THE SPOTTED FROG IN WESTERN OREGON - …soda.sou.edu/awdata/030728a1.pdf · a In the context of this report, western Oregon refers to Oregon west of the crest of the Cascade Mountains

TechnicalReport#94-1-01 rUSPUBLIS

THE SPOTTED FROG IN WESTERN OREGON

Part I - BackgroundPart II - Current Status

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Oregon Department of Fish and WildlifeWildlife Diversity Program

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CONTENTS

PART I - THE SPOTTED FROG IN WESTERN OREGONIntroduction.........................................................................................................................1M e th o d .................................................................................................................................. .................................. 2Biological Synopsis .................................................................... 4

Background and Systematics .................................................................... 4Geographic Distribution .................................................................... 6Systematic Status of the Spotted Frog in Western Oregon ............................ 6Identification............................................................................................................ 10Habitat Requirements and Natural History ...................................................... 16

Historical Change ................................................................... 20Summary and Recommendations ................................................................... 27Acknowledgements............................................................................................................ 29Literature Cited ................................................................... 30Figures

1. The historic distribution of the spotted frog in western Oregon2. Color plate of an adule female western spotted frog3. Comparisons of shank and body length measurements for the northern

red-legged frog and the western spotted frog4. Comparisons of shank and body length measurements for the Cascades frog

and the western spotted frog5. Data on collections of western spotted frogs from western Oregon6. Data on collections of northern red-legged frogs from western Oregon

AppendicesI. BibliographyII. Chronological BibliographyIII. Western Spotted Frog LocalitiesIV. Unverified Localities

PART II - CURRENT STATUS OF THE SPOTTED FROG IN WESTERN OREGONAbstract................................................................................................................................1Introduction.......................................................................................................................1Method................................................................................................................................3Results.................................................................................................................................3Conclusions........................................................................................................................5Recommendations............................................................................................................8Acknowledgements..........................................................................................................9Literature Cited ................................................................... 10Tables

1. Sites checked asssociated with historical spotted frog localities2. Unverified spotted frog locality site checks

Erroneous spotted frog locality site checks SOUTHERN OREGON UNIVERSHTY LIBMAY3. Historical spotted frog site checksASLNOE N9724. Western spotted frog numbers at sampled sites ASHLAND, OREGON 97520

Structure of spotted frog populations in western Oregon5. Western spotted frog detection and search time statistics

Figures1. The historic and current distribution of the spotted frog in western Oregon

AppendicesI. Other western Oregon localities visitedII. Other western spotted frog localities visited

THE SPOTTED FROG (Rana pretiosa) IN WESTERN OREGONa

Marc P. Hayes

Research AssociateDepartment of Biology, Portland State University,

P.O. Box 751, Portland, OR 97207-0751,

andResearch Section, Animal Management Division, Metro Washington Park Zoo,

4001 Canyon Road, Portland, OR 97221-2799

Introduction

Several species of-ranid frogs in the Pacific Northwest are thought to have declined(Hayes and Jennings 1986). Among them, the spotted frog (Rana pretiosa) is generallyregarded as having disappeared from parts of its historic range (St. John 1984a, 1985a,1987a, 1987b; Storm 1986; Marshall 1989; McAllister and Leonard 1990, 1991; Leonardet al. 1993; D. Green, R. Storm, pers. comm.)b. Despite this general agreement, and withfew exceptions (see McAllister and Leonard 1990, 1991 for western Washington),disappearances of the spotted frog lack documentation. The basis for agreement is limitednegative evidence that the spotted frog has not been detected in the absence of systematicsurveys in various parts of its range. Moreover, precise data delineating its historicdistribution are lacking. Absence of significant data was a primary reason that an earlierattempt to list the spotted frog in the state of Oregon (see Marshall 1989) failed (D.

Marshall, C. Puchy, pers. comm.)c. The present work addresses this data gap bydocumenting the historic distribution of the spotted frog in western Oregon and its apparentdecline, and by focusing on those aspects of its biology that help explain its presentcondition. These data are intended to provide the background necessary to conduct surveysneeded to evaluate the status of the spotted frog in western Oregon, to direct management

recommendations, and to organize future research that can refine those recommendations.

a In the context of this report, western Oregon refers to Oregon west of the crest of the Cascade Mountainsand the portion of the Klamath hydrographic basin that lies within the state.

b Additional citations (e.g., Hovingh 1988) discuss disappearances of spotted frogs, but these citationsapply to spotted frogs different from that which occurs in western Oregon. The systematic status ofspotted frogs in western Oregon is addressed in the section by that name.

c Recent attempts to list the spotted frog as Endangered throughout its range in the United States have alsofailed in part for similar reasons (see Miller 1989).

Hayes: Spotted Frog 2

Method

A comprehensive bibliography of the spotted frog was assembled from all available

sources working from the compilation of Turner and Dumas (1972) as a starting point.

WILDLIFE REVIEW and ZOOLOGICAL RECORD were searched in their entirely from 1935

and 1864, respectively. BIOLOGICAL ABSTRACTS was searched back through 1974, the

year its generic index was initiated (which allows a search by scientific name). To ensure

not overlooking recent obscure references, the following computerized indices were also

searched (the years covered indicated in parentheses): AGRICOLA (1979-1991);

BIOLOGICAL AND AGRICULTURAL ABSTRACTS (1983-199 1); and AQUATIC SCIENCES

AND FISHERIES ABSTRACTS (1982-1991). Annual or summary indices of journals likely

to contain articles with data on the spotted frog were also examined over the entire runs of

those journals; these included: THE AMERICAN MIDLAND NATURALIST, AMPHIBIA-

REPTILIA, THE CANADIAN FIELD-NATURALIST, THE CANADIAN JOURNAL OF ZOOLOGY,

COPEIA, ECOLOGY, THE GREAT BASIN NATURALIST, HERPETOLOGICA,

HERPETOLOGICAL REVIEW, JOURNAL OF HERPETOLOGY, JOURNAL OF PARASITOLOGY,

NORTHWEST SCIENCE, OECOLOGIA, OIKOS, and SYSTEMATIC ZOOLOGY. The reference

section of each spotted frog citation was examined for additional sources of data on this

taxon. All references containing original data on the spotted frog, however minimal, were

included in the final alphabetical compilation (Appendix I). References that mentioned the

spotted frog, but lacked either original data or interpretations were not included. The

alphabetical compilation (Appendix I) permitted creation of a chronological compilation

(Appendix II) useful for rapid retrieval of data to identify historical trends and to facilitate

the addition of future references.

Available records for the spotted frog were retrieved from academic, institutional, and

museum collections. Initially, records for all species of ranid frogs known, or suspected,

to occur within western Oregon and areas around its periphery were requested from each

institution. Institutions were then visited to examine the specimens corresponding to these

records; where institutions could not be visited, specimens were borrowed to verify their

specific identities. In all cases except for specimens from Brigham Young University, the

Carnegie Museum, the American Museum of Natural History, and the Academy of Natural

Sciences of Philadelphia, I verified the identity of specimens. For collections at the

California Academy of Sciences, the United States National Museum, the Museum of

Vertebrate Zoology (Berkeley), and the University of Michigan, Dr. Mark R. Jennings also


verified the identity of specimens. Dr. Jennings exclusively verified the material from the

American Museum of Natural History and the Academy of Natural Sciences of

Philadelphia. Dr. John S. Applegarth verified the identity of the Sweet Home specimens.

In many cases, Dr. Robert M. Storm had previously verified the identity of specimens.

Due to logistic difficulties, Carnegie Museum specimens were not verified. The museum

symbolic code and the specimen number from that museum collection follow the spotted

frog records discussed in the text (in brackets). Museum symbolic codes follow Leviton et

al. (1985) for the herpetological collections at the American Museum of Natural History

(AMNH); the Academy of Natural Sciences of Philadelphia (ANSP); Brigham Young

University, Monte L. Bean Museum (BYU); the California Academy of Sciences (CAS or

CAS-SU [old Stanford University collection]); Carnegie Museum of Natural History (CM);

University of Kansas, Museum of Natural History (KU); National Museum of Natural

History (USNM); Museum of Vertebrate Zoology, University of California at Berkeley

(MVZ); The University of Michigan, Museum of Zoology (UMMZ); and University of

Oregon Museum of Natural History (UOMNH). Some institutions lacked a symbolic code

in Leviton et al. (1985); codes (in parentheses) were provided for Oregon State University

Museum of Natural History (OSUMNH); Portland State University (PSU); Southern

Oregon State College (SOSC); James R. Slater Museum, University of Puget Sound

(UPS); Oregon Natural Heritage Database (ONHD); and University of California at Davis

(UCD). In addition, many individuals with field experience or knowledge of frogs were

either queried about spotted frogs, specific records, or other individuals that might have

pertinent knowledge. All individuals contributing information are listed in the

acknowledgments.

Selected measurements were taken to verify the identity of specimens. Only on well-

preserved (minimally distorted) material was measured. Body (snout-urostyle = SUL) and

shank (fibulo-tibia) lengths were measured with calipers to the nearest 0.1 mm. The

flexibility of specimens did not allow taking the body length measurements with the level of

precision the shank length measurements would allow. Thus, all measurements were

rounded to the nearest 0.5 mm, the level of precision that the less precise measurement

(body length) would allow.Data were extracted from the references in Appendix I to develop a synopsis of the

biology of the spotted frog. This synopsis emphasizes the following aspects of spotted

frog life history important to its management: 1) the habitat requirements of different life

stages; and 2) the characteristics of oviposition, refuge, and overwintering sites. As the


work leading to this report was begun, Dr. David M. Green (Redpath Museum, McGill

University, Montreal, Canada) and his student, Jennifer Kearsley, initiated a systematic

revision of the spotted frog based on morphometric data. This work revealed that several

species are represented within what is now recognized as the spotted frog (D. Green, pers.

comm.), and at least two species (perhaps three) occur within the state of Oregon. Thus, a

need existed to identify the probable geographic limits of the spotted frog species found in

western Oregon. This was essential because potential differences in the biologies of the

unrecognized species made it necessary to be able to link previous studies (based on their

geographic origin) to the spotted frogs occurring specifically in western Oregon. For this

reason, a discussion of the probable geographic limits of the spotted frog species occurring

in western Oregon is included. Some references containing data other than that on spotted

frogs were necessary to interpret aspects of the biology and the history of the spotted frogs

in western Oregon. Those references are listed after this text rather than being included in

Appendix I or II. Some biological data obtained from collections were also included in the

synopsis. Where appropriate, the museum symbolic code and collection number indicates

the specimen associated with those data (in brackets).

Biological Synopsis

I. Background and Systematics

The name spotted frog (Rana pretiosa) dates from 1853, when Baird and Girard (1853)

described a series of five frogs [USNM 11409; the five specimens in the type series have

the same number, see Cochran 1961] that the United States Exploring Expedition had

collected in May 1841 in the vicinity of Puget Sound (see Wilkes 1845, Tyler 1968,

Stanton 1975). Reflective of confusion that has long plagued identification of ranid frogs

in the western North America (see Dunlap 1955; Dumas 1966; Case 1978; Kluge et al.

1979; Post and Uzzell 1981; Uzzell and Post 1986; Green 1986a, 1986b), some the

specimens in the type series are not spotted frogs. The type series, which consists of an

adult female, two subadults, and two juveniles, contains two frogs that are northern red-

legged frogs (Rana aurora aurora)d. Until 1939, when James R. Slater of the College of

d The northern red-legged frog (Rana aurora aurora) is easily distinguished from the spotted frog (Ranapretiosa) found in western Oregon based on the longer shank leg, reduced foot webbing, prominent groinmottling, and finer dorsal spotting of the former (see section on identification). The two juveniles in thetype series agree with the northern red-legged frog in these features.


Puget Sound (now the University of Puget Sound) described the Cascades frog (Ranacascadae), that species was included under the name R. pretiosa (see Cope 1889, Meek andElliott 1899, Dickerson 1906, Van Denburgh and Slevin 1915, Grinnell and Camp 1917,Storer 1925, Slevin 1928, Campbell 1929, Grinnell et al. 1930, Fitch 1936, Gordon1939). After Slater described it, acceptance of the Cascades frog as a species distinct fromR. pretiosa was slow, a situation probably resulting from the sparse diagnosis Slater hadprovided. Skepticism about the status of R. cascadae led Stebbins (1954) to regard it as asubspecies of R. aurora until Dunlap (1955) compared Cascades, spotted, and northernred-legged frogs in a definitive manner, and demonstrated their morphological

distinctiveness. Dunlap described differences that were thought to be constant across therange of the three species, even though the samples used for comparison came almostexclusively from a portion of central Oregon lapping both sides of the Cascade Mountains.The fact several species may be contained within R. pretiosa makes Dunlap having focusedon the indicated region advantageous because its includes a large portion of the study areaaddressed here.

The possibility that R. pretiosa includes more than one species is not new; it has longbeen recognized that populations over a significant portion of the western part of the rangeof R. pretiosa consist of individuals with a reddish or red-orange ventral coloration thatcontrasts sharply with the yellow- or yellow-orange-ventered individuals from someinterior populations (see Dunlap 1955, Nussbaum et al. 1983, Storm 1986). Indeed,Thompson (1913) described the subspecies, Rana pretiosa luteiventris, on the basis of aseries of frogs from northern Nevada with a yellow ventral color and one less tubercle onthe feet (described as present in R. p. pretiosa). Difficulties with evaluation of color aswell as different assessments of the distribution of color and tuberculed forms resulted inlack of acceptance of the described subspecies (compare aforementioned references andStebbins 1951, 1954, 1966, 1985; and Turner 1959b) and may have inhibited attempts atstudy to faciliate recognition of different species. Detection of significant chromosome andprotein variation among geographically segregated populations of R. pretiosa (Green1986a, 1986b) was an important impetus to its current systematic reevaluation (D. Green,pers. comm.). The latter methods lack some of the bias inherent in evaluating differencessolely on the basis of external morphology.

As currently understood, R. pretiosa is thought to belong to a-group of closely relatedfrogs termed the Rana boylii group (Case 1978). Although the composition of the Rana

boylii group has been the subject controversy (Case 1978; Kluge et al. 1979; Post and


Uzzell 1981; Hillis and Davis 1983; Uzzell and Post 1986; Green 1986a, 1986b), workersagree that in addition to the spotted frog at least five other frog taxa, all native to the Pacificslope drainages of the western United States, are members of this group. These five taxaare the northern red-legged frog, the California red-legged frog (R. aurora draytonii), thefoothill yellow-legged frog (R. boylfi), the Cascades frog, and the mountain yellow-leggedfrog (R. muscosa). General agreement also exists that R. pretiosa is the most primitive ofthese six taxa (Case 1976; Hillis and Davis 1983; Green 1986a, 1986b). However, sincethe ongoing work of Green and Kearsley suggests that several species are contained withinR. pretiosa, and previous studies attempting to define relationships within the Rana boyliigroup have used spotted frogs that represent more than one of the unrecognized taxa,evaluation of the relationship of these taxa (now R. pretiosa) await their systematicresolution.

II. Geographic Distribution

As currently understood, the spotted frog as a relatively broad geographic range.Stebbins (1951, 1954, 1966, 1985) and Turner and Dumas (1972) mapped the overallrange. That range extends from northeastern California northward through most of Oregon(Gordon 1939), Washington (Slater 1955), and British Columbia (Carl 1943, Green andCampbell 1984) into the Alaskan panhandle (Hock 1953, Logier and Toner 1961, Hodge1976), and eastward through northern Nevada (Linsdale 1940, Banta 1965), northern Utah(Van Denburgh and Slevin 1915, Hovingh 1986), most of Idaho (VanDenburgh and Slevin1921, Waitz 1961), western Wyoming (Test 1891, Cary 1917, Dunlap 1977), westernMontana (Test 1891, Black 1969), and the western edge of Alberta (Stebbins 1985).Records from central Saskatchewan (Logier and Toner 1961) are in error (Cook 1964).This view of the geographic distribution ignores unrecognized taxonomic units within thespotted frog.

III. Systematic Status of the Spotted Frog in Western Oregon

The spotted frog likely comprising more than one species required that a preliminarysystematic determination be made to determine whether specimens from western Oregon allbelong to the same taxonomic entity and, more importantly, to link the biological literatureto the spotted frog species found in western Oregon. Examination of specimens from


collections and descriptions from the literature associated with those specimens led to the

following conclusions:

1) Specimens of the spotted frog from western Oregon appear to represent one species

(all specimens associated with the localities in Appendix I), hereafter referred to as the

western spotted frog. This assessment is based on the fact that all spotted frogs taken or

observed in western Oregon share a prominent, distinctive morphological feature: a

characteristic mottling of some or all of the ventral surface of the abdomen, as described in

the section of this report on identification.

2) For the few specimens for which color data are available, individuals of the spotted

frog from western Oregon are consistently the red/red-orange-ventered color variant (see

Nussbaum et al. 1983). While this feature is not unique to the western spotted frog (see

below), no indications exist that spotted frogs from western Oregon are other than the

red/red-oranged ventered color variant.

3) Outside western Oregon, spotted frogs with the ventral mottling similar to those

found in western Oregon are distributed northward through the state of Washington west of

the Cascade Mountains [AMNH 34-35; USNM 5975, 9421, 11409 (3 specimens), 35638-

9, 45866-7, 312413-4; UPS 1441, 1444, 1446-8, 2069-71, 2100, 2134-7. 2667, 2712-3]

into the coastal plain in extreme southwestern British Columbia (Licht 1969a, 197 lb), and

east of the Cascades in Washington north to at least the vicinity of Trout Lake, Klickitat

County [USNM 61473-4; UPS 5597-5600, 5607, 7372]. Recently, Dennis Paulson

(James R. Slater Museum, University of Puget Sound) discovered spotted frogs which

have this same morphology on Conboy National Wildlife Refuge, just a few miles

northeast of Trout Lake (pers. observ.). Similar spotted frogs also occur in eastern Oregon

through much of the Deschutes River system [MVZ 7464, 7490-6, 26731, 26739, 26764,

26780, 26785-815, 98627-8, 178556-60; USNM 11513, 11521, 11892, 13770], and in

northeastern California from the lower Klamath Basin [CAS 4491-2] and the upper Pit

River system (Modoc Plateau) [MVZ 2098-99; USNM 38806]. Where color data are

available from within the range of similar ventrally mottled frogs outside western Oregon,

they are all red/red-orange in ventral coloration (see Dickerson 1906, Nussbaum et al.

1983, and Storm 1986 in part; see subsequent paragraph for explanation of the latter).

Frogs in the drainages of the closed basin associated with Harney and Malheur Lakes

[CAS-SU 1601, OSUMNH 847-9] have red/red-orange ventral coloration (Storm MSe and

e An unpublished MS by Storm entitled "Remarks on some Oregon amphibians in need of consideration forprotection" dated 1976 and obtained from ODFW files mentions that red-ventered populations


1986; pers. observ.). However, photographs taken by Maurita Smyth of adult spotted

frogs with red-orange ventral coloration from Page Springs (located at the south end of

Malheur National Wildlife Refuge and, thus, within this closed basin system) show no sign

of ventral mottling, a feature shared by the aforementioned specimens from this basin.

Lack of ventral mottling is exhibited by spotted frogs from northeastern Oregon, Idaho,

and points further east. Thus, spotted frogs from the Harney-Malheur system do not

appear to represent the western spotted frog.

4) Ventrally mottled spotted frogs with a red/red-orange ventral coloration clearly differ

biochemically from the spotted frogs elsewhere (D. Green, pers. comm.). Enough data

exist (samples from Trout Lake and Conboy National Wildlife Refuge in Washington, and

Gold Lake, Davis Lake, Wood River (Klamath Basin) and Paulina Marsh in Oregon) to

suggest that the ventral mottling observed in this study is correlated with the above noted

biochemical differences, and thus, that the western spotted frog probably represents a valid

species. Although the eastern limit of the western spotted frog in much of Oregon remains

vague, the allocation of spotted frogs in much of the closed basins between the Klamath

system and Harney-Malheur needs to be established, biochemical data suggests that the

western spotted frog ranges east at least as far as Paulina Marsh (Lake County; D. Green,

pers. comm.). Some spotted frog material does exist in collections for the closed basin

systems south of between Harney-Malheur and Paulina Marsh (i.e., Warner's 2nd Lake

[Hart Lake; ANSP 14592], 12-mile Creek to Warner's 2nd Lake [12-mile Creek near Hart

Lake; ANSP 14593], and at the north end of the Warner Mountains [USNM 26000]), but

all this material is old and cannot be confidently allocated to the spotted frog taxa discussed

here.

5) The western spotted frog is not recorded at elevations above 5,000 ft (1,524 m)

anywhere in Oregon (it closest approach to 5,000 ft is in the central and southern Oregon

Cascades), whereas spotted frogs lacking the morphology of the western spotted frog in

eastern Oregon are recorded up to nearly 8,000 ft (2,438 m) at a latitude which is actualy

somewhat further north that highest elevations reached by the western spotted frog. This

may indicate a fundamental difference in physiology related to the apparent taxonomic

differences between these frogs.

For these reasons, it seems best to regard the spotted frogs in western Oregon as a

single taxon, called here the western spotted frog until as formal taxonomic designation is

(presumably the western form, see subsequent section) occur in the Blitzen [=Donner und Blitzen] Riversystem of southeastern Oregon.

V


provided. This taxon is characterized by a mottled ventral pattern and colored wash which,in adults, is probably consistently red/red-orange. This taxon seems to extend from BritishColumbia to California, and seems to be the only form of the spotted frog west of theCascade Mountains in Oregon and most of Washington (another form of the spottedappears to extend just west of the Cascade crest in the subalpine region of the northCascades in Washington). Verifiable records of the western spotted frog in Oregon arediscontinuously distributed over roughly the western two-thirds of the state (Figure 1).This spotted frog is absent from most of the rest of eastern Oregon; there one or perhapstwo spotted frogs with a bare (unmottled) abdomen are present. In northeastern Oregon,these unmottled frogs are yellow or yellow-orange beneath (Dunlap 1955, Nussbaum et al.1983, Storm 1986).

Distribution of the spotted frog in western Oregon is mapped in Figure 1. Verifiablerecords for the western spotted frog exist for scattered locations over roughly the easterntwo-thirds of the Williamette Valley below ca. 600 ft (180 m) in elevation, and in theremainder of the Willamette hydrographic basin on the west slope of the CascadeMountains between 3,200 ft (975 m) and 5,000 ft (1,524 m). A rather large hiatus existsin elevation between the records on the Willamette Valley floor and those on the western

slope of the Cascades; the reason for this hiatus is unclear, but it may be related to habitatrequirements (see Habitat Requirements and Natural History). The spotted frog is alsorecorded in the Klamath Basin and upper Klamath River system at elevations between ca.4,000 ft (1,220 m) and 4,400 ft (1,340 m). Notably, no verifiable records of either thisform of the spotted frog or any other spotted frog exist for coastal or near coastal areas inwestern Oregon (all coastal region records are either known to represent the northern red-legged frog or are questionable), the higher Cascade Mountains (all records found to datethat are claimed to be spotted frogs from the west slope of the Cascade Mountains above5,000 feet are attributable to the Cascades frog [contra Dunlap 1955]), and the Umpquadrainage basin (see Appendix V). Furthermore, the few records for spotted frogs from theRogue River system are not verified (see 2 and 5 in Appendix IV), so some possibilityexists that spotted frogs are also absent from the Rogue hydrographic basin. The lack ofcoastal and high elevation records for the spotted frog in western Oregon may be related toa need postmetamorphic stages have for warmer (2 200C) water (see Habitat Requirementsand Natural History).

1250 1240 1230 1220 1210

-E)- ~~~~~~~~IIIan46

0- = 460

S

Localities

* Verified 3 450

o Unverified

o Erroneous I

3 ~~~~~~~~~440

t~~~~~ ~14y! < 48

c"- 2 ~~~~~~~~~~~~~~~430

"k~~~i~~~- 6-

0-4

12

420

Dashed line indicates the eastern limit of western Oregon as described in the text.

I I ~ ~ ~~~~I I 11250 1240 1230 122° 121'

Figure 1. The historic distribution of the spotted frog (Rana pretiosa) in western Oregon. Numbers foreach locality correspond to the appended lists for verified (Appendix III), unverified (Appendix IV), anderroneous localities (Appendix VJ. Solid lines are county boundaries.


IV. Identification

Identification here is addressed at two levels: 1) Distinguishing morphologically the

entity I term the western spotted frog from all other spotted frogs; and 2) Distinguishing

morphologically the western spotted frog (and perhaps other spotted frogs) from the other

two congeneric ranids in the Pacific Northwest, the northern red-legged frog (Rana aurora

aurora) and the Cascades frog (Rana cascadae).

1) Distinguishing the western spotted frog from all other spotted frogs - As noted

previously, the western spotted frog can be distinguished morphologically from all other

spotted frogs by the presence of mottling on at least some of the ventral portion of the

abdomen. This mottling, albeit a somewhat faint representation, is illustrated in Figure 2.

Mottling is the result of melanophore clusters such that the ventral skin is partly or entirely

suffused with relatively dark-colored patches, irregular in shape, but with a more or less

regular spacing. In living western spotted frogs, this mottling is evident even if the mottled

region is colored over by the characteristic red/red-orange wash. My own observations of

western spotted frogs in Oregon and Washington indicate that this mottling can range from

dramatically dark to extremely pale in the same individual in the course of a few minutes.

Moreover, a clear lightening of this mottling is apparent after handling a frog for a few

minutes, suggesting that lightening, presumably by retraction of the melanin granules in the

stellate arms of the melanophores, is linked to stress or an excited state. This suggests that

the relatively faint mottling observed in many of the preserved specimens in collections may

be unrelated to the age of the specimen, but rather to the state of the frog just before it died;

it may also well explain the Dickerson (1906) plate of a western spotted frog, in which the

mottling is visible, but faint (see Figure 2). Western spotted frogs I have observed had

darker mottling immediately after capture unless they were not handled for a period of time,

in which case they would darken again. Mottling can be also observed in individuals that

have recently metamorphosed, although a considerably smaller area of the belly may be

mottled. The latter observation and the fact that the larger adults appeared to have the

greatest degree of mottling suggests that mottling may increase in extent and/or density with

age; large spotted frogs may have a prominent ventral mottling reminiscent of that seen in

bullfrogs (Rana catesbeiana).

Features of the larval stage or eggs that might distinguish the western spotted frog from

other spotted frogs have not been examined.

-�- --- -, --- 711

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Jamrc;l

VA 41�4wiv.0 . �41

Figure 2 - Color plate of all adull lemale western spotted frog. Iromi Dickerson (1900).Note the grayish inottling on the venter of the Female and horseshoe-shapedpattern of orange color wash.

Hayes: Spotted Frog ' 11

2) Features used to distinguish the western spotted frog from the northern red-leggedfrog and the Cascades frog - Several features are useful to differentiate the western spottedfrog and its close congeners, the northern red-legged frog and the Cascades frog. Thesefeatures vary considerably in their ability to distinguish these species. As Dunlap (1955)emphasized, character variability can make reliance on any single feature to distinguish thespotted frog from its congeners tenuous, but if used simultaneously, the combination ofcharacters can provide a high degree of confidence in distinguishing the three species.Identification of specimens in collections was based largely on the features Dunlap (1955)originally emphasized with a few caveats. The primary caveat is that the distinguishingcharacteristics apply to the western spotted frog, and especially those found in westernOregonf. These features may allow distinguishing other spotted frogs (other than thewestern spotted frog) from northern red-legged frogs and Cascade frogs, but they shouldbe used cautiously outside of western Oregon until the variation in other spotted frogs isunderstood. A second caveat is that these features apply to the postmetamorphic stagesbecause the only available keys to the larval stages of ranid frogs depend in part ongeographic data and in part on data that likely represents more than one species of spottedfrog (see Altig 1970). Thus, such keys are unable to effectively distinguish between thespotted frog found in western Oregon and its close congeners. Where appropriate, thereliability of each feature for distinguishing the respective species at different post-metamorphic stages is noted. Characteristics are listed based on how reliably theydistinguish these three frogs; the first four (a-d) are more reliable than the latter three (e-g).

a) The length of the lower leg or shank (fibulo-tibia) is significantly shorter relativeto the body length in the spotted frog than in either the Cascades frog or the northernred-legged frog, and reflects the shorter overall length of the legs in the former (Wrightand Wright 1949, Dunlap and Storm 1951, Stebbins 1951, Dunlap 1955). Scatterplotsaid visual comparison of the relative length of the lower leg of the western spotted frogwith each of the Cascades frog and the northern red-legged frog (Figures 3 and 4). Thegreatest difference in the relative length of the lower leg is between the western spottedfrog and the northern red-legged frog (Figure 3). This feature was significant in helpingdetermine that the two smallest individuals in the type series of the spotted frog (blacksquares) were actually northern red-legged frogs (the remaining three frogs in the type

f As noted in the section on distribution, even the careful work of Dunlap (1955) included a few specimensof Rana cascadae in his spotted frog sample. Features of spotted frogs in western Oregon summarizedhere excludes these high elevation specimens attributable to R. cascadae.

Figure 3 - Comparison of shank and body length measurements for the northern red-leggedfrog and the western spotted frog.

50'-

40~~~~~~~~~

30-

Shank TLength V50;w~(mm) 8PQ

20 -cp 09 O Spotted frog

oko Northern red-legged frog

10 3 Spotted frog (types)

* Juvenile "spotted frogs" (types)

0 -o10 20 30 40 50 60 70 80 9 100 110 120

Body Length (mm)

Figure 4 - Comparison of shank and body length measurements for the Cascades frog andthe western spotted frog.

50 I

40

30 -

0

00 4

1atp

8; 0 Cascades fro

a> * Spotted frog4P~~~~~~~~~~~~~

60

0

I

ShankLength(mm)

20 -

10 -

9o

0eo

a IKnown hybrids

a Hidden Lake Specimen

u I I I I I I .0 10 20 30 40 50 60 70 80 90 100 110 120

Body Length (mm)


series [white squares] have lower leg lengths that correspond to the expected range forspotted frogs [black circles]). Less difference exists between the Cascades frog and thespotted frog (Figure 4). Nevertheless, relative leg length can effectively distinguish allbut the smallest Cascades frogs from spotted frogs; juveniles smaller than 40 mm SULwere indistinguishable using solely this character. The two known hybrids between theCascades frog and the spotted frog (see Green 1985) were not consistent in relativelower leg length, one [MVZ 193588] has a relative lower leg length closer to that of theCascades frog, whereas the other [MVZ 193589] is closer to that typical of the spottedfrog (see Figure 4). This result emphasizes the idea that hybrids cannot be identifiedbased on this feature (see Green 1985). Notably, this character helped identify thepeculiarly patterned frog from near Hidden Lake (USNM 294912) that Steve Corn hadidentified as a spotted frog as a Cascade frog because of its relatively longer lower leglength, typical of that species (Figure 4; see also 3 in Appendix IV). A useful estimatoris that for the western spotted frog, the lower leg is generally less than half the bodylength, whereas length of the lower leg generally exceeds half the body length in boththe Cascades frog and the northern red-legged frog. This feature is high reliable aslong as variation because of deformities or leg injuries is recognized (measuring bothshanks can help recognize the latter), and measurements are taken in a uniform manner(one individual taking measurement can considerable reduce variability). This featurecan also sometimes help to distinguish unmetamorphosed northern red-legged frogsfrom unmetamorphosed spotted frogs where sufficient development of the hindlimbshas occurred prior to metamorphosis. This feature was used in lieu of the frequentlyused extended leg-adpressed-to-body measurement to determine leg length by whetheror not the heel extends beyond the eye (e.g., Leonard et al. 1993). The latter, if donerigorously, can damage historically valuable specimens or injure potentially even morevaluable live frogs.

b) Webbing of the hindfeet is more extensive on the spotted frog than in either theCascades frog or the northern red-legged frog (see Nussbaum et al. 1983, Stebbins1985). Specifically, the spotted frog has nearly full webbing on the hind feet, bestviewed when toes III (the middle toe) and IV (the longest toe), and IV and V (theoutermost toe) are spread so that the webbing is reasonably taut. If any indentation ofthe taut webbing in the spotted frog exists, it never dips lower than across from the first


subarticular tubercles on toe IV (the longest toe). In contrast, the webbing of theCascades frog is indented consistently to at least the level of the second subarticular

tubercle, and the webbing of the northern red-legged frog is indented consistently to at

least the level of halfway between the second and third subarticular tubercle and is often

indented below the level of the third subarticular tubercle. This feature is difficult to

assess if the webbing is deformed or torn, so it should be routinely examined on bothfeet to help ensure that at least one set of webbing examined is intact. As with the

relative length of the lower leg, I used this feature to identify unmetamorphosed

western spotted frogs where webbing of the hind feet was developed enough to reflect

the pattern seen in older individuals (e.g., see CAS-SU 5906; 15 in Appendix III).c) The eyes are more upwards oriented in the western spotted frog than in either the

Cascades frog or the northern red-legged frog. Specifically, in the spotted frog, theeyes are oriented upwards so that well over 50% of the surface of the open eyes are

visible when viewing the frog directly from above (bird's eye view). In contrast, the

orientation of the eyes in Cascades or northern red-legged frogs only allows viewingconsiderably less than 50% of the surface of the eyes from the same position. Another

way of assessing this feature is that if frogs are viewed at about a 450 angle from one

side of the body (in the plane perpendicular to the long axis of the body), the cornea

(surface) of the more distant open eye can be easily seen in the western spotted frog,

but it is hidden from view in Cascades and northern red-legged frogs in the same

position. This difference can be difficult to assess without previous experience with the

three species; it is most obvious in a parallel comparison (see Figure 14 in Stebbins

1966; and Figure 13 in Stebbins 1985). This feature is useful for distinguishing all

sizes of postmetamorphic individuals, but metamorphosing individuals of the spotted

frog that have not ungone the dorsal rotation of the eyes cannot be identified based on

this character alone.

d) The mottled pattern of the skin on belly and chest region used to distinguish the

western form of the spotted frog from other spotted frogs (see number 1 in this section)

also constrasts markedly with the unmottled skin of the Cascades frog and the weakly

mottled condition in the northern red-legged frog. Except for the yellow or red pigment,

respectively, the Cascades frog and northern red-legged frog, are almost immaculate over

the central belly and chest.

g Subarticular tubercles are the small bumps beneath the digits of both hind and front limbs that correspondto a point immediately below where the digit bones (phalanges) flex or join.


e) Western spotted frogs typically have inky black, ragged-edged or lichen-shapedspots that often have light centers matching the ground color of the back of the frog

(Dunlap 1955; see also Figure 2). In contrast, Cascades frogs have inky black, but

relatively smooth-edged spots that typically lack light centersh, or less frequently, are

unspotted. Spotting in the northern red-legged frog, if present, consists of small,

diffuse or peppered spots that are typically not prominently inky black. Inky blackmarkings, when they occur on northern red-legged frogs, typically line the dorsolateral

folds that extend from the eyes to the shoulders. This feature is really useful for

identifying frogs > 50 mm SULi; it is rarely useful for distinguishing juveniles of

Cascades frog from those of the spotted frog, both of which often have light-centered,

inky black spots that typically have relatively smooth edges. Even juveniles of northern

red-legged frogs often have light-centered spots, although they are rarely inky black.

Spotting changes with age have not been studied, but the differences between juvenileand adult western spotted frogs examined suggests that the dorsal spots become more

ragged-edged as individuals mature.f) The degree of groin mottling (the pattern on the patch of skin between the upper

hindlimb and the side of the body) is less developed in the spotted frog than either the

Cascades frog or the northern red-legged frog (see Figure 2 in Dunlap 1955). Dunlap

emphasized that the groin is almost unmarked in most spotted frogs in contrast to

weakly marked groin on a honey-colored field displayed by the Cascades frog and the

prominently marked groin on a white, sun yellow, red, or green field in life (white or

pale yellow in preserved specimens) in the northern red-legged frog. However, the

spotted frog sample that Dunlap generalized from included spotted frogs from the MillCreek and John Day River areas of eastern Oregon that are likely not western spotted

frogs, and a few individuals that are actually Cascades frogs from the vicinity of CraterLake. If one considers only the western spotted frogs, in all individuals examined, the

groin area lacks mottling. A key difference Dunlap did not address is that the weak

groin mottling in the Cascades frog contrasts little or not at all with coloration on the

rest of the side. This condition differs sharply with the highly contrasting groin patch

h Adult Cascades frog may have clusters of spots around a light-colored area, but this should not beconfused with a light-centered spot, which occurs infrequently in adult Cascades frogs.

Although Cascades, northern red-legged, and spotted frogs mature at slightly different body sizes (seeStorm 1960, Licht 1974, Briggs 1987), 50 mm is a body size close enough to the minimum adult size ofmales in all three that it represents a good compromise size above which features, like spotting, appearrelatively reliable for identification.


of the northern red-legged frog. In northern red-legged frogs, groin mottling appears

to become pronounced in frogs nearing adult size; thus, it is most useful for to

identifying individuals that are 2 50 mm SUL. This feature may also be useful to

differentiate the western spotted frog from other spotted frogs, but data will probably

have to be gathered from living frogs to establish a satisfactory understanding of what

variation in pattern may exist.

g) Differences in texturing of the skin on the lower back and the dorsal surfaces of

the hindlimbs can also assist diagnosis (Dunlap 1955). These areas are typically quite

rough or pustular in the western spotted frog in contrast to the variable, but generally

intermediately rough condition in the Cascades frog and relatively smooth condition in

the northern red-legged frog. Moreover, the degree of tubercularity or skin texturing

seems to increase with size (age), so comparisons increase in validity where individuals

close to the same size are compared. This feature is also difficult to assess without

previous experience with the three species, and it is most easily identified if one can

examine a western spotted frog next to either a northern red-legged frog or a Cascades

frog. It is a feature best used for diagnosis at a population level when a several frogs

are available for examination.

Color of the undersurfaces of frogs, typically a useful character in life, was useless to

differentiate preserved material. However, if color notes existed associated with specimens

in which it appeared one could have confidence, these were used to help verify their species

identities. In the latter case, the brightly pigmented undersurfaces of the western spotted

frog have a superficial-appearing red or red-orange pigment that has been described as

appearing painted on, whereas those of the Cascades frog were dull to bright yellow or

buff and those of the northern red-legged frogs were a dark to pale carmine, red, or scarlet

(Dunlap and Storm 1951, Dunlap 1955, Nussbaum et al. 1983; pers. observ.). In the latter

two species, the brightly colored pigments appear more deeply imbedded in the skin. The

distribution of pigment may also be useful for distinguishing the western spotted frog from

other taxa as adults. Western spotted frogs often have a incomplete ring of the red/red-

orange pigment around the belly (see Figure 2), a pattern I have not observed in yellow or

red pigmentation of Cascade and northern red-legged frogs. Moreover, the frequency with

which this feature appears in other spotted frogs is not known, so an analysis of the

variation in the color wash of these ranid frogs might reveal additional characters useful in

distinguishing species.


One feature is useful for distinguishing advanced tadpoles (Gosner (1960) stage 2 25and usually at least 30 mm total length) of western spotted frogs from the Cascades andnorthern red-legged frogs (see Altig 1970). Altig used this feature in his key to distinguishtadpoles of all spotted frogs from red-legged frogs (sensu lato; this includes California and

northern red-legged frogs) and other ranid frogs. Since the larvae upon which Altig basedhis key were those of the western spotted frog (R. Altig, pers. comm.), it is not knownwhether this feature will also distinguish other spotted frogs from the Cascades frogs andnorthern red-legged frogs. Altig (1970) describes the spotted frog (sensu lato)as having a

relatively long tail, characterized by a total length-to-body length ratio (see Altig for adefinition) of 2.6 or more, whereas red-legged frogs (sensu lato) have a shorter tail,

characterized by a total length-to-body length ratio of 2.6 or less. I have observed thatwestern spotted frog larvae (n = 17 from three different localities), which are at theadvanced stage I describe above, have a total length-to-body length ratio of 3.0 or more,whereas the Cascade frog and the northern red-legged frog each have total length-to-bodylength ratios of 2.7 or less (n = 41 for 13 different localities for the Cascade frog; n = 67for 17 different localities for the northern red-legged frog). This is feature will distinguishlarvae as long as they are old enough, and if a device (a transparent plastic, water-holdingtrough) is available in which larvae can be measured with some precision.

Only one additional feature is currently available to distinguish the early larval stagesstages of western spotted frog from those of other ranid frogs, a condition largely due tothe lack of detailed study devoted to the early larval stages of frogs. Immediately post-hatching, northern red-legged frog larvae (tadpoles) can be distinguished from larvae ofboth western spotted frogs and Cascade frogs by the much shorter gill filaments possessedby the former. Both of the latter have gill filaments that are 2.5-4 times as long. However,this feature cannot distinguish immediately post-hatching larvae of Cascades frog fromthose of the western spotted frog.

V. Habitat Requirements and Natural History

Most of what is known of the natural history of the western spotted frog is based on thestudies of Licht (1969a, 1969b, 197 1a, 197 1b, 1972, 1974), although a number of otherfield workers have contributed some information. The extensive studies of Turner (1957a,1957b, 1958a, 1958b, 1959a, 1959b, 1959c, 1960a, 1960b) and Morris and Tanner(1969) on spotted frogs in Wyoming and Utah should not be used to evaluate the behavior


of frogs in western Oregon, because these studies probably address different species (D.

Green, pers. comm.). Licht conducted his studies of the spotted frog along the Little

Campbell River in extreme southwestern British Columbia, so differences in the behavior

of the western spotted frogs Licht studied and those at various points in western Oregon are

likely restricted to minor variations, seasonal or otherwise, related to the differences among

these localities in elevation and latitude.

Overwinterine. The western spotted frog is generally inactive during the winter

season, although some individuals may be observed at the water surface on the few

relatively warmer days (Dickerson 1906p. The only overwintering site recorded has been

in the mud bottom of a marshy lake margin in water at least one foot of depth (Dickerson1906). Despite the paucity of data on overwintering sites, underwater overwintering sites

are anticipated because that pattern is typical of many ranid frogs (Emery et al. 1972). The

western spotted frog has been characterized as a highly aquatic species (Jewett 1936,

Nussbaum et al. 1983); as a consequence, the bodies of water that serve as overwintering

sites may be the same ones which the western spotted frog uses for breeding and in which

it spends the summer season, but no data yet exist to verify this supposition.

Reproduction. Emergence from overwintering sites begins as early in the year as the

winter thaw allows (Licht 1969a). Licht believed that a 5.00C (410F) air temperature was

the minimum needed to initiate activity, but because the overwintering site is likely

underwater, water temperatures may be the better predictor of emergence. However, water

temperature measurements during emergence intervals are lacking. Whatever the precise

physical conditions, in southwestern British Columbia and the Puget Sound region,

emergence has taken place from late February to mid-March (Dickerson 1906, Licht

1969a). Emergence dates are lacking for Oregon, but historical records indicate that

western spotted frogs were detected the Willamette Valley floor as early as 8 February

(Nussbaum et al. 1983). These frogs were often moving on wet nights during February

and March, the interval when the Willamette River experiences its freshet (US Army Corps

of Engineers 1982), which flooded shallow wetland areas (see 1, 18, and 20 in Appendix

III); these observations may have occurred soon after emergence. A single water

i Despite the early date of Dickerson's book, her comments on R. pretiosa represent the western formbecause her correspondent regarding this species, Dr. J. F. Illingworth of Seattle, Washington, obtainedall the spotted frog material he sent Dickerson from the vicinity of Seattle in 1905 (e.g., see AMNH 34and 35). Moreover, the color illustration in Dickerson's book of the ventral view of an adult femalespotted frog shows the mottling across the belly characteristic of the western form. The mottling in thisillustration is faint, which is characteristic of spotted frogs in a stressed or excited state, because the darkchromatophores tend to be more retracted in this condition.


temperature taken at 22:30 at night on 25 February 1957, at the time a western spotted frog

was taken ca. 11.2 km [7 mi] east of Corvallis (17 in Appendix III), was 10.60 C [51TF],

which suggests that even early in the active season, the western spotted frog has been

found in relatively "warm" water.

Male western spotted frogs arrive at breeding sites several days before the first females

appear. Breeding sites are located in the shallow (5-15 cm [=2-6 in]) portions of marshes

or ponds or the overflow areas of streams, typically disconnected from the main body of

water (Dickerson 1906, Licht 1969a). In southwestern British Columbia, males have been

observed to begin calling in the afternoon in full sunlight when air temperatures reach at

least 12'C (Licht 1969a); males continue to call into the evening unless temperatures drop to

near freezing, in which case calling ceases. Adult males aggregrate in small calling groups

(8-10 or more individuals), which presumably represent leks, and call while floating with

their heads at the water surface or while sitting above water on mats of vegetation; unlike

northern red-legged frogs and some other western ranids, they have not been observed to

call underwater. Mating calls of males of the western spotted frog are weak, carrying 15-

30 m (50-100 ft) at most; if Pacific treefrogs (Pseudacris (=Hyla) regilla) are calling

simultaneously, the calls of the western spotted frog can be easily masked. Western

spotted frog mating calls consist of 4-26 short bass notes with dominant frequencies of

0.5-1.5 kHz (Licht 1969a). Females appear at breeding sites from a few days to over a

week after the males. When receptive, females approach male calling groups, gain

amplexus with a male, and then deposit eggs in a few inches of water (Licht 1969a). In

southwestern British Columbia and the vicinity of Puget Sound, egg masses were laid in

March; the only Oregon oviposition record is a 10 April date for Portland ([MVZ 28067-8];

see 22 in Appendix III). The dates of oviposition likely vary considerably between years

because the timing at which local water temperatures reach the range suitable for egg laying

are probably highly dependent on the local climatic conditions and temperature regime.

Licht (1971 a) found that western spotted frog embyos have lethal thermal limits of 60 C and

280C, and noted that the water temperature surrounding 19 egg masses deposited in the

field averaged 20.70 C (s = 2.7) over the interval before hatching. Notably, western spotted

frog exhibit "communal" laying; five to 26 egg masses have been observed clustered

together in one small area (Licht 1969a). This may occur because males often call from

locations where other egg masses have already been laid, or because females appear to have

a tendency to deposit new egg masses adjacent to existing ones or both. Masses are

deposited unattached (unsupported by a vegetation brace; Dickerson 1906), often in water


so shallow that only the lower half of each egg mass is submerged, the upper portion beingexposed directly to the air (Licht 1969a). This pattern of oviposition makes mortality ofwestern spotted frog embryos resulting from desiccation or freezing relatively frequent; upto 30% embryonic mortality attributable to desiccation or freezing is not unusual (Licht1974). Oviposition sites may be reused in successive years, indicating that the sites mayhave unique characteristics (Licht 1971a), that they may be limiting, that adults have limitedflexibility to switch sites, or a combination thereof. Whatever the combination, most willmake the western spotted frog particularly vulnerable to oviposition-site modification. Egg

masses themselves are globular and contain several hundred to several thousand moderate-

sized (average diameter = 2.3 mm) eggs. The Cascades frog lays an egg mass very similarto that of the western spotted frog (and perhaps other spotted frogs), so at localities where

the two occur together, one cannot allocate egg masses to a species without additional data.If the oviposition site is a temporary aquatic site (e.g., see Licht 1969a), frogs may move tonearby permanent water before the aquatic site dries (pers. observ.). In British Columbia,larvae can hatch in ca. 5-10 days, require ca. 5-7 months to develop to metamorphosis, andafter metamorphosis, can reach sexual maturity in two (males) to three (females) years

(Licht 197 la, 1974). Based on observing frogs of four or five different sizes at springemergence, Dickerson (1906) thought that four or five years were needed to reach maturity.Because the frogs upon which Dickerson based her conclusion were from the Seattle area,such a long interval to reach maturity is unlikely because western spotted frogs from BritishColumbia at higher latitude grew almost twice as fast. Data on the developmental schedulein Oregon are lacking, but it is anticipated to be somewhat faster at the lower latitude, given

a roughly equivalent elevation, than what Licht observed in British Columbia.Active Season Habitat Requirements. Postmetamorphic stages of the western spotted

frog seem to be daytime active (Dickerson 1906; Licht 1969a, 197 la, 1974; pers. observ.),but observations of spotted frogs made at night (22:00-22:30 hr; R. Storm, pers. comm.)

early in the season and during the summer suggests that frogs may remain active because

warmwater conditions are maintained into the night. Indeed, observations made at.GoldLake Bog, Conboy National Wildlife Refuge, Penn Lake, and the Wood River over thepast two years strongly suggest that postmetamorphic western spotted frogs are somehowtied to warmer water (200-350C) during the late spring and summer season when frogs areactive (pers. observ.; J. Applegarth, D. Green, pers. comm.; see also Licht 1974). If thisinterpretation is correct, it may be the habitat requirement that ties the western spotted frogto warmwater marsh habitats. As noted earlier, the western spotted frog is not recorded


either along the coast of Oregon or Washington (see McAllister and Leonard 1990, 1991)

or at higher elevations in western Oregon (see Figure 1 and discussion in the Systematic

Status of the Spotted Frog in Western Oregon section). This distribution may be related

either to the lack or the reduced availability (time intervals too short) of warmwater habitats

in those areas. Notably, the single feature that united all verifiable spotted frog localities in

western Oregon for which habitat data could be retrieved was that each site had a marsh or

bog. Moreover, these marshes frequently represented overflow areas of a nearby river or

stream (see Comment section of localities in Appendix III). The similarity among these

sites and previously published ones (see Dickerson 1906, Licht 1971 a) suggests that the

requirements of embryonic stages (oviposition sites) and those of the postmetamorphic life

stages (warmwater refuge or foraging sites) may be the constraints that link the western

spotted frog to such warmwater habitats. Use of warmwater habitats by postmetamophic

western spotted frogs would probably make this species significantly more vulnerable to

potential predation by warmwater-loving exotic species (e.g., bullfrogs, Louisiana swamp

crayfish (Procambarus clarkii), and various catfishes (Ameiurus sp., Ictalurus sp.) and

sunfishes (Lepomis sp.)). This possibility would provide a basis for testing the idea that

warmwater exotics are responsible for its rapid historical demise. The impact of exotic

aquatic predators on the larvae of western spotted frogs is not known, but it is suspected of

being negative (see Hayes and Jennings 1986).

Historical Change

Lack of records for the spotted frog in southwestern Oregon, in particular, in the

Umpqua River drainage and most of the Rogue River drainage (even the two records for

the Rogue system are questionable [see 2 and 5 in Appendix IV]) may reflect a historical

absence of the spotted frog from this region. Even if this is not the case, the few records

do not permit addressing historical patterns in any sensible way. Therefore, discussion of

historical changes in the distribution of the spotted frog in western Oregon focuses on the

Willamette and Klamath hydrographic basins, each treated separately. This discussion is

based on 105 specimens from 23 different localities plus a few other data from the literature

and the recollections of selected individuals who have had field experience with this species

in western Oregon.

In the Willamette drainage basin, a near-exhaustive search of historical collections

revealed relatively few specimens (52 from 15 different localities) available to reconstruct


the historical distribution of the western spotted frog (see 1-4 and 13-23 in Figure 1 and

Appendix III). Thirty-three specimens from at least 10 different localities indicate that

spotted frogs were reasonably broadly distributed over roughly the eastern two-thirds of

the lowland (< 600 ft [= 180 m]) Willamette Valley at least into the 1930s. The earliest

comment attributable to the western spotted frog is probably that of Dickerson (1906), who

indicated that, "...in Oregon and Washington, it[northern red-legged frog] is abundant, but

is less so in many parts of these states than is Rana pretiosa." To placed this statement in

the proper context, the spotted frog that Dickerson discusses was included in what is now

recognized as the Cascades frog because Slater did not describe the latter until 1939. On

the other hand, in 1906, what is now recognized as the Cascades frog had not yet been

collected from either Oregon or Washington because most of its habitat (higher elevations)

was inaccessible to the collectors at that time. In contrast, collection was frequent in

lowlands around Puget Sound, from where Dickerson's correspondent in Seattle, Dr. J. L.

Illingworth, gathered most specimens he sent to her. Because the western spotted frog is

mostly a lowland form (especially in Washington) when contrasted with the Cascades frog,

Dickerson's statement likely refers to the western spotted frog. A somewhat similar

statement data was made by Stanley G. Jewett, Jr., in the summary of he wrote about the

amphibians around Portland, Oregon (Jewett 1936). He described the spotted frog as,

"common along the sloughs of the Willamette and Columbia rivers." Jewett's summary

was also published prior to the description of the Cascades frog. Nevertheless, the

comments that Jewett made cannot refer to anything but what is now regarded as the

western spotted frog because the only two species of ranid frogs Jewett ever collected in

the Portland area were the western spotted frog (see 22 in Appendix III) and the northern

red-legged frog; the catalogue that Jewett kept indicates that he consistently distinguished

the two. Moreover, Jewett sent both some of the western spotted frogs and northern red-

legged frogs he collected in the Portland area to Tracy I. Storerk, who verified

identifications for him using the frog names applied at that time (Jewett 1936), and no

Cascades frogs have ever been recorded from the Portland area. Commenting about the

late 1930s, Graf (1939) stated that the [western] spotted frog was locally common in the

upper portion of the lowland Willamette Valley [above Salem]. Graf, now deceased,

further related to Robert Storm (pers. comm.) that spotted frogs were locally abundant in

k Storer was a reknown amphibian biologist then at the University of California at Davis (UCD). Anumber of Jewett amphibian collections still exist in the UCD collection (e.g., see UCD numbers underlocality #21 in Appendix III).


the Willamette Valley into the 1940s. It may be significant that no post-1940 collections of

western spotted frogs exist for the lower portion of the lowland Willamette Valley [below

Salem]; the only pre- 1940 collections known from the lower portion of the lowlandWillamette are a 1937 record from Aumsville and a handful of records from the Portland

area collected between 1905 and 1931 (see 21-23 in Appendix III). Based on his

discussions with Kenneth Gordon and William Graf, Robert M. Storm (pers. comm.)

indicated that little attention was paid to spotted frogs in the interval during which they

apparently disappeared from the lowland Willamette Valley, and that it was only in

retrospect that Storm realized that the few spotted frogs that he saw in the 1950s in the

upper Willamette probably represented the tail of this disappearance. The last spotted frog

that Storm recollected observing in the lowland Willamette Valley was the lone specimen he

collected on 25 February 1957 roughly 11 km [7 mil east of Corvallis (18 in Appendix III).

The last verifiable records of western spotted frogs from the lowland Willamette Valley are

the specimens collected at Sweet Home in 1966 (18 in Appendix III). Except for the single

recent record from Dempsey Creek in Thurston County (Washington; K. McAllister, W.

Leonard, pers. comm.), this dates is close to that of the last record of a western spotted

frog from western Washington (Brush Prairie near Vancouver in 1968 (McAllister and

Leonard 1991)).

No systematic surveys of historical localities of the western spotted frog in the lowland

Willamette Valley have been conducted since the disappearance interval in the 1950s and

1960s suggested by the aforementioned data. Yet, a reasonable probability exists that the

western spotted frog is extinct in the lowland Willamette Valley. This contention has

support in that no verifiable records (photographs or specimens) of spotted frogs have

emerged since 1966, even though a number of herpetologists familiar with spotted frogs

have intermittently worked scattered locations in the lowland Willamette over the past 25

years. Most significant among the latter are the collections Ronald A. Nussbaum made

over the interval 1968-1974 (now housed at the University of Michigan), which include

more than 10,000 amphibian and reptile specimens from western Oregonl. Nussbaum,

widely regarded in herpetological circles as one of the most, if not the most, devastating

collectors ever, did not turn up a single spotted frog in the lowland Willamette over that

interval. The decline in observability of the western spotted frog in the lowland Willamette

I The Oregon State University collection (OSUMNH), for many years under the care of Robert M. Storm,is larger (over 12,000 specimens), but fewer than 10,000 of these specimens come from western Oregonand these were not collected either over as short an interval as the Nussbaum collection nor were mostcollected over the interval after 1966.

ou H~sNorib 63 ~ r/SOUVEKRN) 'a le

Haves: Si)otted Froe - 23Haves: Srotted Fro2 23 ---n __

SOUTHERN ORCUpit,5 l~~QSr-1Ba ASHLAND, NiL

Valley is suggested by the number of specimens collected over the two 25-year intervals,

1941-1966 and 1967-1992, when compared to specimens collected before 1941. Notably,

both the number of specimens collected (Figure 5a) and the number of specimens collected

per locality (Figure Sb) halved from the pre-1941 to the 1941-1966 interval. These data

could be interpreted as reflecting a reduction in collection effort over each interval. This is

probably not the case because if one compares the collections of northern red-legged frogs

from the University of Michigan collected over the same intervals (Figure 6a), that effort

probably increased over the 1967-1992 intervaln. Notably, the ability to collect northern

red-legged frogs based on the numbers collected per locality reflects little change (Figure

6b). This contrasts sharply with the same data for the western spotted frog in the lowland

Willamette Valley (Figure Sb). The limitations of this analysis notwithstanding, the best-

founded interpretation of available data for the lowland Willamette is that an identifiable

reduction in the distribution and numbers of the western spotted frog occurred during the

interval 1941-1966; the data since 1966 suggest that spotted frogs have become extinct in

the lowland Willamette Valley.

Because the decline of spotted frogs in the lowland Willamette was not systematically

traced, one can only speculate on the actual changes that were causal. Yet, examination of

changes that occurred at the localities from which spotted frogs were historically found

through comparison of the differences in conditions before and after they were recorded

there provides useful insights. All changes associated with historical spotted frog localities

can be grouped into two larger categories: 1) changes associated with the vegetation or the

physical structure of spotted frog habitat, and 2) primarily non-structural changes resulting

from novel introductions of an exotic aquatic fauna. With regard to the former, it is

essential to recall that available data suggest that the western spotted frog linked to

warmwater habitats. Changes associated with the two Portland area localities from which

western spotted frogs were recorded (Crystal Springs Lake and Johnson Creek; 22 and 23

in Appendix III) seem to differ from those at the remaining eight western spotted frog

localities in the lowland Willamette Valley, so these are discussed separately first. Both the

Crystral Springs Lake and Johnson Creek areas underwent terrestrial developmental

m These data are slightly biased in that the pre-1941 interval is actually over 25 years in length because thefirst spotted frog specimens collected in the lowland Willamette Valley date from 1905. However, if onecorrects the data to make the pre-1941 interval exactly 25 years (1915-1940), only two specimenscollected between 1905 and 1914 would be removed from the existing number collected before 1941.

n The alternative argument is that the numbers of northern red-legged frogs increased over this interval, butthis contradicts available observations which suggest that they have either declined or disappeared frommany localities where they occurred historically (see Hayes and Jennings 1986).

a

60

50*, Lowland Willamette

rl Cascades slone (Willamette)40--

Number * Klamath Basinof

Specimens 3Collected

20

10

0-1940 1941-1966 1967-1992

Interval

b

6 * Lowland Willamnette

Average [3 Cascade slop~e (Willareteu)Number * Klamath Basin

Of

per

0

-1940 1941-1966 1967-1992

Interval

Figure 5 - Data on collections of western spotted frogs from western Oregon.

a

80

60 -

40 -Number

ofSpecimens

20 -

0 .1.

-1940 1941-1966 1967-1992

Interval

b8

AverageNumber

OfSpecimens

perLocality

4

2 -

0 4--1940 1941- 1966

Interval

1967-1992

Figure 6 - Data on collections of northern red-legged frogs from western Oregon.


changes that eliminated the marsh habitat upon which the western spotted frogs appear to

be dependent, the former as the result of an expansion of the Eastmoreland Muncipal Golf

Course in the 1930s and the latter due to the channelization and development along lower

Johnson Creek in the interval between 1925 and 1940. The alternative that the exotic

aquatic fauna (primarily bullfrogs and warmwater fishes) contributed to the decline of

spotted frogs at these localities cannot be excluded. This fauna was introduced to western

Oregon at early dates, but it is unlikely to have been as significant as the aforementioned

habitat changes because this exotic fauna did not become really well-established in the

Crystal Springs-Johnson Creek system until the late 1940s (R. Macey, pers. comm.). The

remaining eight spotted frog localities in the lowland Willamette Valley have a high

probability of having been significantly affected by both the historical modification of

flooding regimes and the exotic aquatic fauna. Up to the 1940s, the Willamette River and

its tributaries were largely unrestricted and underwent frequent winter flooding, which

maintained numerous shallow-water areas on the valley floor. As of 1941, the US Army

Corps of Engineers began to build a series of flood control structures on the major arms of

the upper Willamette system such that by the late 1960s, the 13 larger reservoirs seen today

were in place (US Army Corps of Engineers 1982, Johnson et al. 1985). As was their

function, these reservoirs markedly reduced the extent of flooding in the valley (US Army

Corps of Engineers 1982), and significant areas of shallow warmwater marsh habitat,

largely dependent on such flooding, were eliminated during this period. Because western

spotted frogs in the lowland Willamette Valley may have been almost exclusively associated

with warmwater marsh habitat, this change would have contributed significantly to

reducing their populations. However, the negative effects that elimination of warmwater

marsh habitat in the lowland Willamette Valley may have had on western spotted frogs is at

least partly confounded with any negative effects that may have been caused by the exotic

aquatic fauna. Bullfrogs, introduced to the Willamette Valley in 1921 for sport and later,

the culinary market for their legs; Louisiana swamp crayfish, introduced as forage for

bullfrogs; and a suite of warmwater fishes, introduced repeatedly by acclimatization

societies and other individuals before and after the turn of the century, were well-

established over much of the lowland Willamette Valley by 1940 due to the fish- and frog-

farming crazes of the previous decade (Lampman 1945). Failure, financial and otherwise,

of all bullfrog-farming and most fish-farming ventures resulted in the "farmers"


abandoning or releasing their stock, and forever changing the composition of the

warmwater aquatic fauna in most of the lowland Willametteo. The warmwater habitat

requirement of western spotted frogs, a feature in which they diverge significantly from the

other native ranid frogs in the Pacific Northwest, may be the factor that ensured a high

degree of contact with the novel warmwater predatorsP. Moreover, western spotted frog

larvae may be placed at a severe disadvantage where they co-occur with bullfrog larvae,

and exotic fishes and crayfishes are the selective agents (see Hayes and Jennings 1986 for a

pertinent discussion).

Nineteen specimens from at least five different localities indicate that spotted frogs are

recorded at scattered locations on the Cascades slope of the Willamette hydrographic basin.

Interpretation of changes in western spotted frog populations on the Cascades slope is more

difficult because records are fewer and only one pre- 1941 collection exists, three specimens

that Stanley G. Jewett, Jr. collected at Little Crater Meadows in 1930 (see 4 in Appendix

III). The temporal distribution of Cascade slope specimens (Figure Sa) supports the fact

that collectors made infrequent attempts to access the montane localities where spotted frogs

existed until after road access to montane areas improved in the 1950s and 1960s (R.

Storm, pers. comm.). Unlike the situation in the lowland Willamette Valley, efficiency of

collection does not appear to have changed (Figure Sb), although the number of specimens

collected in the last 25 years has declined (Figure Sa). Because all collections made since

1967 are from a single locality, Gold Lake Bog (see 13 in Appendix III), and few Cascades

slope collections of the western spotted frog exist, it was not possible to interpret a pattern

with confidence. Just the fact that four Cascades slope localities require verification (see

Figure 1 and Appendix IV) is reflective of the paucity of reliable data on Cascades slope

localities for the spotted frog. The most recent collections of the spotted frog in western

Oregon are from the western Cascades slope. Thus, if the later a collection was made is

reflective of the likelihood that spotted frogs are extant, then the region with the greatest

o In addition, numerous WPA fish "rescue" programs in the Willamette Valley in the 1930s and 1940stransplanted and liberated many thousands of warmwater fishes.

P The warmwater requirements of bullfrogs are well known; their eggs die at water temperatures below 15C(Moore 1942), and adult males rarely give advertisement calls at water temperatures below 21TC (Bury andWhelan 1982). The warmwater requirements of the exotic fishes are similar: bluegill (Lepomismacrochirus) typically will not spawn without water temperatures at least in the 18'-21TC range; forblack crappie (Pomoxis nigromaculatus), the minimum end of the range is 14'-17'C; for largemouth bass(Micropterus salmoides), it is 14O- 16TC; and for black bullhead (Ictalurus melas), it is ca. 20TC (Moyle1973). The range of temperatures required by all these species overlaps significantly with the temperaturerequirements for the spotted frog (see Habitat Requirements and Natural History).


probability of still having western spotted frog populations is the slope of the westernCascades.

The Cascades slope localities from which western spotted frogs are recorded also differsignificantly from parallel locations in the lowland Willamette Valley floor with regard tothe factors that may impact western spotted frog populations. In particular, the exoticwarmwater fauna (bullfrogs, exotic crayfishes, and warmwater fishes) with the potential toseverely negatively impact the western spotted frog is probably absent; the only exoticspresent at any of the montane localities from which the western spotted frog has been

recorded are brook trout (Salvelinusfontinalis) and rainbow trout (Oncorhynchus mykiss),

and the patterns of habitat utilization of these salmonids provides little opportunity for them

to co-occur with western spotted frog life stages. Second, the potential for severe alteration

because of urban or suburban development or hydrological modifications is reduced orabsent. Cascades slope localities are on federal land where such development is generallyexcluded. All are also associated with the portions of drainages upstream from majorhydrological developments where opportunities to significantly modify hydrology arereduced. Examination of historical aerial photographs suggests that the impacts likely tohave most severely affected Cascades slope populations of the spotted frog were cattle and

sheep grazing that visibly altered vegetation structure and perhaps water quality at somemontane marshes from which western spotted frogs were historically recorded. Gold Lake

Bog and Goose Lake are exceptions in this regard in that both localities have historically

had little or no grazing; Gold Lake Bog has benefited from protection as either a Natural

Area or Research Natural Area since 1965 (Kezer 1978; W. Dugas, pers. comm.), andGoose Lake has benefited from its inclusion within the Three Sisters Primitive Area since1937 and its reclassification as the Three Sisters Wilderness Area since 1957. Of the

verified historical localities on the Cascades slope, Gold Lake Bog and Goose Lake likelyhave the greatest probability of still harboring spotted frogs. Juvenile spotted frogs have

been observed near the Salt Creek outflow of Gold Lake, the opposite end of the lake from

which the historical locality (the bog) is situated, as recently as September of 1991 (pers.

observ.).

Fifty-three specimens from eight localities indicate that the western spotted frog is

recorded at scattered localities across the Klamath Basin (Figure 1). Only one recent recordfor the western spotted frog exists for the Klamath Basin has (Figure 5a), two specimens

collected from the upstream end of Little Hyatt Reservoir in 1971 (see 5 in Appendix III).Beyond that isolated record, the most recent records are two 1934 collections made along


the Sprague River (see 11 and 12 in Appendix III). Interpretation of what this means is not

possible because much of the upper Klamath Basin has been infrequently and irregularly

collected. Systematic surveys of both historical localities and localities identified as having

suitable habitat for spotted frogs are necessary to evaluate the current status of the western

spotted frog in the Klamath Basin. Whatever its current status may be, available collections

suggest that historically, the spotted frog was common in the lower Klamath Basin (Figure

5a). More specimens were collected at each locality in the Klamath Basin than anywhere in

the Willamette hydrographic basin (Figure 4b), which likely reflects the fact that the

Klamath Basin historical harbored more shallow, warmwater marshland, the habitat likely

most suited to the western spotted frog, than in any other area in the state. It is outside the

scope of this report to detail the pattern of historical changes in the Klamath Basin, but

significant changes, many similar to those that have taken place in the Willamette Valley,

have occurred in the Klamath Basin (e.g., Sorenson and Schwarzbach 1991). Some of

these changes have undoubtedly impacted the western spotted frog.

Summary and Recommendations

The spotted frog (Rana pretiosa), one of four native ranid frogs found in western

Oregon, likely represents several taxa; one of the putative taxa, termed the western spottedfrog, occurs in western Oregon. This taxon is distinguishable from other spotted frogs onthe basis of a characteristic mottling over much of the surface of the abdomen. In western

Oregon, the western spotted frog can be distinguished from the species with which it maybe most easily confused, the Cascades frog (R. cascadae) and the northern red-legged frog(R. a. aurora) on the basis of its relative shorter legs; nearly full webbing of its feet;distinctly upward-directed eyes; inky black, ragged-edged spots that frequently have light

centers; lack of mottling in the groin; a relatively granular-textured skin; dark belly andchest mottling; and probably red/red-orange pigment on selected undersurfaces.

The most significant finding of this analysis is that, in contrast to the other native ranidfrogs in the Pacific Northwest, which display varying requirements for aquatic habitats thattend to have colder water, the western spotted frog may be closely linked to aquatic habitatsthat have relatively warm water (2 20'C), at least during the summer season. This apparent

specialization may explain the fact that it seems to be recorded only from marsh habitats inthe Willamette and Klamath Basins below 5,000 feet, and that it is unrecorded from coastal

areas where average water temperatures are cooler. This apparent specialization may also


explain its probable vulnerability to exotic aquatic warmwater predators because overlap inthe patterns of habitat utilization between the western spotted frog and this fauna is likely tobe greater than the overlap between any other native ranid frog in the Pacific Northwest andthe same fauna. Scattered indications exist that the loss of marsh habitats or their physicalor biotic modification, particularly as a result of hydrological changes, may also be directlylinked to the apparent decline of the western spotted frog in western Oregon.

The conclusions drawn from these data led to the following recommendations:1) Verification of the apparent decline of the western spotted frog in western Oregon

requires that historical localities be re-examined to determine whether the western spottedfrog still exists at localities from which it was historically recorded. Despite numerousindications that the western spotted frog has declined, its current status in western Oregoncannot be reevaluated in the absence of such an examination. In this re-examination, thelocalities which are verifiable (Appendix III) should have highest priority; unverifiedrecords linked with especially suitable habitat should have second priority (part ofAppendix IV); and other unverified records (part of Appendix IV) and localities withsuitable habitat, but unlinked to unverified records, should have the lowest priority. Thisre-examination should not simply attempt to determine whether spotted frogs are present, itshould also evaluate site-specific conditions that may help explain why spotted frogs arepresent or absent at each locality. Data sufficient to evaluate the spotted frog for listingstatus by the State in this region will be available only after such an examination is made.

2) The pattern of apparent warmwater habitat utilization by the western spotted frogrevealed in this evaluation, a pivotal feature explanatory of numerous aspects of itsdistribution and biology, needs further verification. Examination of temperature variationin western spotted frogs and of the waters in which they occur should be compared totemperature variation in waters where they are absent. These data are necessary to confirmhow tightly linked western spotted frogs are to warmwater habitats, which could greatlyincrease predictive power in the assessment of western spotted frog habitat.

3) Evaluation of the current and historical status of the western form in eastern Oregonin a manner parallel to that done for western Oregon is needed to a evaluate the status of thewestern spotted forg on a statewide basis.

4) Evaluation of the current and historical status of the yellow-ventered form of thespotted frog eastern Oregon and the potentially different red-orange venter spotted frog insoutheastern Oregon in a manner parallel to that done for the western spotted frog is alsoneeded. The partitioning of the spotted frog into several taxa has resulted in reduced ranges


for each taxon, making it more critical to understand the status of each taxon. Indicationsexist that the yellow-ventered form of the spotted frog found in eastern Oregon may have

declined significantly as the result of drought and hydrologic alterations (J. Kirk, A. St.John, pers. comm.). These suppositions need verification.

Acknowledgments

Numerous individuals contributed to my ability to refine this report. Without theirassistance, its completion would not have been possible. Those in charge of collections,

their associates, students or assistants who provided information included: Mark Jenningsand Jens Vindum (CAS, CAS-SU); Clarence McCoy (CM); William Duellman and JohnSimmons (KU); Barbara Stein and David Wake (MVZ); Mark Stern (ONHD); DouglasMarkle (OSUMNH); Brad Shaffer (UCD); Russell Burke, Sheng Hai, and Arnold Kluge(UMMZ); Ernest Karlstrom and Dennis Paulson (UPS); Ron Crombie, Roy McDiarmid,and Robert Reynolds (USNM). John Applegarth, Joseph Beatty, Ellen Benedict, DelbertBlackburn, Andrew Blaustein, Stephen Cross, Richard Forbes, Rebecca Goggans, David

Green, Stephen Hoiland, Kirk Horn, Doris Jewett, James Kezer, James Kirk, FlorenceLehman, David Marshall, James Merzenich, Jane Mesereau, Kelly McAllister, Clea

Neidert, Deanna Olsen, Edward Park, Robert Penson, Claire Puchy, Alan St. John, James

Sjulin, Robert Storm, and Eric Terdahl all contributed data or insights, or assisted infinding bits of crucial information. Jill Mellen provided emotional support and assisted in

editing and proofing the manuscript. The Metro Washington Park Zoo allowed use of their

computer facilities for laser printing.


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Turner, Frederick B. 1959a. Variation in skeletal proportions of Rana p. pretiosa inYellowstone Park, Wyoming. Copeia 1959(1):63-68.

Turner, Frederick B. 1959b. Pigmentation of the western spotted frog, Rana pretiosapretiosa, in Yellowstone Park, Wyoming. American Midland Naturalist 61(l):162-176.

Turner, Frederick B. 1959c. An analysis of the feeding habits of Rana p. pretiosa inYellowstone Park. American Midland Naturalist 61(2):403-413.


Turner, Frederick B. 1960a. Population structure and dynamics of the western spottedfrog, Rana pretiosa pretiosa Baird and Girard, in Yellowstone Park, Wyoming.Ecological Monographs 30(3):251-278.

Turner, Frederick B. 1960b. Tests of randomness in recaptures of Rana pretiosa pretiosa.Ecology 41(1):237-239.

Turner, Frederick B. 1960c. Postmetamorphic growth in anurans. American MidlandNaturalist 64(2):327-338.

Turner, Frederick B. 1962a. The demography of frogs and toads. Quarterly Review ofBiology 37(4):303-314.

Turner, Frederick B. 1962b. An analysis of geographic variation and distribution of Ranapretiosa. American Philosophical Society Yearbook 1962:325-328.

Turner, Frederick B., and Philip C. Dumas. 1972. Rana pretiosa. Catalogue of AmericanAmphibians and Reptiles: 119.1-119.4.

Underhill, J. E. 1967. Report of a trial program of interpretation, Kokanee Creek park -Aug. 1967. Unpublished Report, British Columbia Parks Branch, Victoria. 9 pp.

Uzzell, Thomas, and Thomas J. Post. 1986. Rana temporaria is not a member of the Ranaboylii group. Systematic Zoology 35(3):414-421.

Van Denburgh, John, and Joseph R. Slevin. 1915. A list of the amphibians and reptilesof Utah, with notes on the species in the collection of the Academy. Proceedings ofthe California Academy of Sciences, Series 4, 5(4):99-1 10.

Van Denburgh, John, and Joseph R. Slevin. 1921. List of the amphibians and reptiles ofIdaho, with notes on the species in the collection of the Academy. Proceedings ofthe California Academy of Sciences, Series 4, 11(3):39-47.

Wade, Keith. 1960. Report on the reptiles and amphibians of Wells Gray Park - May andJune, 1960. Unpublished Report, British Columbia Parks Branch, Victoria. 7 pp.

Waits, J. A. 1962. Parasitic helminths as aids in studying the distribution of species ofRana in Idaho. Transactions of the Illinois Academy of Sciences 54:152-156.

Waldman, Bruce, and Michael J. Ryan. 1983. Thermal advantages of communal eggmass deposition in wood frogs (Rana sylvatica). Journal of Herpetology 17(1):70-72.

Wallace, D. G., M. C. King, and A. C. Wilson. 1973. Albumin differences among ranidfrogs: taxonomic and phylogeneic implications. Systematic Zoology 22(1):1-13.

Wells, Kentwood D. 1977. The social behaviour of anuran amphibians. Animal Behavior25(3):666-693.

Whitaker, John O., Jr., Stephen P. Cross, John M. Skovlin, and Chris Maser. 1983.Food habits of the spotted frog (Rana pretiosa) from managed sites in GrantCounty, Oregon. Northwest Science 57(2):147-154.


Wishard, L. 1977. Larval growth in Rana pretiosa: Ecological and genetic factors. MSthesis, University of Montana, Missoula, Montana.

Worthing, Patricia. 1993. Endangered and Threatened wildlife and plants; Finding onPetition to list the spotted frog. Federal Register 58(87):27260-27263.

Wright, Albert H., and Anna A. Wright. 1933. Handbook of frogs and toads - the frogsand toads of the United States and Canada. Comstock Publishing Company, Inc.,Ithaca, New York. 231 pp.

Wright, Albert H., and Anna A. Wright. 1949. Handbook of frogs and toads (3rdedition). Comstock Publishing Associates, Ithaca, New York. 640 pp.

Wright, Richard T. 1972. Wells Gray Provincial Park Naturalist's Report 1972.Unpublished Report, British Columbia Parks Branch, Victoria. 55 pp.

Yarrow, Henry C. 1875. Report upon the collection of batrachians and reptiles made inportions of Nevada, Utah, California, Colorado, New Mexico, and Arizona, duringthe years 1871, 1872, 1873, 1874. Wheeler Survey 5:509-633.

Yarrow, Henry C. 1882. Check list of North American reptilia and batrachia withcatalogue of specimens in U.S. National Museum. United States National MuseumBulletin (24): 1-249.

Zweifel, Richard G. 1954. A new Rana from the Pliocene of California. Copeia1954(2):85-87.


Appendix II

Chronological Bibliography

1853

Baird, Spencer F., and Charles Girard. 1853. [Communication regarding Rana pretiosaand Bufo columbiensisl. Proceedings of the Academy of Natural Sciences ofPhiladelphia 6(10):378-379.

1875

Cope, Edward D. 1875. Check-list of North America Batrachia and Reptilia with asystematic list of the higher groups, and an essay on geographical distribution.Based on the specimens contained in the U.S. National Museum. United StatesNational Museum Bulletin 1(1): 1-104.

Yarrow, Henry C. 1875. Report upon the collection of batrachians and reptiles made inportions of Nevada, Utah, California, Colorado, New Mexico, and Arizona, duringthe years 1871, 1872, 1873, 1874. Wheeler Survey 5:509-633.

1879

Cope, Edward D. 1879. A contribution to the zoology of Montana. American Naturalist13(7):432-441.

1882

Boulenger, George A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in thecollection of the British Museum. 2nd edition, printed by order of the trustees of theBritish Museum of Natural History. 503pp.

Yarrow, Henry C. 1882. Check list of North American reptilia and batrachia withcatalogue of specimens in U.S. National Museum. United States National MuseumBulletin (24):1-249.

1889

Cope, Edward D. 1889. The Batrachia of North America. United States NationalMuseum Bulletin 34:1-525.

1891

Boulenger, George A. 1891. Notes on American Batrachians. Annals and Magazine ofNatural History (Series 6) VIII(48):453-457.

Test, Frederick C. 1891 [1892]. Fish-cultural investigations in Montana and Wyoming.Annotated list of reptiles and batrachians collected. Bulletin of the United StatesFish Commission for 1891:57-59.


1893

Cope, Edward D. 1893. A contribution to the herpetology of British Columbia.Proceedings of the Academy of Natural Sciences of Philadelphia 44(Part 1):181-184.

1896

Cope, Edward D. 1896. Synonymic list of the North American species of Bufo and Rana,with descriptions of some new species of Batrachia, from specimens in the NationalMuseum. Proceedings of the American Philosophical Society 23(124):514-526.

1897

Chamberlain, Frederick M. 1897. Notes on the edible frogs of the United States and theirartificial propagation. pp. 249-261. In: John J. Brice (director), A manual of fishculture based on the methods of the United States Commission of Fish andFisheries. US Government Printing Office, Washington, D.C.

1898

Fannin, John. 1898. A preliminary catalogue of the collections of natural history andethnology in the Provincial Museum, Victoria, British Columbia. British ColumbiaProvincial Museum, Victoria. 196 pp.

1899

Meek, S. E., and D. G. Elliot. 1899. Notes on a collection of cold-blooded vertebratesfrom the Olympic Mountains. Publications of the Field Columbian Museum (31),Zoological Series 1(12):225-236.

1906

Dickerson, Mary C. 1906. The frog book. Doubleday, Page and Company, Garden City,New York. 253pp.

1912

Hollister, N. 1912. List of reptiles and batrachians of the Alpine Club expedition to theMount Robson region. Canadian Alpine Journal 4(special number):45-46.

1913

Thompson, Helen B. 1913. Description of a new subspecies of Rana pretiosa fromNevada. Proceedings of the Biological Society of Washington 26:53-56.

1915

Van Denburgh, John, and Joseph R. Slevin. 1915. A list of the amphibians and reptilesof Utah, with notes on the species in the collection of the Academy. Proceedings ofthe California Academy of Sciences, Series 4, 5(4):99-1 10.

Hayes: Spotted Frog 3 Hayes:

1916

Dice, Raymond L. 1916. Distribution of the land vertebrates of southeastern Washington. Hardy,University of California Publications in Zoology 16(17):293-348.

1917Kermo

Camp, Charles L. 1917. Notes on the systematic status of the frogs and toads ofCalifornia. University of California Publications in Zoology 17:115-125.

TannexCary, M. 1917. Life zone investigations in Wyoming. North American Fauna (42):1-95.

Grinnell, Joseph, and Charles L. Camp. 1917. A distributional list of amphibians and Tanneireptiles of California. University of California Publications in Zoology 17(10): 127-208.

1919Slevin

Boulenger, George A. 1919. Synopsis of American species of Rana. Annals andMagazine of Natural History (Series 9) 111(16):408-416.

1920Camp

Boulenger, George A. 1920. Monograph of the American frogs of the genus Rana.Proceedings of the American Academy of Arts and Sciences 55(9):413-480.

Patch,1921

Blanchard, Frank N. 1921. A collection of amphibians and reptiles from northeasternWashington. Copeia (90):5-6.

Tann(Van Denburgh, John, and Joseph R. Slevin. 1921. List of the amphibians and reptiles of

Idaho, with notes on the species in the collection of the Academy. Proceedings ofthe California Academy of Sciences, Series 4, 11(3):39-47.

1922 Brue!

Patch, Clyde L. 1922. Some amphibians and reptiles from British Columbia. Copeia(111):74-79. Logic

1923

Stejneger, Leonhard, and Thomas Barbour. 1923. A check list of North Americanamphibians and reptiles, 2nd edition. Harvard University Press, Cambridge, BurkMassachusetts. 17lpp.

Stejr1925

Storer, Tracy I. 1925. A synopsis of the amphibia of California. University of CaliforniaPublications in Zoology 27:1-342. Svih

Hardy, George A. 1925. Amphibia of British Columbia. Annual Report of the ProvincialMuseum of Natural History, Victoria for the year 1924:21-24.

3 Hayes: Spotted Frog 4

1927

ngton. Hardy, George A. 1927. Report on a collecting trip to Garibaldi Park, B.C. AnnualReport of the Provincial Museum of Natural History, Victoria for the year 1926:15-26.

Kermode, Francis. 1927. Accessions. Annual Report of the Provincial Museum ofNatural History, Victoria for the Year 1926:38-39.

Tanner, Vasco M. 1927a. Distributional list of the amphibians and reptiles of Utah, No.:1-95. 1. Copeia (163):54-58.

id Tanner, Vasco M. 1927b. An ecological study of Utah amphibia. Proceedings of thel): 127- Utah Academy of Sciences, Arts, and Letters 4:6-7.

1928

Slevin, Joseph R. 1928. The amphibians of western North America. California Academyof Sciences, Occasional Paper (16):1-152.

1929

Campbell, Berry. 1929. Notes on three amphibians. Crater Lake National Park NatureNotes 2(2):8.

Patch, Clyde L. 1929. Some amphibians of western North America. Canadian FieldNaturalist 43(6):138.

1931

Tanner, Vasco M. 1931. A synoptical study of Utah amphibians. Proceedings of the-Is of Utah Academy of Sciences, Arts, and Letters 8:159-198.S of

1932

Brues, C. T. 1932. Further studies on the fauna of North American hot springs.Proceedings of the American Academy of Arts and Sciences 67:185-303.

Logier, E. B. S. 1932. Some account of the amphibians and reptiles of British Columbia.Transactions of the Royal Canadian Institute, XVIII Part 2(53):311-336.

1933

Burke, V. 1933. Bacteria as food for vertebrates. Science 78(2018):194-195.

Stejneger, Leonhard, and Thomas Barbour. 1933. A check list of North Americanamphibians and reptiles, 3rd edition. Harvard University Press, Cambridge,Massachusetts.

Svihla, Arthur, and Ruth D. Svihla. 1933. Amphibians and reptiles of Whitman County,Washington. Copeia 1933(3):125-128.


1933 (continued)

Wright, Albert H., and Anna A. Wright. 1933. Handbook of frogs and toads - the frogsand toads of the United States and Canada. Comstock Publishing Company, Inc.,Ithaca, New York. 231pp.

1934

Myhrman, Herman M. 1934. The nature and extent of the Bidder's organ in Bufo boreasand allied animals. MA thesis, University of Puget Sound, Department of Biology,Tacoma. 54pp.

Slater, James R. 1934a. Notes on northwestern amphibians. Copeia 1934(3):140-141.

Slater, James R. 1934b. Ambystoma tigrinum in the state of Washington. Copeia1934(4): 189-190.

1935

Liu, Ch'eng C. 1935. Types of vocal sac in the Salientia. Proceedings of the BostonSociety of Natural History 41(3):19-40.

Svihla, Arthur. 1935. Notes on the western spotted frog, Rana pretiosa pretiosa. Copeia1935(3): 119-122.

1936

Fitch, Henry S. 1936. Amphibians and reptiles of the Rogue River basin, Oregon.American Midland Naturalist 17(3):634-652.

Ingles, Lloyd G. 1936. Worm parasites of California amphibia. Transactions of theAmerican Microscopical Society 55(1):73-92.

Jewett, Stanley G., Jr. 1936. Notes on the amphibians of the Portland, Oregon, area.Copeia 1936(1):71-72.

Slater, James R., and C. Frank Brockman. 1936. Amphibians of Mt. Rainier NationalPark. Mt. Rainer National Park Nature Notes 14(4):113-138.

Swarth, Harry S. 1936. Origins of the fauna of the Sitkan District, Alaska. Proceedingsof the California Academy of Sciences, 4th Series, 23(3):59-78.

1937

Cowan, Ian M. 1937. A review of the reptiles and amphibians of British Columbia.Annual Report of the British Columbia Provincial Museum of Natural History,Victoria for the year 1936:16-25.

Holland, George P. 1937. Life history and distributional studies of Bell's paintedterrapin, Chrysemys picta belli (Gray). MA thesis, University of British Columbia,Vancouver, British Columbia. 146pp.

Olsen, 0. W. 1937. Description and life history of the trematode Haplometrana utahensissp. nov. (Plagiorchiidae) from Rana pretiosa. Journal of Parasitology 23(1): 13-28.


1938

Olsen, 0. W. 1938. Aplectana gigantica (Cosmocercidae), a new species of nematodefrom Rana pretiosa. Transactions of the American Microscopical Society57(2):200-203.

1939

Brown, Walter C., and James R. Slater. 1939. The amphibians and reptiles of the islandsof the state of Washington. Occasional Papers, Department of Biology, College ofPuget Sound (4):6-13.

Graf, William. 1939. The distribution and habitats of amphibians and reptiles in Lincoln,Benton, and Linn Counties. MS Thesis, Oregon State University, Corvallis. 93pp.

Graf, William, Stanley G. Jewett, Jr., and Kenneth L. Gordon. 1939. Records ofamphibians and reptiles from Oregon. Copeia 1939(2): 10 1-104.

Gordon, Kenneth M. 1939. The amphibia and reptilia of Oregon. Oregon State CollegeMonographs, Studies in Zoology (1):1-82.

Slater, James R. 1939. Description and life history of a new Rana from Washington.Herpetologica 1(6):145-149.

Stejneger, Leonhard, and Thomas Barbour. 1939. A check list of North Americanamphibians and reptiles, 4th edition. Harvard University Press, Cambridge,Massachusetts. 207pp.

1940

Linsdale, Jean M. 1940. Amphibians and reptiles in Nevada. Proceedings of theAmerican Academy of Arts and Sciences 73(8): 197-257.

Stanwell-Fletcher, John F., and Theodora C. Stanwell-Fletcher. 1940. Naturalists in thewilds of British Columbia. Scientific Monthly 50:1-44.

Tanner, Wilmer W. 1940. Notes on the herpetological specimens added to the BrighamYoung University vertebrate collection during 1939. Great Basin Naturalist 1(3-4):138-146.

1942

Booth, Ernest S. 1942. Field key to the amphibians and reptiles of the Northwest.Unpublished Report, Walla Walla College, College Place, Washington. 22pp.

Logier, E. B. S., and G. C. Toner. 1942. Amphibians and reptiles of Canada. CanadianField-Naturalist 56(2): 15-16.

Rodgers, Thomas L., and William L. Jellison. 1942. A collection of amphibians andreptiles from western Montana. Copeia 1942(1):10-13.


1943

Carl, Clifford G. 1943. The amphibians of British Columbia. British ColumbiaProvincial Museum Handbook (2): 1-63.

Carl, Clifford G., and George A. Hardy. 1943. Report on a collecting trip to the Lac LaHache area, British Columbia. Annual Report of the Provincial Museum of NaturalHistory and Anthropology, Victoria for the year 1942:25-49.

Evenden, Fred G., Jr. 1943. Notes on the amphibia of the Cascade mountains in Oregon.Copeia 1943(4):251-252.

Stanwell-Fletcher, John F., and Theodora C. Stanwell-Fletcher. 1943. Some accounts ofthe flora and fauna of the Driftwood Valley region of north central BritishColumbia. British Columbia Provincial Museum Occasional Paper (4): 1-97.

Stejneger, Leonhard, and Thomas Barbour. 1943. A checklist of North Americanamphibians and reptiles. [5th edition] Bulletin of the Museum of ComparativeZoology 93(1): 1-260.

1945

Carl, Clifford G., and Ian M. Cowan. 1945. Notes on some frogs and toads of BritishColumbia. Copeia 1945(1):52-53.

Schonberger, Clinton F. 1945. Food of some amphibians and reptiles of Oregon andWashington. Copeia 1945(2):120-121.

1947

Livezey, Robert L., and Albert H. Wright. 1947. A synoptic key to the salientian eggs ofthe United States. American Midland Naturalist 37(1):179-222.

Pickwell, Gayle. 1947. Amphibians and reptiles of the Pacific States. StanfordUniversity Press, Stanford, California. 236pp.

1948

Mills, Colin R. 1948. A check list of the reptiles and amphibians of Canada.Herpetologica 4(2nd supplement): 1-15.

1949

Creso, Irene 0. 1949. A comparative study of the gross anatomy of the circulatorysystems of the salientia of the Northwest. MSc Thesis, University of Puget Sound,Department of Biology, Tacoma, Washington. 11lpp.

Stewart, Doris M. 1949. Skeletal structures of the genus Rana of the Pacific States. MScthesis, University of Puget Sound, Department of Biology, Tacoma, Washington.42pp.

Wright, Albert H., and Anna A. Wright. 1949. Handbook of frogs and toads (3rdedition). Comstock Publishing Associates, Ithaca, New York. 640 pp.


1951

Bruson, R. Bruce, and Herald A. Demaree, Jr. 1951. The herpetology of the MissionMountains, Montana. Copeia 1951(4):306-308.

Dunlap, Donald G., and Robert M. Storm. 1951. The cascade frog in Oregon. Copeia1951(1):81.

Stebbins, Robert C. 1951. Amphibians of western North America. University ofCalifornia Press, Berkeley, California. 539pp.

1952

Bandy, P. J. 1952. Report on the biological investigations carried out in E. C. ManningPark, British Columbia, during the summer of 1952. Unpublished Report, BritishColumbia Forest Service, Victoria. 193 pp.

Carl, Clifford G., Charles J. Guiguet, and George A. Hardy. 1952. A natural historysurvey of the Manning Park Area, British Columbia. British Columbia ProvincialMuseum Occasional Paper (9): 1-130.

Ferguson, Denzel E. 1952. The distribution of reptiles and amphibians of WallowaCounty, Oregon. Herpetologica 8(3):66-68.

Orton, Grace L. 1952. Key to the genera of tadpoles in the United States and Canada.American Midland Naturalist 47(2):382-395.

Skousen, Don B. 1952. A taxonomic survey of the eggs and larvae of some species ofUtah amphibia. MA thesis, Department of Zoology, Brigham Young University,Provo, Utah.

1953

Carpenter, Charles C. 1953a. An ecological study of the herpetofauna of the GrandTeton-Jackson Hole area of Wyoming. Copeia 1953(3):170-174.

Carpenter, Charles C. 1953b. Trapping technique for aquatic salamanders. Herpetologica8(4):183.

Farner, Donald S., and James Kezer. 1953. Notes on the amphibians and reptiles ofCrater Lake National Park. American Midland Naturalist 50(2):448-462. [No Ranapretiosa in Crater Lake National Park, only R. cascadae]

Hock, Raymond J. 1953. Alaskan zoogeography and Alaskan amphibia. Proceedings ofthe Alaskan Science Conference (Anchorage) 4:201-206.

Schmidt, Karl P. 1953. A check list of North American amphibians and reptiles, sixthedition. American Society of Icthyologists and Herpetologists, University ofChicago Press, Chicago, Illinois. 280 pp.

1954

Carpenter, Charles C. 1954. A study of amphibian movement in the Jackson HoleWildlife Park. Copeia 1954(3): 197-200.


1954 (continued)

Ferguson, Denzel E. 1954. An annotated list of the amphibians and reptiles of UnionCounty, Oregon. Herpetologica 10(3):149-152.

Stebbins, Robert C. 1954. Amphibians and reptiles of western North America. McGraw-Hill Book Company, Inc., New York, New York. 536pp.

Zweifel, Richard G. 1954. A new Rana from the Pliocene of California. Copeia1954(2):85-87.

1955

Brunson, R. Bruce. 1955. Check list of the amphibians and reptiles of Montana.Proceedings of the Montana Academy of Sciences 15:27-29.

Dunlap, Donald G. 1955. Inter- and intraspecific variation in Oregon frogs of the genusRana. American Midland Naturalist 54(2):314-331.

Logier, E. B. S., and G. C. Toner. 1955. Check-list of the amphibians and reptiles ofCanada and Alaska. Contributions of the Royal Ontario Museum of Zoology andPalaeontology (41):1-88.

Moore, J. E., and E. H. Strickland 1955. Further notes on the foods of Albertaamphibians. American Midland Naturalist 54(1):253-256.

Slater, James R. 1955. Distribution of Washington amphibians. Occasional Papers,Department of Biology, College of Puget Sound (16): 120-154.

1956

Committee [Conant, Roger; Fred R. Cagle, Coleman J. Goin, Charles H. Lowe, Jr.,Wilfred T. Neill, M. Graham Netting, Karl P. Schmidt, Charles E. Shaw, RobertC. Stebbins, and Charles M. Bogert]. 1956. Common names for North Americanamphibians and reptiles. Copeia 1956(3): 172-185.

Ferguson, Denzel E. 1956. The distribution of Rana sylvtica cantabrigensis Baird inwestern Canada and Alaska. Herpetologica 12(2): 132.

1957

Burns, Douglas M. 1957. A new locality for the leopard frog in Oregon. Herpetologica13(2):211.

Carpenter, Charles C. 1957. Aggregation behavior of tadpoles of Rana p. pretiosa.Herpetologica 9(2):77-78.

Dumas, Philip C. 1957. Rana sylvatica Le Conte in Idaho. Copeia 1957(2):150-151.

Manville, Richard H. 1957. Amphibians and reptiles of Glacier National Park, Montana.Copeia 1957(4):308-309.


1957 (continued)

Middendorf, L. J. 1957. Observations on the early spring activities of the western spottedfrog (Rana pretiosa pretiosa) in Gallatin County, Montana. Proceedings of theMontana Academy of Sciences 17:55-56.

Turner, Frederick B. 1957a. The ecology and morphology of Rana pretiosa inYellowstone Park, Wyoming. PhD dissertation, University of California,Berkeley, California.

Turner, Frederick B. 1957b. The ecology and morphology of Rana pretiosa inYellowstone Park, Wyoming. American Doctoral Dissertations 18(7): 146. [listingonly]

1958

Ferguson, Denzel E., K. E. Payne, and Robert M. Storm. 1958. Notes on theherpetology of Baker County, Oregon. Great Basin Naturalist 18(2):63-65.

Turner, Frederick B. 1958a. Life-history of the western spotted frog in YellowstoneNational Park. Herpetologica 14(2):96- 100.

Turner, Frederick B. 1958b. Some parasites of the western spotted frog, Rana p.pretiosa, in Yellowstone Park. Journal of Parasitology 44(2):182.

1959

Corbit, Carl D., and Larry Nelson. 1959. Interorderal amplexus. Herpetologica15(2): 104.

Dunlap, Donald G. 1959. Notes on the amphibians and reptiles of Deschutes County,Oregon. Herpetologica 15(4): 173-177.

Gosner, Kenneth L. 1959. Systematic variations in tadpole teeth with notes on food.Herpetologica 15(4):203-210.

Ruibal, Rodolfo. 1959. The ecology of a brackish water population of Rana pipiens.Copeia 1959(4):315-322.

Turner, Frederick B. 1959a. Variation in skeletal proportions of Rana p. pretiosa inYellowstone Park, Wyoming. Copeia 1959(1):63-68.

Turner, Frederick B. 1959b. Pigmentation of the western spotted frog, Rana pretiosapretiosa, in Yellowstone Park, Wyoming. American Midland Naturalist 61(1):162-176.

Turner, Frederick B. 1959c. An analysis of the feeding habits of Rana p. pretiosa inYellowstone Park. American Midland Naturalist 61(2):403-413.

1960

Metter, Dean E. 1960. The distribution of amphibians in eastern Washington. MScthesis, Washington State University, Department of Zoology, Pullman. 89pp.


1960 (continued)

Turner, Frederick B. 1960a. Population structure and dynamics of the western spottedfrog, Rana pretiosa pretiosa Baird and Girard, in Yellowstone Park, Wyoming.Ecological Monographs 30(3):251-278.

Turner, Frederick B. 1960b. Tests of randomness in recaptures of Rana pretiosa pretiosa.Ecology 41(1):237-239.

Turner, Frederick B. 1960c. Postmetamorphic growth in anurans. American MidlandNaturalist 64(2):327-338.

Wade, Keith. 1960. Report on the reptiles and amphibians of Wells Gray Park - May andJune, 1960. Unpublished Report, British Columbia Parks Branch, Victoria. 7 pp.

1961

Cochran, Doris M. 1961. Type specimens of reptiles and amphibians in the United StatesNational Museum. United States National Museum Bulletin (220):1-291.

Logier, E. B. S., and G. C. Toner. 1961. Check list of the amphibians and reptiles ofCanada and Alaska. Contribution of the Life Sciences Division, Royal OntarioMuseum (53):1-92.

1962

Parson, Thomas S., and Ernest E. Williams. 1962. The teeth of the amphibia and theirrelation to amphibian phylogeny. Journal of Morphology 110(4):375-389.

Turner, Frederick B. 1962a. The demography of frogs and toads. Quarterly Review ofBiology 37(4):303-314.

Turner, Frederick B. 1962b. An analysis of geographic variation and distribution of Ranapretiosa. American Philosophical Society Yearbook 1962:325-328.

Waits, J. A. 1962. Parasitic helminths as aids in studying the distribution of species ofRana in Idaho. Transactions of the Illinois Academy of Sciences 54:152-156.

1963

Gorham, Stanley W. 1963. The comparative number of species of amphibians in Canadaand other countries. HI. Summary of species of anurans. Canadian Field-Naturalist 77(1):13-48.

Hebard, William B., and R. Bruce Brunson. 1963. Hind limb anomalies of a westernMontana population of the Pacific tree frog, Hyla regilla Baird and Girard. Copeia1963(3):570-572.

1964



1964 (continued)

Dumas, Philip C. 1964. Species-pair allopatry in the genera Rana and Phrynosoma.Ecology 45(1):178-181.

Lehmann, Donald L. 1964 [1965]. Intestinal parasites of northwestern amphibians.Yearbook of the American Philosophical Society 1964:284-285.

Metter, Dean E. 1964. Morphological and ecological comparison of two populations ofthe tailed frog, Ascaphus truei Stejneger. Copeia 1964(1):181-195.

Slater, James R. 1964a. A key to the adult amphibians of Washington State. OccasionalPapers, Department of Biology, University of Puget Sound (25):235-236.

Slater, James R. 1964b. County records of amphibians for Washington. OccasionalPapers, Department of Biology, University of Puget Sound (26):237-242.

-1965

Banta, Benjamin H. 1965. A distributional checklist of recent amphibians inhabiting thestate of Nevada. Occasional Papers of the Biological Society of Nevada (7): 1-4..

Johnson, Oliver W. 1965a. Early development, embryonic temperature tolerance and rateof development in Rana pretiosa lutiventris Thompson. PhD dissertation, OregonState University, Corvallis, Oregon.

Johnson, Oliver W. 1965b. Early development, embryonic temperature tolerance and rateof development in Rana pretiosa luteiventris Thompson. Dissertation Abstracts25(12):7427. [abstract]

Schell, S. C. 1965. The life history of Haematoloechus breviplexus Stafford, 1902(Trematoda: Haplometridae). Journal of Parasitology 51(4):587-593.

1966

Dumas, Philip C. 1966. Studies of the Rana species complex in the Pacific Northwest.Copeia 1966(1):60-74.

Stebbins, Robert C. 1966. A field guide to western reptiles and amphibians. HoughtonMifflin Company, Boston, Massachusetts. 279pp.

Storm, Robert M. 1966. Amphibians and reptiles. Northwest Science 40(4):138-141.

1967

Carlson, Dennis S. 1967. A checklist of amphibians and reptiles of Washington. Bulletinof the Pacific Northwest Herpetological Society 2(1):7-8.

Underhill, J. E. 1967. Report of a trial program of interpretation, Kokanee Creek park -Aug. 1967. Unpublished Report, British Columbia Parks Branch, Victoria. 9 pp.


1968

Chantell, Charles J. 1968. Some osteological observations on the genus Rana. Journal ofHerpetology 1(1-4):121. [abstract]

1969

Altig, Ronald G. 1969a. Developmental cranial osteology of three species of Rana(Anura). PhD dissertation, Oregon State University, Corvallis, Oregon. 81pp.

Altig, Ronald G. 1969b. Developmental cranial osteology of three species of Rana(Anura). Dissertation Abstracts International 30B:468.

Black, Jeffrey H. 1969. The frog genus Rana in Montana. Northwest Science 43(4): 191-195.

Clark, Glen W., Jack Bradford, and Ronald Nussbaum. 1969. Blood parasites of somePacific Northwest amphibians. Bulletin of the Wildlife Disease Association5(2):117-11-8.

Lardie, Richard L. 1969. Checklist of the amphibians of Pierce County, Washington withreference to those found on McChord Air Force Base. Bulletin of the PacificNorthwest Herpetological Society 4(2):17-21.

Licht, Lawrence E. 1969a. Comparative breeding behavior of the red-legged frog (Ranaaurora aurora) and the western spotted frog (Rana pretiosa pretiosa) insouthwestern British Columbia. Canadian Journal of Zoology 47(6):1287-1299.

Licht, Lawrence E. 1969b. Palatability of Rana and Hyla eggs. American MidlandNaturalist 82(1):296-298.

Morris, R. L., and W. W. Tanner. 1969. The ecology of the western spotted frog, Ranapretiosa pretiosa Baird and Girard, a life history study. The Great Basin Naturalist29(2):45-81.

1970

Altig, Ronald G. 1970. A key to the tadpoles of continental United States and Canada.Herpetologica 26(2):180-207.

Chantell, Charles J. 1970. Upper Pliocene frogs from Idaho. Copeia 1970(4): 654-664.

Cochran, Doris M., and Coleman J. Goin. 1970. The new field book of reptiles andamphibians. C. P. Putnam's Sons, New York, New York. 359pp.

Dronen, Norman 0. 1970. The life history of a species of Cephalogonimus (Trematoda:Cephalogonimidae) from Ambystorna tigrinum (Green) of eastern Washington.MSc thesis, Eastern Washington State College, Department of Biology, Cheney,Washington. 50pp.

Haertel, John D., and Robert M. Storm. 1970. Experimental hybridization between Ranapretiosa and Rana cascadae. Herpetologica 26(4):436-446..


1971

Grass, Al. 1971. Wells Gray Park Naturalist's Report - 1971. Unpublished Report,British Columbia Parks Branch, Victoria. 24 pp.

Hodge, Robert P. 1971. Often seen...little understood! Washington Wildlife 23(3):10-11.

Licht, Lawrence E. 197 la. Breeding habitat and embryonic thermal requirements of thefrogs, Rana aurora aurora and Rana pretiosa pretiosa, in the Pacific Northwest.Ecology 52(1):116-124.

Licht, Lawrence E. 1971b. The ecology of coexistence in two closely related species offrogs (Rana). PhD dissertation, University of British Columbia, Vancouver,British Columbia. 155pp.

Paulson, Dennis. 1971. Key to Washington State reptiles and amphibians. Bulletin of thePacific Northwest Herpetological Society 5(1):27-33.

Ryder, Glen R. 1971. Pelican Park-Naturalist's report for 1971. Unpublished Report,British Columbia Parks Branch, Victoria. 91 pp.

1972

Bols, Niels C. 1972. Studies on sperm histones in amphibia and chondrichthytes. MScthesis, University of British Columbia, Department of Zoology, Vancouver. 74pp.

Bols, Niels C., and H. E. Kasinsky. 1972. Basic protein composition of anuran sperm: Acytochemical study. Canadian Journal of Zoology 50(2): 171-177.

Licht, Lawrence E. 1972. The ecology of coexistence in two closely related species offrogs (Rana). Dissertation Abstracts International 33B(6):2563.

Paulson, Dennis. 1972. Reptiles and amphibians of Washington. Bulletin of the PacificNorthwest Herpetological Society 5(2): 13-15.

Turner, Frederick B., and Philip C. Dumas. 1972. Rana pretiosa. Catalogue of AmericanAmphibians and Reptiles: 119.1-119.4.

Wright, Richard T. 1972. Wells Gray Provincial Park Naturalist's Report 1972.Unpublished Report, British Columbia Parks Branch, Victoria. 55 pp.

1973

Goward, Trevor. 1973. Naturalist's Report 1973 - Champion Lakes Provincial Park.Unpublished Report, British Columbia Parks Branch, Victoria: 1-50.

Larsen, John H., Jr. 1973. Effects of cytochalasin B on the ultrastructure of Ranapretiosa tadpole epidermis. Anatomical Record 177(3):427-440.

Wallace, D. G., M. C. King, and A. C. Wilson. 1973. Albumin differences among ranidfrogs: taxonomic and phylogeneic implications. Systematic Zoology 22(1):1-13.


1974

Altig, Ronald G., and William L. Pace. 1974. Scanning electron photomicrographs oftadpole labial teeth. Journal of Herpetology 8(3):247-25 1.

1974 (continued)

Brown, Vinson. 1974. Reptiles and amphibians of the west. Naturegraph PublishersInc., Healdsburg, California. 79pp.

Dronen, Norman O., and Bruce Z. Lang. 1974. The life cycle of Cephalogonimussalamandrus sp. n. (Digenea: Cephalogonimidae) from Ambystoma tigrinum(Green) from eastern Washington. Journal of Parasitology 60(1):75-79.

Green, David. 1974. Reptiles and amphibians of Shuswap Area - An annotated list.Unpublished Report, British Columbia Parks Branch, Victoria. 2 pp.

Gorham, Stanley W. 1974. Checklist of World amphibians up to January 1, 1970. NewBrunswick Museum, Saint John. 172pp.

Haertel, John D., Alfred Owczarzak, and Robert M. Storm. 1974. A comparative studyof the chromosomes from five species of the genus Rana (Amphibia: Salientia).Copeia 1974(1):109-114.

Licht, Lawrence E. 1974. Survival of embryos, tadpoles, and adults of the frogs Ranaaurora aurora and Rana pretiosa pretiosa sympatric in southwestern BritishColumbia. Canadian Journal of Zoology 52(5):613-627.

Parkin, T. W. 1974. Kokanee Glacier Provincial Park - Fisheries and Wildlife Inventory,July - August 1974. Unpublished Report, British Columbia Parks Branch,Victoria. 38 pp.

Perry, Alfred E. 1974. A survey of terrestrial vertebrate fauna of the Touchet Riverdrainage, Washington. Unpublished Report, Walla Walla College, College Place,Washington. 107pp. [A report submitted to US Department of the Interior, Bureauof Reclamation, Boise, Idaho]

1975

Brown, Herbert A. 1975. Reproduction and development of the red-legged frog, Ranaaurora, in northwestern Washington. Northwest Science 49(4):241-252.,

Dowling, Herndon G. 1975. A classification and checklist of the species of amphibiansand reptiles found in the United States and Canada. pp. 175-189. In: Herndon G.Dowling (editor), 1974 Yearbook of Herpetology, Volume 1. HerpetologicalInformation and Search Systems Publications in Herpetology No. 8, New York.256pp.

Green, David M. 1975. Reptiles and amphibians of Shuswap area 1975. UnpublishedReport, British Columbia Parks Branch, Victoria. 3 pp.

Gilboa, Itzchak. 1975. Karyotypes of amphibians and reptiles: a bibliographic review.pp.9 1-156. In: Herndon G. Dowling (editor), 1974 Yearbook of Herpetology,Volume 1. Herpetological Information and Search Systems Publications inHerpetology No. 8, New York. 256pp.


1975 (continued)

Licht, Lawrence E. 1975. Comparative life history features of the western spotted frog,Rana pretiosa, from low- and high-elevation populations. Canadian Journal ofZoology 53(9):1254-1257.

Osmond-Jones, E., J. Bonner, M. Sather, B. Smith, and M. MacColl. 1975. A fisheriesand wildlife survey of the Burnie Lakes Park proposal. Unpublished Report,British Columbia Parks Branch, Victoria. 44 pp.

1976

Hazelwood, W. Grant. 1976. Tweedsmuir Park initial Wildlife and Fisheries inventory(Area A to D). Unpublished Report, British Columbia Parks Branch, Victoria.220 pp.

Hodge, Robert P. 1976. Amphibians and reptiles in Alaska, the Yukon, and NorthwestTerritories. Alaska Northwest Publishing Company, Anchorage, Alaska. 89pp.

Maser, Chris. 1976. Frog preys on tiger beetle. Cicindela 8(1): 10.

Robinson, Douglas A. 1976. A survey of the amphibians and reptiles of WhitmanCounty, Washington, and Latah County, Idaho. Unpublished Report, WashingtonDepartment of Game, Olympia. 62pp.

Swift, Pat. 1976. Paul Lake Provincial Park. Unpublished Report, British ColumbiaParks Branch, Victoria. 24 pp.

1977

Bernard, Stephen, and Kenneth F. Brown. 1977. Distribution of mammals, reptiles, andamphibians by BLM physiographic regions and A. W. Kiichler's associations forthe eleven western states. United States Department of the Interior, Bureau of LandManagement Technical Note (301):1-169.

Cannings, Robert A. 1977. Biological investigation of the Blue River headwaters area,Wells Gray Park. Unpublished Report, British Columbia Parks Branch, Victoria.28 pp.

Dunlap, Donald G. 1977. Wood and western spotted frogs (Amphibia, Anura, Ranidae)in the Big Horn Mountains of Wyoming. Journal of Herpetology 11(1):85-87.

Osmond-Jones, E., M. Sather, W. G. Hazelwood, and B. Ford. 1977. Wildlife andfisheries inventory of Spatsizi Wilderness and Tatlatui Provincial Parks BritishColumbia. Unpublished Report, British Columbia Parks Branch, Victoria. 292pp-

Stephens, J. C. 1977. Hydrologic reconnaissance of the Tule Valley drainage basin, Juaband Millard Counties. Utah Technical Publication Division of Water Rights (56).

Thomas, Robert A. 1977. Selected bibliography of certain vertebrate techniques. UnitedStates Department of the Interior, Bureau of Land Management Technical Note(306): 1-88.


1977 (continued)

Wells, Kentwood D. 1977. The social behaviour of anuran amphibians. Animal Behavior25(3):666-693.

Wishard, L. 1977. Larval growth in Rana pretiosa: Ecological and genetic factors. MSthesis, University of Montana, Missoula, Montana.

1978

Bowman, Irene. 1978. Status of herptiles in Canada. Unpublished Report, OntarioMinistry of Natural Resources, Ottawa. 166pp.

Case, Susan M. 1978. Biochemical systematics of members of the genus Rana native towestern North America. Systematic Zoology 27(3):299-31 1.

Collins, Joseph T., James E. Huheey, James L. Knight, and Hobart M. Smith. 1978.Standard common and scientific names for North American amphibians andreptiles. Society of the Study of Amphibians and Reptiles, Herpetological Circular(5): 1-65.

Daugherty, Charles H., L. Wishard, and L. R. Daugherty. 1978. Sexual dimorphism inan anuran response to severe thermal stress (Amphibia, Anura). Journal ofHerpetology 12(3):431-432.

Ferguson, Denzel E., and Nancy E. Ferguson. 1978. Oregon's Great Basin Country.Gail Graphics, Burns, Oregon. 178 pp.

Kezer, James. 1978. The Gold Lake bog. Nature trails. Eugene Natural History SocietyNewsletter 12(6):5-7.

Miller, J. D. 1978. Observations on the diets of Rana pretiosa, Rana pipiens, and Bufoboreas from western Montana. Northwest Science 52(3):243-249.

Orchard, Stan A. 1978. Amphibians of British Columbia. Unpublished Report, BritishColumbia Fish and Wildlife Branch, Victoria. 92 pp.

Schaub, David L., and John H. Larsen, Jr. 1978. The reproductive ecology of the Pacifictreefrog (Hyla regilla). Herpetologica 34(4):409-416.

Tanner, Wilmer W. 1978. Zoogeography of reptiles and amphibians in the IntermountainRegion. pp. 43-53. In: Intermountain biogeography: A symposium. Great BasinNaturalist Memoir 2.

1979

Behler, John L., and F. Wayne King. 1979. The Audubon Society field guide to NorthAmerican reptiles and amphibians. Alfred A. Knopf, Inc., New York, New York.7l9pp.

Bury, R. Bruce, J. H. Wolfheim, and Roger A. Luckenbach. 1979. Restoration ofMedical Lake: Responses of wildlife. Unpublished Report, WashingtonDepartment of Game, Olympia. 6pp.


1979 (continued)

Campbell, R. Wayne. 1979a. Check-list of amphibians and reptiles of British Columbia.Unpublished Report, British Columbia Provincial Museum, Vertebrate ZoologyDivision, Victoria. 5 pp.

Campbell, R. Wayne. 1979b. Checklist of British Columbia amphibians and reptiles.Unpublished Report, British Columbia Provincial Museum, Vertebrate ZoologyDivision, Victoria. 4 pp.

Case, Susan M. 1979. Observations on some cranial foramina in the Ranidae. Copeia1979(2):346-348.

Gregory, Patrick T. 1979. Predator avoidance behavior of the red-legged frog (Ranaaurora). Herpetologica 35(2): 175-184.

Kluge, Arnold G., James S. Farris, and Mary F. Mickevich. 1979. Paraphyly of theRana boylii species group. Systematic Zoology 28(4):627-634.

Leviton, Alan E. 1979. Reptiles and amphibians of North America. Doubleday andCompany, Inc., New York. 250pp.

Taylor, J. Mary. 1979. Amphibians, reptiles and mammals of British Columbia.Department of Zoology, University of British Columbia, Vancouver. 144pp.

1980

British Columbia Fish and Wildlife Branch. 1980. Preliminary amphibian managementplan for British Columbia. British Columbia Ministry of Environment, Fish andWildlife Branch, Victoria. 29 pp.

Cook, Francis R. 1980. Checklist of amphibians and reptiles of Canada. CanadianAmphibian and Reptile Conservation Society Newsletter 18(2):1-6.

Green, David. M., and R. Wayne. Campbell. 1980. The amphibians of British Columbia.Unpublished Report, British Columbia Provincial Museum, Vertebrate ZoologyDivision, Victoria. 105 pp.

Kennedy, Murray J. 1980a. Host-induced variations in Haematoloechus buttensis(Trematoda: Haematoloechidae). Canadian Journal of Zoology 58(3):427-442.

Kennedy, Murray J. 1980b. Geographical variation in some representatives ofHaematoloechus Looss, 1899 (Trematoda: Haematoloechidae) from Canada and theUnited States. Canadian Journal of Zoology 58(6):1151-1167.

MacIntyre, D. H., and R. V. Palermo. 1980. The current status of the amphibia in BritishColumbia. pp. 146-151. In: Richard Stace-Smith, Lois Johns, and Paul Joslin(editors), Proceedings of the Symposium on Threatened and Endangered Speciesand Habitats in British Columbia and the Yukon. British Columbia Ministry ofEnvironment, Fish and Wildlife Branch, Victoria.

McDiarmid, Roy W. 1980. U.S. National Museum specimens of reptiles and amphibiansfrom the United States for the state of Washington. Unpublished Report, UnitedStates National Mtseum, Washington, D.C. 55 pp.


1980 (continued)

Reinking, Larry N., Charles H. Daugherty, and Lynn B. Daugherty. 1980. Plasmaaldosterone concentrations in wild and captive western spotted frogs (Ranapretiosa). Comparative Biochemistry and Physiology 65A(4):517-518.

Schueler, Frederick W., Francis R. Cook, and Donald H. Rivard. 1980. BritishColumbia distribution data from the herpetology collection of the national museumsof Canada. Unpublished Report, National Museums of Canada, HerpetologySection, Ottawa, Ontario. 6pp.

1981

Kennedy, Murray J. 1981. A revision of species of the genus Haematoloechus Looss,1899 (Trematoda: Haematoloechidae) from Canada and the United States.Canadian Journal of Zoology 59(9):1836-1846.

Post, Thomas J., and Thomas Uzzell. 1981. The relationships of Rana sylvatica and themonophyly of the Rana boylii group. Systematic Zoology 30(2):170-180.

1982

St. John, Alan D. 1982. The herpetology of the Wenaha Wildlife Area, Wallowa County,Oregon. Oregon Department of Fish and Wildlife, Nongame Wildlife Program,Technical Report (82-4-03):1-23.

1983

Hillis, David M., and Scott K. Davis. 1983. Evolution of ribosomal DNA: Fifty millionyears of recorded history in the frog genus Rana. Evolution 40(6):1275-1288.

Hillis, David M., John S. Frost, and David A. Wright. 1983. Phylogeny andbiogeography of the Rana pipiens complex: a biochemical evaluation. SystematicZoology 32(2):132-143.

Nussbaum, Ronald A., Edmund D. Brodie, and Robert M. Storm. 1983. The amphibiansand reptiles of the Pacific Northwest. University of Idaho Press, Moscow.

Waldman, Bruce, and Michael J. Ryan. 1983. Thermal advantages of communal eggmass deposition in wood frogs (Rana sylvatica). Journal of Herpetology 17(1):70-72.

Whitaker, John O., Jr., Stephen P. Cross, John M. Skovlin, and Chris Maser. 1983.Food habits of the spotted frog (Rana pretiosa) from managed sites in GrantCounty, Oregon. Northwest Science 57(2):147-154.

1984

Green, David. M., and R. Wayne. Campbell. 1984. The amphibians of British Columbia.British Columbia Provincial Museum Handbook Series (45):1-102.


1984 (continued)

Hovingh, Peter. 1984. Western spotted frog (Rana pretiosa) distribution in the BonnevilleBasin of western Utah and aquatic resource description of Tule Valley (WhiteValley) of western Utah. Unpublished report submitted with spotted frog (Ranapretiosa) petition to the U.S. Fish and Wildlife Service, Denver, Colorado. 1 May1989. 42 pp.

St. John, Alan D. 1984a. The herpetology of Jackson and Josephine Counties, Oregon.Oregon Department of Fish and Wildlife, Nongame Wildlife Program, TechnicalReport (84-2-05):1-78.

St. John, Alan D. 1984b. The herpetology of the upper John Day River drainage,Oregon. Oregon Department of Fish and Wildlife, Nongame Wildlife Program,Technical Report (84-4-05):1-34.

St. John, Alan D. 1984c. The status of the tiger salamander (Ambystoma tigrinum) innorthern Harney County, Oregon with notes on additional herpetofauna. OregonDepartment of Fish and Wildlife, Nongame Wildlife Program, Technical Report(84-5-03): 1-23.

1985

Green, David. M. 1985. Natural hybrids between the frogs Rana cascadae and Ranapretiosa (Anura: Ranidae). Herpetologica 41(3):262-267.


St. John, Alan D. 1985b. The herpetology of the Owyhee River drainage, MalheurCounty, Oregon. Oregon Department of Fish and Wildlife, Nongame WildlifeProgram, Technical Report (85-5-03): 1-83.

Stebbins, Robert C. 1985. A field guide to western reptiles and amphibians. Secondrevised edition, Houghton Mifflin Company, Boston, Massachusetts. 336pp.

1986

Beneski, John T., Jr., Edward J. Zalisko, and John H. Larsen, Jr. 1986. Demographyand migratory patterns of the eastern long-toed salamander, Ambystomamacrodactylum columbianum. Copeia 1986(2):398-408.

Bols, Niels C., M. Mann, and H. E. Kasinsky. 1986. Detection of sperm histonediversity among vertebrates by alkaline fast green staining. Stain Technology61(2):111-119.

Filipski, G. T., and M. V. H. Wilson. 1986. Nerve staining using Sudan Black B and itspotential use in comparative anatomy. Royal Ontario Museum, Life SciencesDivision, Miscellaneous Publication:33-36.

Green, David. M. 1986a. Systematics and evolution of western North American frogsallied to Rana aurora and Rana boylii: karyological evidence. Systematic Zoology35(3):273-282.


1986 (continued)

Green, David. M. 1986b. Systematics and evolution of western North American frogsallied to Rana aurora and Rana boylii: electrophoretic evidence. SystematicZoology 35(3):283-296.

Hayes, Marc P., and Mark R. Jennings. 1986. The decline of ranid frog species inwestern North America: Are bullfrogs (Rana catesbeiana) responsible? Journal ofHerpetology 20(4):490-509.

Hayes, Marc P., and Dana M. Krempels. 1986. Vocal sac variation among frogs of thegenus Rana from western North America. Copeia 1986(4):927-936.

Hovingh, Peter. 1986. Biogeographic aspects of leeches, mollusks, and amphibians inthe intermountain region. Great Basin Naturalist 46(4):736-744.

Licht, Lawrence E. 1986a. Comparative escape behavior of sympatric Rana aurora andRana pretiosa. The American Midland Naturalist 1 15(2):239-247.

Licht, Lawrence E. 1986b. Food and feeding behavior of sympatric red-legged frogs,Rana aurora, and spotted frogs, Rana pretiosa, in southwestern British Columbia.The Canadian Field-Naturalist 100(1):22-31.


Uzzell, Thomas, and Thomas J. Post. 1986. Rana temporaria is not a member of the Ranaboylii group. Systematic Zoology 35(3):414-421.

1987

Briggs, Jeffrey L., Sr. 1987. Breeding biology of the Cascade frog, Rana cascadae, withcomparisons to R. aurora and R. pretiosa. Copeia 1987(1):241-245.

Hovingh, Peter. 1987a. Investigations of Great Basin springs and wetlands for leeches,mollusks and amphibians. Proceedings of the Bonneville Chapter of the AmericanFisheries Society 1987:27-40.

Hovingh, Peter. 1987b. Current status of the western spotted frog (Rana pretiosa) in theBonneville Basin. Unpublished MS, 15 September 1987. 13 pp.




1988

Blaustein, Andrew R. 1988. Ecological correlates and potential functions of kinrecognition in anuran larvae. Behavioral Genetics 18(4):449-464.

Hovingh, Peter. 1988. Amphibian and wetland survey and analysis of the StrawberryRiver drainage in Wasatch County, Utah. Report to the Division of WildlifeResource, State of Utah. 28 pp.

Kirk, James J. 1988. Western spotted frog (Rana pretiosa) mortality following forestspraying of DDT. Herpetological Review 19(3):51-53.

O'Hara, Richard K., and Andrew R. Blaustein. 1988. Hyla regilla and Rana pretiosatadpoles fail to display kin recognition behavior. Animal Behavior 36(3):946-948.

1989


Miller, James L. 1989. Endangered and Threatened Wildlife and Plants; Finding onPetition to List the Spotted Frog. Federal Register 54(199):42529.

Macartney, J. Malcolm. 1989. Diet of the northern Pacific rattlesnake, Crotalus viridisoreganus, in British Columbia. Herpetologica 45(3):299-304.

1990

Lindeman, P. V. 1990. Life history notes: Rana pretiosa. Herpetological Review21(2):38.


Rannala, Bruce H. 1990. Electrophoretic evidence concerning the relationship betweenHaplometrana and Glypthelmins (Digenea: Plagiorchiiformes). Journal ofParasitology 76(5):746-748.

1991


1992

O'Grady, Richard T. (in press). A preliminary report on the life cycle of Glypthelminscaliforniensis (Cort, 1919) Miller, 1930 (Digenea: Plagiorchidae), a parasite ofranid frogs in western North America. Journal of the Helminthological Society ofWashington


1993


Worthing, Patricia. 1993. Endangered and Threatened wildlife and plants; Finding onPetition to list the spotted frog. Federal Register 58(87):27260-27263.

In Press

Leonard, William, and Kelly R. MGAllister. The rediscovery of the spotted frog (Ranapretiosa) in western Washington. Northwest Naturalist.


Appendix III

Western Spotted Frog Localities

1) Benton County: 12 mi (19.2 km) south of Corvallis, McFadden Swamp. Legal: T13S,R5W, extreme south end of Sec 33 [but also probably historical habitat in T14S, R5W,Sec 4]. Specimens: (3) OSUMNH 1426, 5826-7. Collector: Robert M. Storm.Collector's number: None provided. Collection date(s): 15 February 1950 (1426), 8February 1951 (5826-7). Life stage: Adults. Sex: Male (1426), Unknown (5826-7).Comments: Data in the catalogue for OSUMNH 1426 states, "Ca. 10:00 PM, in waterat edge of road (north side) between hunting shack and first bridge at McFadden'sMarsh, 0.5 mi (0.8 km) west of Bruce Station, ca. 12 mi (19.2 km) south ofCorvallis." For OSUMNH 5826-7, the catalogue indicates, "Ca. 10:00 PM, frogs inwater along north edge of road running through marsh, between the corner and firstbridge." Robert Storm (pers. comm.) indicated that is locality is along the north side ofBruce Road, approximately 0.2 mi (0.3 km) west of the north-south jog in Bruce Roadlocated 0.8 mi (1.3 km) west of Highway 99W, and 0.2 mi (0.3 km) east of the bridgeon Bruce Road that crosses Muddy Creek. Storm stated that, in both cases, the frogswere collected on the north side of Bruce Road at a point where the water typicallyflowed across the road in a shallow overflow. USGS topo quad shows McFaddenSwamp as marshy ground ca. 1.5 mi long (north-south) by ca. 0.25 mi wide (east-west). Dunlap (1955) referred to R. pretiosa specimens from this locality that areprobably these specimens, but no listing of specimens was provided. Alan St. John(pers. comm.) indicated that he refers to this locality as Finley National WilldlifeRefuge (St. John 1987a). Ownership: United States Fish and Wildlife Service, FinleyNational Wildlife Refuge. USGS Topo Quad: 7.5' Greenberry [1969, photoinspected1975]. Elevation: ca. 260 ft (80 m).

2) Benton County: Corvallis. Legal: T1 1-12S, RSW, insufficient data to determinesection. Specimens: (1) OSUMNH 10127. Collector: A. C. Chandler. Collector'snumber: None provided. Collection date: 27 April 1965. Life stage: Subadult. Sex:Unknown. Comments: Discussion with Robert Storm (pers. comm.) indicated that thecollector, A. C. Chandler, was a well known parasitologist associated with OregonState University. Chandler could not be located to potential obtain additional data onthis locality. Ownership: Probably private, but locality data currently too vague toknow precisely. USGS Topo Quad: 7.5' Corvallis [1969, photorevised 1986].Elevation: ca. 220-240 ft (67-73 m).

3) Clackamas County: Clackamas Lake, at edge of lake. Legal: T 5 S, R 8.5 E, NE 1/4 ofNE 1/4 of Sec 35 and NW 1/4 of NW 1/4 of Sect 36 [but probably collected in Section35. Specimens: (1) OSUMNH 4451. Collector: Ed Park. Collector's number: Noneprovided. Collection date: 22 July 1956. Life stage: Adult. Sex: Unknown.Comments: The collector, Ed Park (P.O. Box 887, Bend, OR 97709; Ph: 576-2242),was contacted, but a stroke recently deprived Park of his memory, so no further detailsconcerning this collection could be obtained from the collector. Clackamas Lake is asmall lake on the Oak Grove Fork of the Clackamas River located approximately 1.6 mi(2.5 km) in a straight line almost due southeast of Timothy Lake.

A specimen at the Carnegie Museum of Natural History listed simply as Rana sp.[CM 32730] from Clackamas Lake, Oregon Skyline Road collected on 26 June 1952by R. H. McCauley may be a western spotted frog. At this juncture, no pressingattempt to obtain this specimen for verification because the OSUMNH specimen for thislocality was verified as being a western spotted frog and the Carnegie specimen wascollected over four years earlier, so even if the Carnegie specimen was a spotted frog, itcould not improve on the date of the most recent collection; and no more precise locality


was associated with the Carnegie specimen, so no locality data would be gained byverifying the specimen. Nevertheless, it would be worthwhile to verify the identity ofthis specimen at a later juncture to gain a more precise understanding of the variabilityamong spotted frogs west of the Cascades. Ownership: Forest Service, Mt. HoodNational Forest, Bear Springs District. USGS Topo Quad: 7.5' Mt. Wilson [1985provisional edition]. Elevation: 3329 ft (1015 m).

4) Clackamas County: Crater Meadows; (TSS, R81/2E, N1/2 for PSU specimen only).Legal: TSS, R8.5E, east portion of Sec 11. Specimens: (3) OSUMNH 0613-4,PSU 1095. Collectors: Stanley G. Jewett, Jr (OSUMNH), Clea M.Neidert (PSU).Collector's number: For OSUMNH specimens only: SJ 276 (0613), SJ 278 (0614).Collection date(s): 9 October 1930 (OSUMNH), 8 October 1966 (PSU). Life stage:Juvenile (OSUMNH 0613), Adult (OSUMNH 0614), Adult (PSU 1095). Sex:Unknown (OSUMNH 0613), Male (OSUMNH 0614), Female (PSU 1095).Comments: The collector of the two OSUMNH specimens, Stanley G. Jewett, Jr.,passed away early in 1991. His widow, Doris Jewett, gave permission to examine theannotated catalogue of the amphibian and reptile specimens he had collected in Oregon.Matching Jewett's original specimen numbers (SJ 276, 278) and the collection dates,the catalogue indicated that the two frogs were collected at Crater Meadows. CraterMeadows, also known as Little Crater Meadows, are located over a rather broad areaalong the inflow point of Crater Creek into Timothy Lake. In the OSUMNH catalogue,the locality associated with these specimens was erroneously transcribed as CraterLake, Klamath County. However, the numbers and dates in Jewett's catalogue showthe latter to be in error.

Neither Clea M. Neidert (pers. comm.), who collected the PSU specimen and isnow an instructor at Portland City College (Sylvan Campus), nor Ellen Benedict (pers.comm.), a collecting colleague on the same trip, recalled details of this collection.

Robert Penson, then Assistant District Biologist for the Bear Springs RangerDistrict of the Mt. Hood National Forest, indicated that Little Crater Meadows iscurrently about 40 ac (ca. 15 ha) of low emergent marsh and meadow. A beaver pondis present on the north inflow stream into the meadow. Ownership: Forest Service, Mt.Hood National Forest, Bear Springs District. USGS Topo Quads: 7.5' Wapinita Pass[1985 provisional edition] and 7.5' Wolf Peak [1985 provisional edition]. Elevation:Ca. 3300 ft (1006 m).

5) Jackson County: NW end of Little Hyatt Reservoir. Legal: T39S, R3E, Sec 20.Specimens: (2) SOSC 823-4. Collector: J. Brown. Collector's number: 94 (823), 83(824). Collection date(s): 10 May 1971. Life stage: Adult (823), postmetamorphicjuvenile (824). Sex: Female (823) and unknown (824). Comments: No notes existassociated with the specimens, but Dr. Steve Cross indicated that the northwest end ofthis reservoir has a small area of marsh. Ownership: Bureau of Land Management,Medford District, Ashland Resource Area and private. USGS Topo Quad: 7.5' HyattReservoir [1988 provisional edition]. Elevation: Ca. 4625 ft (1409 m).

6) Klamath County: Fort Klamath. Legal: T33S, R7.5E, SE 1/4 of Sec 23. Specimens:(23) CAS 44365-78, USNM 14728 (9 specimens with the same number). Collector:Joseph R. Slevin (44365-78), Dr. J. C. Merrill (14728). Collector's number: None.Collection date(s): 24 June 1918 (44365-78), 29 October 1887 (14728). Life stage:Adults and subadults. Sex: Females, males, and unknown. Comments: No notes existassociated with the specimens, but examination of old topographic engineers mapsindicates that significant marshes occurred in the vicinity of Fort Klamath along FortCreek and the Wood River, the likely collection location of these specimens.Ownership: Private or Forest Service (Winema National Forest, Klamath Ranger


District). USGS Topo Quads: 7.5' Fort Klamath [1985 provisional edition].Elevation: Ca. 4155 ft (1266 m).

7) Klamath County: Mountains near Fort Klamath. Legal: T33S, R7.5E, locality tooimprecise to provide a section. Specimens: (1) USNM 11531. Collector: Henry W.Henshaw. Collector's number: None. Collection date: 29 August 1878. Life stage:Subadult. Sex: Female. Comments: Specimen described in text of explorations westof the 100th meridian. Description in text and journals too imprecise to retrieve detailsof locality. Ownership: Probably Forest Service, Winema National Forest, KlamathRanger District. USGS Topo Quads: 7.5' Fort Klamath [1985 provisional edition].Elevation: Ca. 4100-4200 ft (1250-1280 m).

8) Klamath County: Klamath Falls. Legal: T38S, R9E, Sec 31. Specimens: (14) CAS20414-27. Collector: John Van Denburgh. Collector's number: None. Collectiondate(s): 8 September 1909 (20414-22), 15 September 1909 (20423-27). Life stage:Adults. Sex: Females (20414-22), unknown (20423-27; condition of specimens toopoor to sex). Comments: No notes were associated with the specimens. Examinationof old maps indicates marshes were present in the vicinity of Klamath Falls at the timeof collection, and were likely the site of collection. Ownership: Probably private, butwithout further data, it is impossible to understand ownership of this locality precisely.USGS Topo Quads: 7.5' Klamath Falls [1985 provisional edition]. Elevation: Ca.4150 ft (1265 m).

9) Klamath County: Klamath Lake. Legal: T38S, R9E, locality too imprecise to provide asection. Specimens: (2) USNM 3360, ANSP 14532. Collector: Lieutenant Willamson(USNM 3360), Edward Drinker Cope (ANSP 14532). Collector's number: None.Collection date: No day, No month 1850s (USNM 3360); no day, no month, 1879(ANSP 14532). Life stage: Subadult (USNM 3360), juvenile (ANSP 14532). Sex:Unknown. Comments: Willamson's journals indicate the exploring party skirted whatis now the east side of Upper Klamath Lake, and probably collected this specimen inthe vicinity of what is now known as Hank's Marsh [= Upper Klamath NationalWildlife Refuge] below Bald Hill. No notes exist for the Cope specimen. Somepossibility exists that the Cope and Williamson specimens were collected on LowerKlamath Lake, when this lake was significant prior to 1912, but no independent dataexist to attempt to verify this alternative. Ownership, elevation, and quad are based onthe assumption the collection took place along the subsequently well-traveled route thatWilliamson took along the east margin of Upper Klamath Lake.Ownership: Probablyprivate, but without further data, it is impossible to understand ownership of thislocality precisely. USGS Topo Quads: 7.5' Wocus [1985 provisional edition].Elevation: Ca. 4150 ft (1265 m).

10) Klamath County: Sprague River vicinity of Spring Creek. Legal: T36S, R12W, northhalf of Sec 14. Specimens: (1) CAS 44299. Collector: Joseph R. Slevin. Collector'snumber: None. Collection date: 15 June'1918. Life stage: Adults. Sex: Females(20414-22), unknown (20423-27). Comments: No notes exist associated with thespecimens, but examination of topographic maps indicates marshes were present alongthe Sprague River just northeast of Beatty, the site of collection. Ownership: Probablyprivate, but without further data, it is impossible to understand ownership of thislocality precisely. USGS Topo Quads: 7.5' Beatty [1988 provisional edition].Elevation: Ca. 4315 ft (1315 m).

11) Klamath County: North fork of Sprague River above junction with S Fork. Legal:T36S, R14E, NW 1/4 of NW 1/4 of Sec 18. Specimens: (1) UMMZ 77901.Collector: Carl L. Hubbs and family. Collector's number: 120A. Collection date: 9


August 1934. Life stage: Subadult. Sex: Unknown. Comments: Carl Hubbs notesassociated with these specimens indicated the presence of a vast shallow marsh at thislocality. This marsh is indicated on current aerial photographs, but considerablymodified and reduced in areal extent. The site of the original collection is near the pointwhere Campbell Road crosses the North fork of the Sprague River. Ownership:Private, without further data, it is not possible to know on which of the properties ofcurrent landowners Hubbs made his 1934 collection. USGS Topo Quads: 7.5' Bly[1988 provisional edition]. Elevation: Ca. 4320 ft (1317 m).

12) Klamath County: Sprague River opposite Ferguson Butte [= Mountain]. Legal: T36S,R13E, Sec 15 or 16. Specimens: (1) UMMZ 77902. Collector: Carl L. Hubbs andfamily. Collector's number: 121. Collection date: 10 August 1934. Life stage: Adult.Sex: Female. Comments: Carl Hubbs notes associated with these specimens indicatedthe presence of marshy ground at this locality. The site of the original collection is onthe Sprague River south of Ferguson Mountain (then called Ferguson Butte) near apoint about six miles east of Beatty along state highway 140. Ownership: Private,without further data, it is not possible to know on which of the properties of currentlandowners Hubbs made his 1934 collection. USGS Topo Quads: 7.5' FergusonMountain [1988 provisional edition]. Elevation: Ca. 4315 ft (1315 m).

13) Lane County: Gold Lake Bog, at edge of lake (for OSUMNH specimen); above north-west corner of Gold Lake [Gold Lake Bog]. Legal: T22S, R6E, north 1/3 of Sec 29.Specimens: (12) OSUMNH 10218; MVZ 186428, 193594-5; UMMZ 151846[includes 8 specimens]. Collector: Ronald G. Altig and John D. Haertel (10218),David M. Green (186428), John H. Applegarth (193594-5), Ronald G. Altig(151846). Collector's number: None (10218), 1247 (186428), DMG 1612-3 (193594-5), A-12 (151846). Collection date(s): August 1966 (10218), date of collection notprovided; October 1982 (186428), date of collection not provided; 26 September 1984(193594-5); 5 August 1968 (151846). Life stage: Subadults (all collections except151846], 151846 includes two metamorphosing individuals and six tadpoles. Sex:Unknown. Comments: Gold Lake and Gold Lake Bog lie on Salt Creek, a tributary ofthe Middle Fork of the Willamette River. The bog is located on the upstream end of thelake and is as large or slightly larger than the lake itself. The Cascades frog (Ranacascadae) is also present at this locality. This is the locality from which Green (1985)reported natural hybrids [MVZ 193588-9] between the spotted frog and the Cascadesfrog that John Applegarth had collected. Ownership: Forest Service, WillametteNational Forest, Oakridge Ranger District. USGS Topo Quads: 7.5' Waldo Lake[1986 provisional edition]. Elevation: 4807-4830 ft (1465-1472 m).

14) Lane County: Goose Lake, Mink Lake Basin. Legal: T19S, R7E, Sec 8 and 9.Specimens: (7) UMMZ 151848 [includes 7 specimens]. Collector: Ronald G. Altig.Collector's number: A-212. Collection date: Date of collection not provided. Lifestage: Juveniles. Sex: Unknown. Comments: In the Michigan catalogue, this localityis erroneously indicated as being in Deschutes County because the entire Mink LakeBasin is in Lane County. A marsh adjacent to Goose Lake is indicated on the USGStopo quad. Storm (1986) makes allusion to this record as Mink Lake rather than theMink Lake Basin. Spotted frogs are unlikely to be present in Mink Lake, which at5,034 ft is a relatively barren-edged lake and has habitat characteristic of R. cascadae.This record also appears in the Oregon Natural Heritage Database. Ownership: ForestService, Willamette National Forest, McKenzie Ranger District. USGS Topo Quads:7.5' Packsaddle Mountain [1963, photorevised 1981]. Elevation: Ca. 4758 ft (ca.1450 m).


15) Lane County: Oregon State Highway 36, 2.5 mi (4 km) west of junction with Highway99, 4.5 mi (7.2 km) southwest of Junction City. Legal: Ti iS, R5W, center of eastedge of Sec 12. Specimens: (3) CAS-SU 5906-8. Collector: Margaret Storey.Collector's number: none. Collection date: 24 June 1940. Life stage: Metamorph withtail (5906), larvae with back legs (5907-8). Sex: Unknown. Comments: This localityis just west of where Highway 36 crosses Amazon Creek. The collector, MargaretStorey, is no longer alive, so details of the locality were obtainable only from old aerialphotographs. Aerial photographs indicate that this area had an extensive low emergentvegetation marsh that flooded during high water flows from Amazon Creek and theLong Tom River. These specimens represent the locality that St. John (1987a) refers toas the vicinity of Junction City (St. John, pers. comm.). Ownership: Private, details ofownership unknown. USGS Topo Quads: 7.5' Junction City [1967, photorevised1986]. Elevation: Ca. 325 ft (99 m).

16) Linn County: Albany. Legal: T1OS, R3W, middle of Sec 44 [based on 7.5' USGStopo quad] or Sec 33 [based on September 1967 Metsker Atlas of Linn County].Specimens: (1) USNM 45870. Collector: D. D. Streeter, Jr. Collector's number:None. Collection date: 3 October 1909. Life stage: Juvenile. Sex: Unknown.Comments: The collector, D. D. Streeter, Jr., who worked for the Bureau of theBiological Survey, probably made the collection in the vicinity of what is now calledSecond Lake. Ownership: Presumably private, but without further data, it isimpossible to understand ownership of this locality precisely. USGS Topo Quads:7.5' Albany [1970, photorevised 1975]. Elevation: Ca. 180 ft (55 m).

17) Linn County: North end of Big Lake. Legal: T14S, R7.5E, central south 1/2 of Sec 2.Specimens: (2) PSU 1874, 1875. Collector: James J. Kirk. Collector's number. 491(1874), 492 (1875). Collection date: 4 July 1965. Life stage: Juveniles. Sex:Unknown. Comments: Kirk (pers. comm.) the north end of Big Lake has a moderate-sized, shallow, warm water marsh. Ownership: Forest Service, Willamette NationalForest, McKenzie Ranger District. USGS Topo Quads: 7.5' Three Fingered Jack[1988 provisional edition]. Elevation: Ca. 4644 ft (1415 m).

18) Linn County: Ca. 7 mi (11.2 km) east of Corvallis. Legal: TilS, R4W, Sec 34.Specimens: (1) OSUMNH 4693. Collector: Robert M. Storm. Collector's number:None. Collection date: 25 February 1957. Life stage: Adult. Sex: Male. Comments:Collection data in the OSUMNH catalogue states, "Collected 10:30 PM, in pond alongrailroad, water temperature 10.60 C." This is the last specimen of R. pretiosa thatRobert Storm (pers. comm.) remembers seeing in the Willamette Valley. This area alsoflooded when Lake Creek, a local tributary of the Calapooia River, was at high water.This locality and the Oregon Electric Railroad (see #19) are those that St. John (1987a)refers to as the Tangent area (St. John, pers. comm.). Dunlap (1955) discussed thetownship, range, and section coordinates (Ti iS, R4W, Sec34) of at least one specimenthat undoubtedly refers to this locality. Ownership: Probably private, but withoutfurther data, it is impossible to understand ownership of this locality precisely. USGSTopo Quads: 7.5' Riverside [1969, photorevised 1975]. Elevation: Ca. 220 ft (67 m).

19) Linn County: Sweet Home. Legal: T13S, RIE, locality too imprecise to provide asection. Specimens: (5) BYU 31435-9. Collector: Don R. Harris. Collector's number:None available. Collection date: 25 August 1966. Life stage: Subadults. Sex:Unknown. Comments: This may be the locality that St. John (1987a) refers to as nearCrawfordsville, but St. John (pers. comm.) could not recollect the details of thiscitation. This probably also represents the record that Gordon (1939) lists as SouthSantiam River, and which Dunlap mapped, but did not discuss. John Applegarth(pers. comm.) verified Ihe identity of these specimens as R. pretiosa. Ownership:


Probably private, but without further data, it is impossible to understand ownership ofthis locality precisely. USGS Topo Quads: 7.5' Sweet Home [1984 provisionaledition]. Elevation: Ca. 600 ft (183 m).

20) Linn County: Oregon Electric Railroad. Legal: T12S, R4W, middle of Sec 7.Specimens: (1) OSUMNH 1461. Collector: Robert M. Storm and Richard Pimentel.Collector's number: None provided. Collection date: 17 March 1950. Life stage:Adult. Sex: Female. Comments: Collection data in the OSUMNH catalogue states,"Along drainage ditch ca. 5 ft (1.5 m) wide running through a pastured area. By pondlocated northwest of railroad just southwest of point where road running from roadbetween Orleans school and 99E to Hulburt School crosses the railroad." OregonElectric Railroad on older maps is the same as the Burlington Northern Railroad ofnewer maps. Historically, when the Calapooia River was at highwater, this areaflooded. Ownership: private, specific owner unknown. USGS Topo Quads: 7.5'Riverside [1969, photorevised 1975]. Elevation: Ca. 218 ft (66 m).

21) Marion County: Aumsville. Legal: T8S, R1-2W, locality too imprecise to providesection data. Specimens: (1) MVZ 25288. Collector: Henry S. Fitch. Collector'snumber: 4158. Collection date: 13 October 1937. Life stage: Adult. Sex: Female.Comments: Henry Fitch's field notes on the date of collection were examined, but noadditional data was provided regarding this frog. Aumsville is on Mill Creek.Examination of historical aerial photographs indicate Mill Creek in the vicinity ofAumsville consisted of an extensive low, emergent marsh strongly influenced by thehigh water flows of the Santiam River a few miles to the south during high water.Ownership: Presumably private, but without further data, it is impossible to understandownership of this locality precisely. USGS Topo Quads: 7.5' Stayton [1969,photorevised 1986]. Elevation: Ca. 350 ft (107 m).

22) Multnomah County: Portland, south end of Crystal Springs Lake. Legal: T 1 S, R 1 E,north half of NW 1/4 of Sec 24 [based on 1944 Metsker Atlas of Multnomah County]or middle of Sec 49 [based on 7.5' USGS topo quad]. Specimens: (1) MVZ 197537.Collector: P. H. Pope. Collector's number: none. Date: 21 March 1922. Life stage:Adult. Sex: Female. Comments: A marsh was historically present on the west side ofCrystal Springs Lake; this marsh was destroyed with the development of the MunicipalCourse in the 1930s, and was likely the habitat that harbored the local population ofspotted frogs. Based on the Metsker Atlas of Multnomah County, the marsh west ofCrystal Springs Lake would have had a legal of T 1 S, R 1 E, SW 1/4 of Sec 13 andSE 1/4 of SE 1/4 of Sec 14.

Pope also collected some spotted frog eggs on 10 April 1922, from which he rearedand preserved two lots of larvae, one lot (MVZ 28067) was preserved on 14 April 1922and the second (MVZ 28068) was preserved on 17 May 1922. This collection likelycame from the same locality.

A series of four specimens (OSUMNH 0616, 1946; CAS 71411, UCD 3374) thatStanley G. Jewett, Jr. collected on 29 March 1930 are also from very close to thislocality; OSUMNH 0616 is the only specimen in this series with a Jewett cataloguenumber (SJ 20). All are adult females except UCD 3374, which is a juvenile ofunknown sex. The collector, Stanley G. Jewett, Jr., could not be contacted since hepassed away early in 1991. However, Doris Jewett, Stanley's widow, gavepermission to examine the catalogue of amphibian and reptile specimens Stanley hadcollected. Although he had provided no specimen number, matching the collectiondate, the catalogue indicated that frogs collected on this date had the notation "Found inpond near Crystal Creek." Historical aerial photographs indicate that Crystal SpringsLake or the marsh immediate adjacent the lake was the only significant spotted froghabitat in the Crystal Creek system and the probable site of collection. A fifth

- ---- -I


specimen, an adult female (OSUMNH 1947 [Jewett catalogue number SJ 54]) collectedon 12 September 1931 may also be from this locality, but the Jewett catalogue revealedno additional data on this collection.

Additionally, a series of 10 specimens (UCD 3361, 3584-92) that Leonard M. Scottcollected on 24 August (3584), 13 September (UCD 3585,3591), 21 September (UCD3592), 10 October (UCD 3361), and no day, no month (UCD 3586-88) 1931 mayhave come from this locality or the non-specific "sloughs along the Columbia andWillamette" in the Portland area that Jewett (1936) refers to. UCD 3361, 3586, and3591 are adult males, the remainder are females except for UCD 3587-9, which aresubadults. Scott was a close friend of Jewett's, who often went collecting with him(D. Jewett, pers. comm.).

Dunlap (1955) referred to R. pretiosa specimens from Portland that may representthese specimens, but did not provide anything to verify specimen identities. Thesespecimens are some of the records for Portland that St. John (1987a) referred to (St.John, pers. comm.). Ownership: Eastmoreland Municipal Golf Course. USGS TopoQuads: 7.5' Lake Oswego and Portland. Elevation: Ca. 50 ft (15 m).

23) Multnomah County: Portland, Johnson Creek. Legal: TlS, RlE, locality too impreciseto provide section data. Specimens: (1) USNM 36049. Collector: M. W. Lyon.Collector's number: None. Date: No day, May 1905. Life stage: Adult. Sex: Female.Comments: The collector, M. W. Lyon, worked for the Bureau of the BiologicalSurvey, but no notes were available on this collection. Johnson Creek flows westalong the southern edge of east Portland toward the Willamette River. Historical aerialphotographs indicate that a significant portion of the lower Johnson Creek drainageconsisted of marshes subject to periodic flooding. Dunlap (1955) referred to R.pretiosa specimens from Portland that may represent these specimens, but did notprovide anything to verify specimen identities. These specimens are among the oldrecords for Portland that St. John (1987a) referred to (St. John, pers. comm.).Ownership: Presumably private, but without further data, it is impossible to understandownership of this locality precisely. USGS Topo Quads: 7.5' Lake Oswego.Elevation: Ca. 60 ft (18 m).

Hayes: Spotted Frog I

Appendix IV

Unverified Localities

1) Clackamas County: Stone Creek tributary, T6S, R8E, SW 1/4 of Sec 2. Observer:Robert Penson. Collector's number: Not applicable. Date: 26 July 1991. Life stage:Adult. Sex: Probable female. Comments: Robert Penson, who made this sight record,is the Assistant District Biologist for the Bear Springs Ranger District of the Mt. HoodNational Forest (Ph: 328-621 1). This stream is a tributary of Stone Creek, which is onthe Oak Grove Fork of the Clackamas River ca. 2-3 mi (3.2-4.8 km) south of TimothyLake. Penson (pers. comm.) provided the following description. He indicated that thefrog he found was about 4 inches (ca. 100 mm) long and red orangish on itsundersurfaces. He further indicated that the coloration appeared superficial as though itwas painted on and the inky black dorsal spots had ragged edges. He described thehabitat as a shallow (3-4 inches [8-10 cm] deep), perennial creek, 4-8 ft (1.2-2.4 m)wide, with some pooling, but mostly a steady flow of water running at a slow tomoderate rate. Vegetation on the surrounding slopes at the location of the sighting wasDouglas fir (Pseudotsuga menziesii), western red cedar (Thuja plicata), westernhemlock (Tsuga heterophylla). Alders (Alnus sp.) and willows (Salix sp.) werehanging over the creek. The creek is mostly shaded with intermittent openings thatvary in size. The creek substrate is rocky with small pebbles and scattered, but fewlarger rocks. The description of the frog agrees with R. pretiosa, but the habitatdescription agrees more closely with that of R. a. aurora. The occurrence of historicalrecords for the spotted frog at nearby Clackamas Lake and Little Crater Meadows makethis record worthy of further checking. Ownership: Forest Service, Mt. Hood NationalForest, Bear Springs Ranger District. USGS Topo Quad: 7.5' Timothy Lake [1985provisional edition]. Elevation: Ca. 3,440 ft (1049 m).

2) Klamath County: vicinity of Sky Lakes Wilderness - headwaters of the South Fork ofthe Rogue River. Comments: Steve Cross (pers. comm.) mentioned having a second-hand record of red-orange ventered frogs in vicinity of the Sky Lakes Wilderness, butcould not recall the details of the sighting. Some areas in the Butte Falls Ranger Districtportion of this Wilderness or nearby may have the appropriate habitat for this species;Black Bear Swamp is one possibility that should be checked. Ownership: ForestService, Rogue River National Forest, Butte Falls Ranger District. USGS Topo Quad:7.5' Imnaha Creek [1985 provisional edition]. Elevation: 4,040 ft (1231 m) for BlackBear Swamp.

3) Lane County: Near Hidden Lake, 11 mi (17.6 km) south and 3.5 mi (5.6 km)west ofMcKenzie Bridge, T18S, R5E, Sec 8. Specimens: (1) USNM 294912. Collector:Douglas W. Smith. Collector's number: USNM-FH 133321. Date: 22 August 1983.Life stage: Adult. Sex: Female. Comments: This is a specimen with somewhatpeculiar spotting reminiscent of the yellow-ventered R. pretiosa in eastern Oregon, butits leg dimensions and its foot webbing agree with R. cascadae. Thus, it is either a R.cascadae with peculiar spotting or a hybrid. Although the potential hybrid status of thisspecimen can ultimately only be verifed genetically, one can have a high degree ofconfidence that it is not a hybrid if it agrees in its morphometrics with the Cascade frog(D. Green, pers. comm.). A tiny (ca. 2 ac) bog exists along the north side of HiddenLake. Ownership: Forest Service, Willamette National Forest, Blue River RangerDistrict. USGS Topo Quad: 7.5' Cougar Reservoir [1989 provisional edition].Elevation: 3,285 ft (1001 m).

Hayes: Spotted Frog 2.

4) Linn County: Jorn Lake. Comments: Evenden (1943) listed this locality as having bothsubspecies of the spotted frog, namely, the western spotted frog (Rana pretiosapretiosa) and the Nevada spotted frog (Rana pretiosa luteiventris). Evenden indicatesthat he referred to Gordon (1939) for identification of various species. But becauseGordon did not recognize the Cascades frog, and the Nevada spotted frog wasdescribed as having yellow undersurfaces essentially identical to the Cascades frog,Evenden's observations of R. p.luteiventris are referable to the Cascades frog. Dunlapand Storm (1951) empasized this point, stating numerous literature records of R. p.luteiventris in lakes and streams of the high Cascades undoubtedly refer to Ranacascadae. These authors reported records of Rana cascadae, including those for LakeJorn, for which several verifiable specimens exist [OSUMNH 5407-16; n = 10).However, Evenden (1943) also reported taking two specimens of Rana pretiosapretiosa in the grasses at the edge of Jorn Lake, but the specimens that are the basis ofthis record have not been found. If this represents a valid record, it would be thehighest elevation at which R. pretiosa had been recorded west of the Cascade crestOwnership: Forest Service, Willamette National Forest, Detroit Ranger District. USGSTopo Quad: 7.5' Marion Lake [1988 provisional edition]. Elevation: 5,070 ft(1545 m).

5) Jackson County: Vicinity of Union Creek. Comment: St. John (1984a) commented onrecords from this locality. This is near the Whiskey Creek locality reported by Fitch(1936) and represented by specimens of R. cascadae (see Appendix V). Steve Cross(pers. comm.) indicated that a spotted frog specimen from this vicinity used to be in thecollection at Southern Oregon State College (Ashland), but the specimen was lost.However, Cross indicated he had seen red-orange ventered frogs from this vicinity thathe considered spotted frogs. A Carnegie Museum specimen from along the RogueRiver near Union Creek is a R. cascadae [CM 32802]. The Thousand Springs area onthe Jackson-Klamath county line [T 31 S, R4 E, SE 1/4 of Sec 12 and NE 1/4 of 13;and T31S, R5E, SW 1/4 of Sec 7 and NW 1/4 of Sec 8] should be checked forpotentially suitable habitat, but the elevation suggests suitability may be borderline.Ownership: Forest Service, Prospect Ranger District, Rogue River National Forest.USGS Topo Quad: 7.5' Thousand Springs [1988 provisional edition]. Elevation:4,920-5,000 ft (1500-1524 m).

6) Multnomah County: Marsh in vicinity of Big Bend Mountain. Comment: This record isin the Oregon Natural Heritage Database, and is based on the sight record of KirkHorn, formerly Wildlife Biologist for the Mt. Hood National Forest. The locality ofthe 1974 sighting is a small marsh surrounding a small lake ca. 1 mi SSE of Big BendMountain (K. Horn, pers. comm.). The habitat and elevation agree with the possibilitythat this is a valid spotted frog record. Horn indicated that he had discussed thepossibility of this locality harboring spotted frogs with Ronald Nussbaum and RobertStorm, both of whom indicated they considered it a reasonable possibility, but neitherof whom ever had the opportunity to verify their occurrence there. Ownership: ForestService, Mt. Hood National Forest, Zigzag Ranger District. USGS Topo Quad: 7.5'Hickman Butte [1962, photorevised 1980]. Elevation: 3,084 feet (940 in).


Appendix V

Erroneous Localities

1) Clatsop County: Gearhart. Comments: Slevin (1928) listed this record, which Gordon(1939) and Dunlap (1955) accepted without examining associated specimens (e.g., seeFigure 1 in Dunlap). Only northern red-legged frogs (R. a. aurora) have beencollected from this locality [CAS 29858-9; n = 2].

2) Coos County: Myrtle Point. Comments: Slevin (1928) reported this locality and hiscomment has been the basis of quotes by others (Gordon 1939, St. John 1985)without examination of specimens (e.g., see list of literature records accepted byDunlap (1955) in his Figure 1). This record is also listed in ONHD. The recordSlevin reported is based on five specimens [CAS-SU 1224, CAS 4988-91] JamesOtterbein Snyder and a colleague collected in 1897 and 1899, and identified simply asRana sp. in the Academy catalogues. Four of the specimens [CAS 4988-9 1] are notedin the catalogue as having been lost in the earthquake and fire of 1906; the fifthspecimen is apparently also lost although no notation indicating such exists. The fifthspecimen noted as Rana sp. indicates that, like the others, it was among the specimensthat John Van Denburgh, then Curator at the Academy, was holding until he couldexamine them for identification, but which were lost in the fire of 1906 (J. Vindurn,pers. comm.). Slevin had to identify these specimens as Rana pretiosa after they werelost since Van Denburgh hired him at the Academy to assist with collecting in 1905and he did not become experienced in identification of material until well after 1906(M. Jennings, pers. comm.). Slevin, at least partly dependent on the notes VanDenburgh had taken on specimens lost in 1906 for the material he treated in his 1928opus, likely lacked notes on these specimens because other unidentified material in thecatalogue listed as lost in 1906 represented material that Van Denburgh never had theopportunity to identify before it was lost. In addition, Slevin had taken a large seriesof northern red-legged frogs (Rana aurora) at Myrtle Point on 9-11 June 1911,specimens which still exist [CAS 29466-91; n = 26]. Slevin likely interpreted theunidentified Rana specimens that Van Denburgh left as something other than Ranaaurora, perhaps assuming that Van Denburgh had actually gotten a chance to comparethem to red-legged frogs, and that Van Denburgh had concluded they were different,but did not get a chance to finalize his identification before they were lost in 1906.However, no reason exists not to believe that these five specimens had simply beensitting until Van Denburgh would have had the chance to examine them and were lostbefore he ever examined them at all. Based on the locality, Rana pretiosa was the onlyalternative Rana that these specimens could reasonably be had they not been R. aurora,so this was probably the guess that Slevin made. However, it is very unlikely that theseries of lost specimens is R. pretiosa because R. aurora is the only spotted ranid frogthat either Slevin or anyone else collected in that region, and for the ecological andbiogeographic reasons provided earlier, it is unlikely that R. pretiosa would haveoccurred either so near the coast or in this general region of southwestern Oregon. Inthe absence of material upon which this collection can be verified, it is consistent toregard it as an erroneous record.

3) Columbia County: Nehalem River drainage, vicinity of Vernonia. Ownership: Private,specific ownership unknown. Comments: This record is in the Oregon NaturalHeritage Database and is based on a sight record. Jim Merzenich (pers. comm.), theindividual who made the sight record, indicated that he observed a frog on land in awell-shaded copse of alders with an understory of lady fern and sword fern along asmall tributary stream of the Nehalem in the vicinity of Vemonia in late August of1976. The adjacent stream was described as clear, cold, 2 meters wide and 5

"I 11 -_- __1 -- I----- � - - -11 '


centimeters deep. Further discussion with Merzenich, a botanist, revealed that areasonable probability existed that the frog he had observed was a northern red-leggedfrog. His description of the frog and the well-shaded, cold stream habitat make itlikely that a northern red-legged frog, rather than a spotted frog, was observed. PhilGaddis (pers. comm.) field checked this locality in 1982, but was uncertain whetherhe relocated Merzenich's original locality. Nevertheless, Gaddis was unable to locateany spotted frogs. The relatively coastal position of this locality make it unlikely forthe spotted frog. In the absence of material upon which this collection can be verified,it is consistent to regard it as an erroneous record.

4) Curry County: 11 mi (17.6 km) above mouth of Rogue River. Comments: This recordis based on a single specimen (KU 50362) Henry S. Fitch (his number 2446) collectedon 20 May 1935; this specimen is a subadult northern red-legged frog (R. a. aurora).

5) Douglas County: Diamond Lake. Comments: This locality is represented by a series ofspecimens from the University of Kansas (KU 23545-8, 23550-8; n = 13) collected byDSF (collector's number 410701-2) on 1 July 1941; the entire series consists of R.cascadae.

6) Jackson County: Whiskey Creek. Comments: This record is based on workers(Dunlap 1955, St. John 1984a) quoting the original record of Fitch (1936) withoutreexamination of specimens themselves (e.g., see list of literature records accepted byDunlap (1955) in his Figure 1). The Fitch record consists of two specimens of Ranacascadae [MVZ 17185-6], but was published prior to the description of the latter (seeSlater 1939). Other specimens recorded from Whiskey Creek are also Rana cascadae[KU 192060-3].

7) Klamath County: all localities within Crater Lake National Park. Ownership: NationalPark Service. Comments: Rana cascadae is represented at all known localities forwhich specimens exist within Crater Lake National Park (contra Dunlap 1955; see alsoDunlap and Storm 1951); although the Union Peak-Red Blanket Motorway Junction[CAS-SU 3852-5] specimens are spotted in a fashion similar to the peculiar HiddenLake specimen (see #3 in Appendix IV). No specimens of Rana pretiosa have everbeen collected within Crater Lake National Park, and the large majority of the Parkbeing over 5,000 ft (Farner and Kezer 1953), most habitat there is likely too high forthis species. Rana cascadae is documented at the following localities by one or morespecimens: Crater Lake, no other specifics [OSUMNH 610-612, KU 128084-6];Boundary Spring, west-northwest Park Boundary [CAS-SU 3865]; Crater Spring [KU30219, 128079-83, 206431]; Government Headquarters [CAS-SU 9940-3]; MunsonSpring [KU 23544]; Munson Meadows [CAS-SU 3863-4, KU 23549, MVZ 50264-5]; Munson Valley, 0.5 mi (0.8 km) Government Camp [MVZ 17606]; South Fork ofCastle Creek [CAS-SU 3856]; and Videa Falls [CM 32792].

8) Lane County: Mud Lake, Mink Lake Basin. Comments: This locality represents alapsus in recording the correct Mud Lake. The Mud Lake with R. pretiosa [OSUMNH1835-40, 1845-7, 5152-65, 8340-1, 8360-1; n = 27) is located in Deschutes Countyand on current maps is known as Hosmer Lake. A different Mud Lake does exist in theMink Lake Basin, but only R. cascadae have been recorded at the latter, the topographyand elevation of which make it unlikely that spotted frogs would occur there.

9) Linn County: Blue Lake. Comments: Evenden (1943) listed this locality as having theNevada subspecies of the spotted frog, namely Rana pretiosa luteiventris. Evendenindicates that he referred to Gordon (1939) for identification of various species. Butbecause Gordon did not recognize the Cascades frog, and the Nevada spotted frog was


described as having yellow undersurfaces similar to the Cascades frog, Evendenobservations are referable to the Cascades frog, particularly since he discussesobserving the red-orange ventered subspecies (Rana pretiosa pretiosa) at nearby LakeJorn. Dunlap and Storm (1951) emphasized this point, stating that numerous literaturerecords of Rana pretiosa luteiventris in lakes and streams of the high Cascadesundoubtedly refer to Rana cascadae.

10) Linn County: Bowerman Lake. Comments: Evenden (1943) listed this locality ashaving the Nevada subspecies of the spotted frog, namely Rana pretiosa luteiventris.The same comments apply as for the Blue Lake locality (see #9).

11) Linn County: Red Butte Lake. Comments: Evenden (1943) listed this locality ashaving the Nevada subspecies of the spotted frog, namely Rana pretiosa luteiventris.The same comments apply as for the Blue Lake locality (see #9).

12) Linn County: Santiam Lake. Comments: Evenden (1943) listed this locality as havingthe Nevada subspecies of the spotted frog, namely Rana pretiosa luteiventris. Thesame comments apply as for the Blue Lake locality (see #9).

13) Linn County: Along South Santiam River, 100 yds (ca. 90 m) below Coyote Creek.Comments: Virgil Morris (Ph: 367-5168), Biologist for the Sweet Home RangerDistrict of the Willamette National Forest, indicated that Jim Gillette (Ph: 484-0517) ofEugene made this sight record. Gillette (pers. comm.) indicated that a well-usedswimming hole is located at this locality (T13S, R3E, Sec 32 or 33). A frog that fromhis description sounds more like a red-legged frog than a spotted frog was found inshallow riffles at the lower end of the pool which is the swimming hole. I visited thislocality in late February 1992 and found three northern red-legged frogs (Rana aurora)in the vicinity of the swimming hole. The edge habitat is quite shaded and habitatconditions are not what one would expect as suitable for the spotted frog. Based on thehabitat, the description of the frog by Gillette, and the red-legged frogs observed at thesite by myself, this record should be considered erroneous.

14) Tillamook County: Garibaldi. Comment: Slevin (1928) listed this record, whichGordon (1939) and Dunlap (1955) accepted without examining associated specimens(e.g., see Figure 1 in Dunlap). Only the northern red-legged frog (R. a. aurora) hasbeen collected from this locality [CAS 29732; n = 1]

15) Washington County: Gales Creek. Comments: St. John (1987a) referred to old R.pretiosa specimens from this locality, but he could not recall the source of thoserecords, but thought they might be based on a comment that Robert Storm made (A. St.John, pers. comm.). Storm knew of no R. pretiosa records from the Gales Creek area,although he mentioned that R. a. aurora had been recorded from that vicinity severaltimes. Further, no specimens or other literature that were examined refer to records ofR. pretiosa from this site, although specimens of the northern red-legged frog (R. a.aurora) are known from this area [MVZ 61764-5; n = 2]. The relatively coastalposition of this locality makes it doubtful for R. pretiosa, and this record should beconsidered erroneous.

16) Yarnhill County: McMinnville, vicinity of 835 Ashwood. Comment: This record wasthe McMinnville record of St. John (1987a). Alan St. John (pers. comm.) indicatedthat the basis of this record was a photograph that naturalist friends, the Fenders, had intheir living room, which he saw when he was 11 years old (ca. 1959). Dorothy Fender(still at 835 Ashwood, McMinnville; Ph: 472-6905) remembered the photograph, butinsisted it was a red-legged frog, and accurately described that species. Fender also


indicated that she had sent the photograph to Robert Storm for verification, but wasunable to relocate it. Without further data, this record should be considered erroneous.

CURRENT STATUS OF THE SPOTTED FROG (Rana pretiosa)

IN WESTERN OREGON

Marc P. Hayes

Research Associate

Department of Biology, Portland State University,


and

Research Section, Animal Management Division, Metro Washington Park Zoo,


Abstract: Survey of localities where the spotted frog (Rana pretiosa) previously occurred

in western Oregon, a taxon that has been characterized as distinctive (the western spotted

frog), indicates that, conservatively, it has disappeared from over 90% of its range in this

region. In addition to the two historical localities when the western spotted frog is extant,

one previously unrecorded locality for this taxon was found in western Oregon. Localities

where the western spotted frog is extant are all over 1219 m (4,000 ft) in elevation, and are

located in areas where hydrological modification and exotic macropredators, the two factors

believed to be most responsible for the disappearance of this taxon at low elevations, are

reduced. Nevertheless, prognosis for two of the three remaining populations is poor, as

the size structure of those populations samples suggests recruitment failure or high adult

mortality. The severe reduction of its range and the high vulnerability of remaining western

Oregon populations strongly supports the recommendation that the western spotted frog

should be listed as Endangered in the state.

CURRENT STATUS OF THE SPOTTED FROG (Rana pretiosa)

IN WESTERN OREGONa

Marc P. Hayes

Research Associate

Department of Biology, Portland State University,


and

Research Section, Animal Management Division, Metro Washington Park Zoo,


Introduction

Previous work has detailed the historic distribution of the spotted frog (Rana pretiosa)

in western Oregon; indicated that the spotted frog in western Oregon is a morphologically

identifiable form to which the label, western spotted frog has been applied; and suggested a

link between the western spotted frog and warmwater marsh habitats (Hayes 1994). Yet,

despite numerous suggestions that this frog has declined (St. John 1984a, 1985a, 1987a,

1987b; Storm 1986; Marshall 1989; McAllister and Leonard 1990, 1991; Leonard et al.

1993; D. Green, R. Storm, pers. comm.), systematically collected data needed to verify

that possibility in western Oregon were lacking. Specifically, verification required that

historical localities be examined to determine over what proportion of its historic range the

western spotted frog still occurs, the pivotal recommendation of previous work (Hayes

1994). Moreover, verification that warmwater marshes were the active-season habitat

needed verification. This report summarizes the work that provides verification for both.

Method

Every verifiable locality for the western spotted frog in western Oregon was examined

(Table 1; data for each locality are provided elsewhere; see Appendix I of Hayes (1994)).

Except for Goose Lake, each locality was visited three times. Where the historical locality

could be precisely identified (n = 14; Localities 1, 3, 4, 5, 10, 11, 12, 13, 14, 16, 17, 18,

a In the context of this report, western Oregon refers to Oregon west of the crest of the Cascade Mountainsand the portion of the Klamath hydrographic basin that lies within the state.

Table 1 - Sites Checked Associated With Historical Spotted Frog Localities

Locality (County) Locality (County)

1) McFadden's Marsh (Benton)

2) Corvallis (Benton)

a) Hewlett-Packard Pond

b) Aggregate plant ponds

c) Oak Creek Marsh near 53rd Street

d) Near Marys River, south of Highway 20

3) Clackamas Lake (Clackamas)

4) Little Crater Meadows (Clackamas)

5) Little Hyatt Reservoir (Jackson)

6) Fort Klamath (Klamath)

a) Wood River Ranch

b) Wood River at Highway 62

c) Wood River at Loosley Road

d) North-to-east turn in Weed Road

e) Wood River at Weed Road

7) Mountains near Fort Klamath (Klamath)

a) Sevenmile Creek Marsh

b) Jack Spring (Fourmile Creek)

c) Mares Egg Spring outflow

8) Klamath Falls (Klamath)

a) Marshy edge of Lake Ewauna

b) Marshy area along Link River

c) Klamath Game Management Area

9) Klamath Lake (Klamath)

a) Hank's Marsh

b) Marshy area near Modoc Point

c) Marshy area near Pelican City

10) Sprague River at Spring Creek (Klamath)

11) North fork of Sprague River (Klamath)

12) Sprague River near Ferguson Mountain (Klamath

13) Gold Lake Bog (Lane)

14) Goose Lake (Lane)

15) Southwest Junction City (Lane)

a) Flat Creek junction with Highway 36

b) Amazon Creek junction with Highway 36

c) Amazon Creek arm junction with Highway 36

16) Albany, Second Lake (Linn)

17) North end of Big Lake (Linn)

18) Tangent, Oregon Electric Railroad (Linn)

19) Sweet Home (Linn)

a) Marshy area near Crawfordsville

b) Ponds near Santiam River

c) Marshy area near Holley

20) Oregon Electric Railroad (Linn)

21) Aumsville (Marion)

a) Beaver Creek at Highway 22

b) Marshy area south of Mill Creek Road

c) Mill Creek near Golf Club Road

22) Southwest of Crystal Springs Lake (Multnomah)

23) Johnson Creek (Mulmnomah)

a) Near Lara Drive west of Southeast 17th Avenue

b) Near junction with Crystal Springs Creek

c) Near Johnson Creek Boulevard crossing

d) Near Stanley Avenue crossing

e) Near Linwood crossing

f) Near Bell Avenue crossing

Hayes: Spotted Frog Status in Western Oregon 2

20, and 22; Table 1), each site visit consisted of a search lasting at least two hours. At sites

where the historical locality could not be precisely determined (n = 9; localities 2, 6, 7, 8,

9, 15, 19, 21, and 23; Table 1), 3-6 sites with characteristics thought to be most favorable

for the western spotted frog (see Hayes 1994) were examined in the general vicinity of each

historical locality. Searches of the latter varied in length from 20 minutes to three hours,

depending on site size. One to four individuals participated in the searches. Prior to a site

search, available aerial photographs and topographic maps were examined in detail to help

identify the areas of a site most likely to have suitable habitat, often with the assistance of

agency biologists familiar with the area. All searches were conducted during the day under

favorable climatic conditions, at least partial sun with minimal wind to allow surface water

to warm. At each locality, data were recorded on the number and species of amphibians

present, associated animal species observed, and the physical and vegetational structure of

the habitat. and climatic conditions at the time of the site reconnaissance. In many cases,

agency biologists assisted in the onsite identification of fishes through some sort of

sampling, electroshocking or netting; individuals who assisted in the latter are listed in the

acknowledgments. Where it was not possible to coordinate onsite sampling with agency

biologists, locality-specific historic or current data on fishes and other aquatic fauna were

retrieved from available sources. If the western spotted frog was encountered, as many

individuals as possible were captured, processed, and subsequently released to provide

data on population structure. For those individuals that could not be captured, size was

estimated wherever possible in order to assist the population structure analysis. Data taken

included a body size measurement or estimate, a determination of sex or estimate, and a

measurement of mass. Shank lengths (fibulo-tibia) were also generally taken. Particular

effort was made to take shank length measurements if any ambiguity existed as to whether

the frogs captured were really western spotted frogs (see Hayes 1994).

Unverified localities (i.e., localities where spotted frogs had been claimed to occur, but

for which no data existed upon which to either confirm or reject this claim; see Hayes

1994) were visited at least twice; some were visited three times (Table 2a). All unverified

localities had precise locality data, so the pattern of searching was not altered as was done

for some of the historical localities. Data recorded at unverified localities was identical to

that recorded for verifiable localities. One unverified locality (see 2 in Table 2a) was not

visited because the specifics of the locality record were not retrieved until after the field

season was over.

Table 2a - Unverified Spotted Frog Locality Site Checks

Site Visits

Locality (County) 1 2 3 OFa Bfb FishesC

1) Stone Creek Tributary (Clackamas) 0 0 0 T 0 CuUs*

2) Sky Lakes Wilderness (Klamath) - - - - - -

3) Hidden Lake (Lane) 0 0 0 CP 0 ?

4) Jorn Lake (Linn) 0 0 0 C,P 0 Br*

5) Union Creek (Jackson) 0 0 0 CP 0 Br,Cu*

6) Big Bend Mountain (Multnomah) 0 0 - CP 0 0

Table 2b - Erroneous Spotted Frog Locality Site Checks

Site Visits

Locality (County) 1 2 3 OFa Bfb Fishesc

a) Vicinity of Vernonia (Columbia) 0 0 - N,P 0 CuUs*

b) South Santiam River near Coyote Creek (Linn) 0 0 - N,P 0 ?

The code + (Present) refers to detections of the western spotted frog, whereas 0 (Not found) refers to notfinding the indicated species category. A (-) indicates the listed visit was not made. Under the othercolumns, a (?) indicates one or more fish species were observed, but not identified, and an (*) indicatesthat the indicated fishes are probably only a subset of the species present.

a OF = Other native frogs; codes are: C = Cascades frog (Rana cascadae), N = Northern red-legged frog (Ranaaurora aurora), P = Pacific (treefrog) chorusfrog (Pseudacris (=Hyla) regilla), and T = Tailed frog(Ascaphus truei).

b Bf = Bullfrog (Rana catesbeiana).c Fishes codes: Br = Brook trout (Salvelinus confluentus), Cu = Cutthroat trout (Oncorhynchus clarki), and

Us = Unidentified sculpin (Cottus sp.).


Erroneous localities (i.e., localities where the western spotted frog was claimed to

occur, but where that claim was based on misidentified specimens, a misinterpretation

associated with the original specimens, or other data that suggested the localities were

erroneous; see Hayes 1994) were visited only if an a priori reason existed that the absence

of the western spotted frog could not be eliminated without at least one site visit. A priori

reasons for visiting such localities included: 1) knowledge of the habitat present could not

exclude the possibility that the western spotted frog was absent; or 2) absent of knowledge

of the habitat present could not eliminate the possibility that the western spotted frog was

absent. Two such localities were each visited twice (see a,b in Table 2b).

A few additional localities in western Oregon not falling into one of the three above-

mentioned categories were visited because they matched the habitat characteristics of those

localities known to harbor western spotted frogs (Appendix I). Because of time constraints

and the relatively short summer season in 1993, three of these localities were not revisited.

Because so few localities with western spotted frogs were found in western Oregon, a few

reference localities known to harbor western spotted frogs from outside the study area were

also visited (see Appendix II). Data from these localities was used in analyses designed to

validate the search method and verify the postulated link between the western spotted frog

and warmwater marsh habitats (see Hayes 1994). For all such localities, data were

recorded as previously described. In addition, the length of the search interval up to the

point the first western spotted frog was found was recorded. A few of the data presented

here were collected during 1992, but most were collected during 1993.

Results

Western spotted frogs were encountered at two of the 23 localities (9%) from which

they were historically recorded (Table 3; Figure 1). Because nine localities were imprecise,

the actual number of localities searched to have a high degree of confidence that the sites

from which western spotted frogs were historical taken were actually covered during these

searches was 47. Among these localities, the western spotted frog was found at only in the

vicinity of Gold Lake (Willamette River basin) and on the Wood River Ranch (Klamath

River basin). The Wood River Ranch locality was conservatively assumed to represent the

historical locality identified as Fort Klamath (6a in Tables 1 and 3). If this assumption is

incorrect, this locality would represent a new record and the western spotted frog would

have been encountered at only one of the 23 localities (4%) at which they were historically

Table 3 - Historical Spotted Frog Site Checks

Site Visits


1) McFadden's Marsh 0 0 0 N,P + C,BL,M,Us*

2) Corvallisa) Hewlett-Packard Pond 0 0 0 P + BL,M*b) Aggregate plant ponds 0 0 0 P + C,B,L,M,Us*c) Oak Crk Marsh near 53rd Street 0 0 0 P 0 M,Us*d) Near Marys River, south Highway 20 0 0 0 P + Us*

3) Clackamas Lake 0 0 0 C,P 0 Br,Cu*

4) Little Crater Meadows 0 0 0 C,P 0 Cu*

5) Little Hyatt Reservoir 0 0 0 P 0 B,B1,Br,_Q,L

6) Fort Klamatha) Wood River Ranch + + + P 0 Bc,F*b) Wood River at Highway 62 0 0 0 P 0 Rt,Us*c) Wood River at Loosley Road 0 0 0 P 0 Rt,Tc*d) North-to-east turn in Weed Road 0 0 0 0 0 0e) Wood River at Weed Road 0 0 0 P 0 Rt,Tc*

7) Mountains near Fort Klamatha) Sevenmmile Creek Marsh 0 0 0 C,P 0 0b) Jack Spring (Fourmile Creek) 0 0 0 P 0 Rt,Us*c) Mares Egg Spring outflow 0 0 0 P 0 Rt,Us*

8) Klamath Fallsa) Marshy edge of Lake Ewauna 0 0 0 P + B, BI, L,M,Tc*b) Marshy area along Link River 0 0 0 P + L,M,Ucy*c) Klamath Game Management Area 0 0 0 P + Bl,GrL

10) Sprague River at Spring Creek 0 0 0 P + BcTc*


a OF = Other native frogs or toads; codes are: C = Cascades frog (Rana cascadae), N = Northern red-leggedfrog (Rana aurora aurora), and P = Pacific (treefrog) chorusfrog (Pseudacris regilla).

b Bf = Bullfrog (Rana catesbeiana).c Fishes codes: B = Brown bullhead (Ameiurus nebulosus), Bc = Blue chub (Gila coerulea), Bl = Bluegill

(Lepomis macrochirus), Br = Brook trout (Salvelinus confluentus), C = Carp (Cyprinus carpio), Cu =Cutthroat trout (Oncorhynchus clarki), F = Fathead minnow (Pimephales promelas), L = Largemouthbass (Micropterus salmoides), M = Mosquitofish (Gambusia affinis), Rt = Rainbow trout(Oncorhynchus mykiss), Tc = Tui chub (Gila bicolor), and Us = Unidentified sculpin (Cottus sp.).Underlined abbreviation for a fish species indicates data on the presence of this species was obtainedfrom a source other than direct observation.

Table 3 - Historical Spotted Frog Site Checks (continued)

Site Visits


11) North fork of Sprague River 0 0 0 P + Tc*.

12) Sprague River near Ferguson Mountain 0 0 0 P + BcTc*

13) Gold Lake Bog + + + CPW 0 Br,Cu,Rt

14) Goose Lake 0 0 - CP 0 Br

15) SW Junction Citya) Flat Creek junction Highway 36 0 0 0 0 + CUcy*b) Amazon Creek junction Highway 36 0 0 0 P + G*c) Amazon Creek arm at Highway 36 0 0 0 N,P + M,Gr*

16) Albany, Second Lake 0 0 0 0 + L,M*

17) North end of Big Lake 0 0 0 C,P,W 0 Br,CuK

18) Tangent, Oregon Electric Railroad 0 0 0 P + M

19) Sweet Homea) Marshy area near Crawfordsville 0 0 0 P + MUs*b) Ponds near Santiam River 0 0 0 0 + B,B1,L*c) Marshy area near Holley 0 0 0 P + Bl,M*

20) Oregon Electric Railroad 0 0 0 P 0 0

21) Aumsvillea) Beaver Creek at Hwy 22 0 0 0 P + CuM*b) Marshy area s of Mill Creek Road 0 0 0 0 + ?c) Mill Creek near Golf Club Road 0 0 0 P 0 M,Us*


a OF = Other native frogs or toads; codes are: C = Cascades frog (Rana cascadae), N = Northern red-leggedfrog (Rana aurora aurora), P = Pacific (treefrog) chorusfrog (Pseudacris regilla), and W = Western toad(Bufo boreas).

b Bf = Bullfrog (Rana catesbeiana).c Fishes codes: B = Brown bullhead (Ameiurus nebulosus), Bc = Blue chub (Gila coerulea), Bl = Bluegill

(Lepomis macrochirus), Br = Brook trout (Salvelinusfontinalis), C = Carp (Cyprinus carpio), Cu =Cutthroat trout (Oncorhynchus clarki), G = Goldfish (Carassius auratus), Gr = Green sunfish (Lepomiscyanellus), K = Kokanee (Oncorhynchus kisutch), L = Largemouth bass (Micropterus salmoides), M =Mosquitofish (Gambusia affinis), Tc = Tui chub (Gila bicolor), Ucy = Unidentified cyprinid, and Us =Unidentified sculpin (Cottus sp.). Underlined abbreviation for a fish species indicates data on thepresence of this species was obtained from a source other than direct observation.

---

Table 3 - Historical Spotted Frog Site Checks (continued)

Site Visits


22) Southwest Crystal Springs Lake 0 0 0 P + C,Ls,Rs,Ts

23) Johnson Creek (Multnomah)a) Near Lara Drive west of 17th Avenue 0 0 0 0 0 Ls,Rs*b) Near juncttion Crystal Springs Creek 0 0 0 0 0 SSUs*c) Near Johnson Creek Boulevard crossing 0 0 0 0 0 ?d) Near Stanley Avenue crossing 0 0 0 0 0 ?e) Near Linwood Avenue crossing 0 0 0 0 0 Ts,Us*f) Near Bell Avenue crossing 0 0 0 0 0 Ch,Re,Rs,Us*


a OF = Other native frogs or toads; code is: P = Pacific (treefrog) chorusfrog (Pseudacris regilla).b Bf = Bullfrog (Rana catesbeiana).c Fishes codes: Ch = Chiselmouth (Acrocheilus alutaceus), Ls = Largescale suckers (Catostomus

macrocheilus), Re = Reticulate sculpin (Cottus perplexus), Rs = Redside shiners (Richardsoniusbalteatus),S = Silver salmon (Oncorhynchus kisutch), St = Steelhead (Oncorhynchus mykiss), Ts =Threespine stickleback (Gasterosteus aculeatus), and Us = Unidentified sculpin (Cottus sp.).

1250 124° 1230 1220 1210

_il t Km - 460

S

Localities

* Extant 345

0 Probably Extinct 0

, 17<~~2 17-9

440

0~~ > 149\"} L - 44°

13-.-

1~~~~ ~430

6~~~~

5~-o 9

8 --_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 4 2 0

C M P H~y. 1992

Dashed line indicates the eastern limit of western Oregon as described in the text.

1250 1240 1230 1220 1210

Figure 1. The historic and current distribution of the spotted frog (Rana pretiosa) in western Oregon.Numbers for each locality correspond to the lists in Tables 1 and 3. Solid lines are county boundariesexcept that the dashed line representing the eastern limit of western Oregon follows county boundariesnorth of the vicinity of Crater Lake National Park. The dot labeled A is the new Penn Lake locality.


found. Exotic warmwater macropredators (bullfrogs and various fishes) were encountered

15 of the 23 localities at which the western spotted frog was historically found, and neither

were recorded from localities where the western spotted frog was currently found (Table

3). One locality where the western spotted frog had been historically recorded no longer

has significant aquatic habitat present (20 in Tables 1 and 3). The Cascades frog (Rana

cascadae) and the northern red-legged frog (Rana aurora aurora) were found at six and two

localities, respectively, during these searches. More than one species of ranid frog was

found at only a single locality, Gold Lake; both the Cascades frog and the western spotted

frog were present there.

No western spotted frogs were encountered at five of the six unverified localities at

which they had been claimed to have been found (Table 2). The Cascades frog was found

at four of these unverified localities, and the tailed frog (Ascaphus truei) was found at one

of the unverified localities.

No western spotted frogs were encountered at either of the two erroneous localities

visited, but northern red-legged frogs were observed at both localities (Table 2).

If the Wood River Ranch locality (6a in Table 3) is considered to be a previously

recorded locality for the western spotted frog, the western spotted frog was found at one

previously unrecorded locality in western Oregon (Penn Lake, Three Sisters Wilderness,

Willamette River basin; see Appendix I).

Of the three localities where the western spotted frog is known to be extant in western

Oregon, only one (Gold Lake) may harbor a significant population (13 in Table 3) that has

a population structure with all size groups represented (Table 4). Despite the fact that all

three localities at which the western spotted frog was found within the study area lack

exotic warmwater macropredators (Table 3 and pers. observ.), some exotic fishes were

present at each locality. Brook trout (Salvelinusfontinalis) were common in Gold Lake,

and occur in the deeper water of its associated bog, where most of the western spotted

frogs that occur there are found. Brook trout were also abundant in Penn Lake. Although

the fish fauna of the Wood River Ranch locality remains incompletely understood,

apparently at least fathead minnows (Pimephelas promelas) are common there. Notably,

Gold Lake Bog is the only one of the three, that because of large areas of isolated shallow

water, appears to offer significant refuge from fish predators. All three localities are similar

in that each had a development of warm water (consistently over 20'C) during the summer

in a bog or marsh (albeit modified in the case of Wood River Ranch) that is located in the

upper portions of their respective stream systems. The Gold and Penn Lakes localities are

Table 4 - Western Spotted Frog Numbers at Sampled Sites

Site Visit

1 2 3

Locality (County) n f/hr n f/hr n f/hr

1) Trout Lake marsh (Klickitat) 1 6.0 11 4.8 9 3.6

2) Conboy NWR (Klickitat) 8 10.7 - - - -

3) Big Marsh (Klamath) 33 16.5 35 15.4 27 13.7

4) Davis Lake, Odell Crk inflow (Klamath) 1 0.4 2 0.6 - -

5) Odell Crk above NFD 4660 (Klamath) 3 1.5 4 1.7 - -

6) Wood River Ranch (Klamath) 1 2.0 3 1.5 5 2.2

7) Gold Lake Bog (Lane) 113 22.6 153 25.5 101 27.3

8) Penn Lake (Lane) 11 2.4 - - - -

Under the site visits (I1-3), n is the number of western spotted frogs observed and f/hr is the number ofwestern spotted frogs observed per hour of search time.

Structure of Spotted Frog Populations in Western Oregon

Size Class (Snout-urostyle length (SUL) in mm)

Locality Visit# <35 mm 36-50 mm 51-65 mm 66-80 mm >80 mm

1) Wood River Ranch 2 0 1 2 0 0

3 0 3 2 0 0

2) Gold Lake Bog 1 2 44 56 11 0

2 18 48 72 15 0

3 23 36 30 12 0

3) Penn Lake 1 0 1 1 5 4

Number of frogs in each sample in the indicated size classes is shown.

Hayes: Spotted Frog Status in Western Oregon 5s

the uppermost or next to the uppermost in their respective systems. All three localities are

also similar in that the bog or marsh system is at least four hectares in size. In their

minimum size and in their warmwater condition during summer months, these localities

parallel the five reference sites where western spotted frogs were found outside the study

area (Appendix II).

For all localities sampled at least twice where western spotted frogs were encountered,

western spotted frogs were found on every site visit. At no site were western spotted frogs

found after the first visit if they had not be found on the first visit; similarly, at no site were

western spotted frogs found on the first site visit, but then not found on subsequent visits.

At each site visited where western spotted frogs were found, the first western spotted frog

was encountered within the first 18 minutes onsite, and often much sooner (Table 5).

Conclusions

The precise number of historical localities for the western spotted frog in western

Oregon may be somewhat vague because of the way some historical localities were

recorded, but even the most conservative estimate, which treats each historical record from

the same vicinity as one locality and potentially new localities as historical ones if they

occur within an extended "vicinity", leads to the conclusion that the species has disappeared

from over 90% of its historic range in this region. No western spotted frogs were

encountered at any of the 10 historical localities (represented by a search of 27 localities;

see Table 3) in the lowland Willamette Valley, which supports previous claims that this

taxon is extirpated there. Extirpation is based on negative evidence, so this finding should

especially not be used to discourage additional searches for this frog in the lowland

Willamette Valley, especially where appropriate habitat pockets may exist that lack the

exotic warmwater macropredators probably detrimental to this taxon. Nevertheless,

substantial information on habitat change in the lowland Willamette River system (e.g.,

US Army Corps of Engineers 1982) indicating widespread alteration unfavorable to the

western spotted frog suggests that a low likelihood exists that this species will be

encountered in the lowland Willamette hydrographic basin in the future. All localities

where the western spotted frog is extant occur at elevations above 1219 m (4,000 ft),

where the combined impacts of exotic warmwater macropredators and hydrologic

modification are limited or absent. Since the known elevation range of the western spotted

frog in Oregon does not exceed 1524 m (5,000 ft; see Hayes 1994), a key pattern is that

Table 5 - Western Spotted Frog Detection and Search Time Statistics

Site Visit

1 2 3

Locality (County, State) dt tst dt tst dt tst

1) Trout Lake marsh (Klickitat, WA) 3 0.2* 7 2.3 5 2.5

2) Conboy NWR (Klickitat, WA) 9 0.7* - - -

3) Big Marsh (Klamath, OR) 0.5 2.0 0.8 2.3 0.6 2.0

4) Davis Lake, Odell Crk inflow (Klamath, OR) 17 2.5 14 3.3 - -

5) Odell Crk above NFD 4660 (Klamath, OR) 15 2.0 12 2.4 - -

6) Wood River Ranch (Klamath, OR) 10 0.5* 5 2.0 3 2.3

7) Gold Lake Bog (Lane, OR) 0.3 5.0 0.3 6.0 0.4 3.7

8) Penn Lake (Lane, OR) 16 4.6 - - - -

Under the site visits (1-3), "dt" is the detection time or the number of minutes after beginning the search ofthe site that western spotted frogs were detected and "tst" or total search time is the total time spentsearching for western spotted frogs during a site visit.

The asterisked (*) site visits do not follow the minimum two-hour search protocol because they wereconducted before the protocol was established or by other parties. Site visits to Conboy NWR and thefirst site visit to Trout Lake marsh were conducted in 1992, and the first site visit at the Wood RiverRanch is the original reconnaissance when Ron Hicks (BLM) rediscovered the western spotted frog inthe Wood River system.


this taxon is being crowded up against the upper limit of its elevation range, a pattern which

has been observed in other amphibians formerly common in lowland habitats in the western

United States (Schaffer and Stanley 1992: California tiger salamander (Ambystoma

californiense); Hayes, unpubl. data: California red-legged frog (Rana aurora draytonii)). If

spotted frogs still occur in as yet unidentified localities in western Oregon, they have the

highest probability of occurring in habitat at these higher elevations because such sites are

mostly likely to possess habitat matching their requirements. Nevertheless, spotted frogs

are now unquestionably rare in the region.

Demographic data suggest that of the populations of western spotted frog encountered

in western Oregon, only the Gold Lake Bog population appears to have both successful

recruitment and adequate adult survivorship (see Table 4). Data on the size structure of the

Penn Lake and Wood River Ranch populations suggest that both are threatened; the former

by a factor that may be curtailing recruitment, and the latter by a factor that may be reducing

adult survivorship (see Table 3). Penn Lake harbors an extremely dense population of

exotic brook trout (aggregations of 100-200 brook trout 20-40 cm long were observed at

several places, even in water less than 0.75 m deep, along some edges of Penn Lake), and

metamorphosis of the demonstrably unpalatable western toad in Penn Lake was extremely

successful (over 300,000 recently metamorphosed western toadlets were observed along

the northeast shoreline of Penn Lake during the survey for western spotted frogs). These

observations may indicate that brook trout are the selective agent suppressing recruitment of

the frogs in Penn Lake because the Pacific chorus frog (=treefrog), a species with highly

palatable larvae that was observed to be extremely abundant in nearby lakes in this basin

(Mink Lake basin), was also observed in very limited numbers in Penn Lake. The Wood

River Ranch locality displays significant habitat alteration characteristic of the vast shallow

marshes historically associated with the Wood River system. In particular, a large degree

of small- and large-scale channelization has greatly reduced the total area of water surface

that was present historically. This change is coupled with a measurably reduced water

quality (R. Hicks, pers. comm.) because of eutrophication from heavy livestock use.

These modifications probably reduce refuge habitat, especially for adult western spotted

frogs, which require an open, shallow water refuge, and likely results in their reduced

survivorship.

Survey of historical collections of the Cascades frog indicates that it was sympatric,

although perhaps not syntopic with the western spotted frog, at at least four Cascade slope

localities besides Gold Lake Bog. The Cascades frog is still extant at all four (see 3, 4, 14,


and 17 in Table 3). Similarly, historical collections of the northern red-legged frog suggest

that they were sympatric with the western spotted frogs at perhaps at all ten of the lowland

Willamette Valley localities (assuming the locality imprecision is allowed for). Northern

red-legged frogs may be extant at only two of these localities (see 1 and 15c in Table 3).

These patterns underscore the more significant habitat alteration that has occurred the

lowland Willamette hydrographic basin when compared to areas at higher elevations.

The data continue to be consistent with the previously formulated hypothesis that the

western spotted frog is associated with warmwater marshes (see Hayes 1994). Data on

water temperatures taken next to active spotted frogs from all sites (n = 8) where this taxon

was encountered averaged 28.60 C (n = 287 individual temperatures), and the <5% of these

temperatures that were below 20'C were taken during early morning intervals when the

surface water had not yet warmed. The warmwater temperature relationship may be linked

to the data on marsh size, which suggests that a minimum area size that may not be very

small may be needed to support a western spotted frog population. An aquatic site may

have to be a minimum size just to attain the warm water temperatures in which western

spotted frogs are observed, particularly if the inflow to the marsh consists of cool or cold

water, as is the general pattern in much of the Pacific Northwest. Moreover, the more

aquatic habit of western spotted frogs may restrict them to their preferred aquatic habitat

more than other frogs, a condition that may cause them to experience greater predation

pressure (the Cascades frog or the northern red-legged frog are both known to be capable

of using relatively small and cold aquatic sites in which to breed, and both are known to

engage in movements which consistently remove adults from the high predation conditions

present at the breeding site). Thus, western spotted frogs may require a larger aquatic site

simply to maintain a viable population; presumably, the larger site can maintain the larger

population that can sustain greater predation pressure.

Finally, the pattern of detection of western spotted frogs suggests that the protocol used

was adequate to identify even apparently low-density populations (e.g., Davis Lake; see

Table 3). The diurnal activity and close tie to shallow warm water sites of western spotted

frogs facilitated surveying for this species. It needs to be emphasized, however, that the

success rate in detection might not have been as high had not significant preparation

involving both examination of aerial photographs and topographic maps to determine the

potential most favorable areas been done beforehand. The latter is absolutely necessary to a

successful search that minimizes total search and the number of days searched.


Recommendations

Verification that the western spotted frog was extirpated or near-extirpated over much

of western Oregon provides ample justification to list this species, especially since only one

of the three western spotted frog populations (e.g., Gold Lake) found may not be in

decline. Moreover, even if a few additional undetected populations of the western spotted

frog exist in western Oregon, a regional-level catastrophe could easily eliminate the few

remaining populations in the region. In Oregon, the Deschutes system and selected closed

basins in south central Oregon have 20-odd historical localities that remain to be surveyed

for this species (see Hayes 1994), but scattered anecdotal evidence suggests that this taxon

is no better off there. No extant localities for the western spotted frog below 1219 m

(4,000 ft) are known from eastern Oregon, so this taxon may be equally restricted to the

headwaters of the systems in which it occurred historically in this region as well. Further,

the broad distribution of brook trout in lakes possessing habitat for the western spotted frog

in eastern Oregon may threaten those populations in a manner similar to that postulated for

some of the remaining western Oregon populations. Thus, few remaining populations and

potentially serious threats to remaining populations indicate that the western spotted frog

should be listed as Endangered in the state.

Verification of the habitat requirements suggested by the data in this study need to be

pursued. In particular, the potentially significant relationship between water temperature,

minimum size of the aquatic site, and predation levels needs to be much better understood.

This is crucial to the effective management of this taxon, especially to identify sites that can

maintain viable populations and to limit hydrological modifications that may be detrimental.

The relative long winter season in 1992-1993 and the correspondingly shorter summer

season that followed prevented surveying as many additional sites with the characteristics

suitable for western spotted frog as desired. Several sites not surveyed should be surveyed

as soon as the opportunity arises. These include (county indicated in parentheses): 1) the

Dwarf Lakes complex of the Sky Lakes Wilderness (Klamath), 2) Aspen Lake (Klamath),

3) Jackson-Irving Creeks Marsh along Willamson River (Klamath), 4) Sycan Marsh (Lake-

Klamath), 5) Sellers Marsh (Klamath), 6) Wildcat Swamp (Lane), and 7) Big Meadows

complex in Warm Springs Indian Reservation, but on the upper west slope drainage of the

Cascades (Clackamas).

In the course of fieldwork, it was recognized that the greater sandhill crane appears to

often overlap with the western spotted frog in its use of habitat, a pattern which should be


investigated for its historical congruence as well. The nature of this relationship is unclear,

but greater sandhill cranes are known to require a marshy prairie of minimum size in which

to forage or reproduce, probably to minimize predation on themselves (Stern et al. 1987).

Cranes eat frogs (M. Stern, pers. comm.), so the relationship might also be a trophic one.

Whatever the case, the relationship needs investigation as potential exists to manage both

species simultaneously. In a similar vein, a reasonable likelihood exists that selected marsh

plants, because of their shallow warmwater requirements, may serve as excellent indicators

of the presence of western spotted frogs. This possibility needs investigation.

The two remaining recommendations of Hayes (1994) also need to be pursued. These

are: 1) evaluation of the current status of the western spotted frog in eastern Oregon in a

manner parallel to that done for western Oregon is needed to determine the status of the

western spotted frog statewide; and 2) evaluation of the current and historical status of the

other spotted frog taxa eastern Oregon in a manner parallel to that done for the western

spotted frog as these may have an even more restricted range than the western spotted frog

within the state.

Finally, a minimum of two days of diurnal search with a minimum search time of two

hours per locality is the recommended search protocol for this species if data on specifics of

site suitability are gathered beforehand from aerial photographs and topographic maps. If

such data is not gathered beforehand, three days of diurnal search should be conducted.

Search days should be temporally separated by intervals of at least one month. Searches

should be conducted absolutely no earlier than 15 April nor later than 15 October of any

year, and it would be preferable to conduct them between 15 May and 15 September, with

roughly a month shift later the suggested initiation interval for surveys at elevations above

914 m (= 3,000 ft). Searches should be conducted between 10:00 and 16:00 of each

search day. Surveyors should use a water thermometer and should focus efforts in waters

exceeding 20'C during the above-noted hours and months. Following this protocol will

maximize the probability of detecting the western spotted frog throughout its geographic

range, if it is present.

Acknowledgments

Many individuals contributed to my ability to develop this report. Acknowledgments

for the many individuals who contributed to the original Phase I report is listed therein

(Hayes 1994). Charlotte Corkran, David Corkran, Tim Fox, Cheryl Friesen, Larry Gangle


III, Rebecca Goggans, Ron Hicks, Mark Jennings, David Jepsen, John-Paul Leeming,

Lisa Lyon, Jill Mellen, Steve Mamoyac, David Marshall, Maura Naughton, Ralph Opp,

Dennis Pate, Margie Pate, Robert Penson, David Shepherdson, Mark Stern, Eric Terdahl,

Laurie Turner (formerly Church), Beth Waterbury, and the biologists and biological crews

at the Zigzag Ranger District of the Mt Hood National Forest and the Klamath Ranger

District of the Klamath National Forest either assisted in the field, contributed data or

insights, or assisted in finding bits of crucial information. Ron Hicks (Bureau of Land

Management, Klamath Falls) deserves special mention for his rediscovery of the western

spotted frog in the Wood River system. Steve Mamoyac and Paul Scheerer (Oregon

Department of Fish and Wildlife, Corvallis) and Mark Buchner (Bureau of Reclamation,

Klamath Falls) assisted in verifying some of the fish identifications. Jill Mellen provided

emotional support and assisted in editing and proofing the manuscript.

Literature Cited

Hayes, Marc P. 1994. The spotted frog (Rana pretiosa) in western Oregon. Final Reportto the Oregon Department of Fish and Wildlife. 30 pp.+ appendices





St. John, Alan D. 1984a. The herpetology of Jackson and Josephine Counties, Oregon.Oregon Department of Fish and Wildlife, Nongame Wildlife Program, TechnicalReport (84-2-05): 1-78.


Hayes: Spotted Frog Status in Western Oregon I11



Shaffer, H. B., and S. Stanley. 1992. Final report to California Department of Fish andGame; California tiger salamander surveys, 1991--Contract (FAG 9422). CaliforniaDepartment of Fish and Game, Inland Fisheries Division, Rancho Cordova,California.

Stern, Mark, Carroll D. Littlefield, and Geoff Pampush. 1987. The status and distributionof greater sandhill cranes in Oregon, 1986. A co-operative project conducted by theNature Conservancy in co-operation with the Nongame Division of the OregonDepartment of Fish and Wildlife, the Bureau of Land Management, and the US Fishand Wildlife Service.


U.S. Army Corps of Engineers. 1982. Willamette River projects, hydrologic andtemperature effects. Portland District, Portland, OR. 111 pp.

Appendix I - Other Western Oregon Localities Visited

Site Visits


1) Trapper Creek Meadows (Clackamas) 0 0 0 N,P 0 0

2) Head of Willarnson River (Klamath) 0 0 - P 0 ?

3) Klamath Forest NWR (Klamath) 0 0 - P 0 0

4) Lake of the Woods marsh (Klamath) 0 0 - PW + L,Usn*

5) Penn Lake (Lane) + - - PW 0 Br

6) Mink Lake (Lane) 0 - - CP 0 Br*

7) Latourelle Prairie (Multnomah) 0 - - N,P 0 Cu*


a OF = Other native frogs; codes are: C = Cascades frog (Rana cascadae), N = Northern red-legged frog (Ranaaurora aurora), and P = Pacific (treefrog) chorusfrog (Pseudacris (= Hyla) regilla).

b Bf = Bullfrog (Rana catesbeiana).c Fishes codes: Br = Brook trout (Salvelinus confluentus), Cu = Cutthroat trout (Oncorhynchus clarki), L =

Largemouth bass (Micropterus salmoides), and Usn = Unidentified sunfish.

Appendix II - Other Western Spotted Frog Localities Visited

Site Visits

Locality (County, State) 1 2 3 OFa Bfb Fishesc

1) Trout Lake marsh (Klickitat, WA) + + + P 0 ?

2) Conboy NWR (Klickitat, WA) + - - P H ?

3) Big Marsh (Klamath, OR) + + + PW 0 Cu*

4) Davis Lake, Odell Crk inflow (Klamath, OR) + + - PW 0 ?

5) Odell Crk above NFD 4660 (Klamath, OR) + + - PW 0 ?


a OF = Other native frogs and toads; codes are: P = Pacific (treefrog) chorusfrog (Pseudacris (= Hyla) regilla)and W = Western toad (Bufo boreas).

b Bf = Bullfrog (Rana catesbeiana). An (H) indicates that bullfrogs were historically present in this system,but have not been observed recently at the sites where western spotted frogs have been seen.

c Fishes codes: Cu = Cutthroat trout (Oncorhynchus clarki).

SOUTHERN GREGO'N S-, iI lL'it LASHLAND, OPEGON 97520

DO*SOMETHI NC-W!LD

SUPPORTOREGON'SNONGAMEWILDLIFEOregon Departmient of Fish & Wildlife

OREGON Oregon Department of Fish and WildlifeL~~~ 2501 SW First Ave., P.O. Box 59~~73 Portland, Oregon 97207

0*.haw Uw.4.

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THE SPOTTED FROG IN WESTERN OREGON - …soda.sou.edu/awdata/030728a1.pdf · a In the context of this report, western Oregon refers to Oregon west of the crest of the Cascade Mountains