel (Scourse et al. 2009), Profile C being

situated in a marginal backwater in which the vegetation cover was more dense, and in which plant detritus

accumulated.

The plant macrofossil data from Unit C1 and the bone scrape samples indicate a water body possessing a modest

aquatic flora, possibly dominated by stoneworts (calcareous algae; family Characeae). These are typical colonizers

of newly created freshwater ponds and lakes of calcareous status. In contrast the high, and increasing, concentration

of Pediastrum (Figures 3 and 4), dominated by P. boryanum typicum, in Unit C1, is indicative of eutrophic water

with high nutrient levels and low oxygen content (Salmi 1963). The absence of P. duplex in Unit C1 may be

significant in that it is not tolerant of highly eutrophic conditions (Salmi 1963; Cronberg 1982).

There were probably fringing plant communities of limited extent with sedges (Carex), club-rush (Scirpus

lacustris) and marsh cinquefoil (Potentilla palustris). Terrestrial plants, whose remains must have been blown in or

washed into the basin through the fluvial channel which was active at this time (Scourse et al. 2009), include a few

taxa, which today would be considered weeds of cultivated and waste places, indicating unstable soils in the

vicinity. Thus, outside the basin itself the vegetation was dominated by grasses with a variety of taxa characteristic

of unstable minerogenic soils; these include allseed (Chenopodium polyspermum), goosefoots in C. Section

Pseudoblitum (probably C. glaucum or C. rubrum), orache (Atriplex sp.) and some of the Rumex sp(p). These

records, in association with thrift (Armeria maritima), are all indicative of a treeless environment. Lack of trees may

reflect cold climatic conditions though none of the taxa recorded are obligate arctic-alpine plants at the present day.

It is likely, however, that the remains of willows (Salix sp(p).) were all from dwarf forms typical of arctic-alpine

habitats. Alternatively, treelessness might be a response to extensive grazing pressure.

One of the plant taxa recorded is of interest from a palaeobotanical point of view. The pondweed from the

Profile Z samples, Potamogeton rutilus, appears not to have been recorded from the British fossil flora (there are

no records in Godwins (1975) compilation). It is commonly known as the Shetland pondweed, being recorded

within the British Isles only from those islands and the Outer Hebrides. Its distribution across Europe is in the

northern, central and eastern parts, as far north as northern Russia, but it is absent from Scandinavia. Such a

distribution lends support to the suggestion that Unit C1 accumulated in an essentially cool temperate to boreal

treeless environment.

The single pollen spectrum from Unit C3 indicates a very similar environment to that characterizing Unit C1;

in the correspondence analysis it plots very close to the C1 samples. The Unit C3 sample has relatively higher

values of Gramineae and lower values of Cyperaceae, and thus, shows a closer affinity to

Documents

Allen Et Al. (2009) -- Palaeoenvironmetal Context of Woolly Mammoth at Candover, UK