Late Quaternary mammalian zoogeography of eastern Washington

QUATERNARY

RESEASRCH

Late Quaternary

20, 360-373 (1983)

Mammalian

Zoogeography

of Eastern Washington

R. LEE LYMAN AND STEPHANIE D. LIVINGSTON* Department

of Anthropology, Anthropology

Oregon State DH-05, University

University. Corvallis, Oregon 97331 and *Department of Washington, Seattle. Washington 98195

of

Received October 11, 1982 The late Quatemary mammalian zoogeographic history of eastern Washington as revealed by archaeological and paleontological research conforms to a set of past environmental conditions inferred from botanical data. During the relatively cool and moist late Pleistocene andearly Holocene, Cervus cf. elaphus, Ovis canadensis. Vulpes vulpes, Martes americana, Alopex lagopus, and perhaps Rangifer sp., taxa with ecological preferences for mesic steppe habitats, were present in the now xeric Columbia Basin. As the climate became progressively warmer and drier during the late Pleistocene and early Holocene, Antilocapra americana, Onychomys leucogaster, Spermophilus townsendii, and Neotoma cinerea, taxa with ecological preferences for xeric steppe habitats, appear in the Columbia Basin. Bison sp. and Taxidea taxus may have been presentin eastern Washington for the last 20,000 yr. Middle and late Holocene records for Oreamnos americanus. Spermophilus columbianus, S. townsendii, Lagurus curtatus, and Urocyon cinereoargenteus in central eastern Washington suggest fluctuations in the ranges of these taxa that conform to a middle Holocene period of less effective precipitation and a ca. 3500-yr-old period of more effective precipitation before essentially modem environmental conditions prevailed.

INTRODUCTION

Unlike many other parts of North America, little attempt has been made to provide analytic studies of the late Pleistocene and Holocene mammalian history of eastern Washington and adjacent Oregon. During the past 40 years archaeological investigations and sporadic paleontological studies have generated sufficient information to make such a study possible. In this paper we compare the available mammalian zoogeographic information to the environmental record derived from botanical data to shed light on the historical development of the modern distributions of mammals in this region and to further illuminate the paleoenvironmental record. STUDY AREA

We shall be concerned with those parts of Washington State and adjacent Oregon that lie east of the crest of the Cascade Range. This area is quite diverse physiographically (Franklin and Dyrness, 1973; Hunt, 1967; McKee, 1972). It includes the 0033-5894183 $3.00 Copyright @ 1983 by the University of Washmgton. Au rights of reproduction in any form reserved.

360

northern three fourths of the Columbia Basin as defined by Freeman et al. (1943, the eastern flanks of the Cascade Range, the Okanogan Highlands, the western edge of the Selkirk Mountains, and the northern portion of the Blue Mountains (Fig. 1). Mean annual precipitation ranges from less than 20 cm in the central Columbia Basin to more than 60 cm in northeast Washington and 250 cm near the crest of the Cascades. January mean temperature minima range from - 13” to -5”C, changing with elevation and latitude. July temperature maxima range from 22” to 34”C, also changing with elevation and latitude. Vegetation varies along temperature, precipitation, and altitudinal gradients from relatively hot, dry, shrub-steppe communities to subalpine coniferous forests. Local edaphic conditions result in a wide variety of habitat types occurring in extremely small areas (Daubenmire and Daubenmire, 1968; Daubenmire, 1970). The diversity of habitats provides for both a complex fauna1 history and a rich, extant mammalian fauna. Hall (1981) re-

MAMMALIAN 121*

ZOOGEOGRAPHY 120°

361

OF WASHINGTON

1190

ll?O

118O

rk-

48’

I

Yw,-COLUMBIA

PLATEAU

FR 22

7 : + 47O

46

1210

1200

ll9O

ll8O

1170

FIG. 1. Eastern Washington fauna1 regions (after Dalquest, 1948) and geographic place names and sites mentioned in the text. (1) 35WS4 (Road Cut); (2) 35UM7 (Cold Springs); (3) 45BN3 (Berrian Island) and 45BN254 (Umatilla Mammoth); (4) Prescott; (5) Delight; (6) 45FR50 (Marmes Rockshelter); (7) 45WT2; (8) 45COl (Iltcannon); (9) 45WT7; (10) 45GA17; (11) 45WT39 (Wawawai) and 45WT41 (Granite Point); (12) 45AS78 (Pig Farm), 45AS80, and 45AS82 (Timothy’s Village); (13) Jeppson Locality; (14) 45KT241; (15) Cox’s Pond (45DO172); (16) Indian Dan (450K58); (17) 450K145 (Coulee Creek Rockshelter); (18) 45FE24; (19) Ksunku (45FE45); (20) 45GR97 (Lind Coulee): (21) 45AD2; (22) 45LIl and 45LI3 (Meyer Cave).

cords 93 mammalian species representing 58 genera in the study area today (not counting introduced domesticates), with 6 or 7 additional species marginally present. Of these, 40 species representing 35 genera have been identified in the Holocene fauna1 record.

PREHISTORIC

FAUNAL RECORD

Paleontological research in the area of concern has been minimal and has focused largely on Pliocene and early Pleistocene faunas (Gustafson, 1978; Fry and Gustafson, 1974). Most late Pleistocene faunas

362

LYMAN

AND

LIVINGSTON

Because the entire taphonomic history of a prehistoric fauna is difficult to ascertain, the extent to which a sample of bones resembles the original fauna cannot be determined (Grayson, 1981). It is reasonable, however, to assume that taxa represented in a single sample lived in the area of the site. Samples from multiple sites are required to determine whether the absence of a taxon in an area reflects its absence in the original fauna or sampling error (Grayson, 1981, 1982; Guilday et al., 1978; King and Graham, 1981). In the analysis to follow, we have necessarily assumed that the present ecology of specific mammals is the same as the ecology of those mammals in the past. In this analysis we circumvent the problem that the assumption may be false (Grayson (1981) and references therein) by using suites of taxa the members of which have essentially the same ecological tolerances and preferences. This approach allows us to infer past environmental conditions more confidently on the bases of the presence and absence of mammalian taxa. In 1948 Walter W. Dalquest presented an hypothetical zoogeographic history for the state of Washington. In this history, Dalquest (1948, pp. 55-109) speculated on Holocene dispersal chronologies and routes for many of Washington’s mammalian species (Table 1). In brief, Dalquest hypotheMATERIALS AND METHODS sized that with the recession of late WisData for this study were derived by a consin glaciers, early Holocene warming, search of published and unpublished ar- and the adjustment of habitat distributions, chaeological and paleontological reports shifts in the ranges of taxa occurred until pertaining to the study area and sur- modern conditions were attained. The zoorounding regions. Critical analyses of the geographic history reflects Dalquest’s relationship between relative taxonomic (1948) understanding of modern habitat and abundances and other paleoenvironmental fauna1 distributions and the potentially sigindicators like palynological and alluvial nificant variables affecting the dispersal of chronological data indicate serious defimammalian taxa. He definedfuunal regions ciencies in many fauna1 samples (R. L. (hereafter FR) on the basis of the distribuLyman, unpublished data). Only those tion of life zones in conjunction with the faunas of special zoogeographic and pa- courses of the Columbia, Snake, Okaleoenvironmental significance will be con- nogan, and Yakima rivers in eastern Washsidered in detail here. ington (Fig. 1). Dalquest speculated that

have not been dated by radiometric techniques. However, two recently reported late Pleistocene faunas have been tentatively dated on stratigraphic bases (Gilbow, 1981; Martin et al., 1982). Archaeological research in this area has been largely concerned with establishing cultural chronologies. Preoccupation with the temporal and spatial distributions of prehistoric peoples has seriously influenced the recovery and analysis of fauna1 remains collected from excavated sites. There are few 14C dates stratigraphically associated with known faunas. Most dating is based on artifact typologies, and because of differences between local cultural chronologies, significant zoogeographic events may be obscured. Spatial patterns are also difficult to ascertain because most archaeological work has been done under the auspices of cultural resource management programs and most data dating to the Holocene are restricted to major river canyons where hydroelectric and other water management projects have been located. Early archaeological site reports usually include a list of represented taxa, but detailed data concerning the spatial-temporal distributions of these taxa generally are not reported. Later reports are, however, characterized by greater emphasis on the cultural and stratigraphic associations of taxa.

MAMMALIAN TABLE

1. SUMMARY

OF DALQUEST’S

ZOOGEOGRAPHY (1948)

OF WASHINGTON

ZO~GE~GRAPHIC

MODEL

363

FOR EASTERN WASHINGTON”

1. Present in Columbia Plateau FR and/or SE Washington FR before and/or during the last glacial age Taxidea taxus Odocoileus hemionus*a Ovis canadensis (?)* Ondatra zibethicus (?)* Sylvilagus nuttallii (?)*

Lepus townsendii Spermophilus washingtoni Perognathus parvus Thomomys talpoides Microtus pennsylvanicus* Procoyon lotor

2. Forced south into eastern Washington by glacier advance Castor canadensis Gulo luscus Phenacomyr intermedius Martes americana

Lynx canadensis* Synaptomys borealis Martes pennanti

3. Late Pleistocene refugium in Blue Mountains FR Lepus americana* Canis lupus* Felis concolor* Vulpes vulpes* Mustela erminea* Tamiasciurus hudsonicus* Clethrionomys gapper? Ovis canadensis*

Spermophilus columbianus Spermophilus lateralis Microtus montanus* Microtus richardsoni* Mus tela frena ta* Eutamias amoenus* Glaucomys ,sabrinus* Zapus princeps*

4. Late Pleistocene refugium in southern Cascade Range FR Ochotona p,rinceps Microtus montanus* Oreamnos americanus Odocoileus hemionus* Mustela erminea* Eutamias amoenus* Tamiasciurus hudsonicus* Glaucomys sabrinu? Mustela frenata*

Lepus americana* Lynx canadensi? Ovis canadensis* Mephitis mephitis* Vulpes vulpes* Clethrionomys gapper? Microtus richardson? Zapus princeps*

5. Late Pleistocene refugium in Rocky Mountains Marmota monax Marmota caligata Microtus pennsy/vanicus* Lepus americana* Canis lupus* Felis concolor* Martes americana Mu&la erminea* Eutamias amoenus* Tamiasciurus hudsonicus*

Ursus americanus Vulpes vulpes* Lynx canadensis* Ovis canadensis* Cervus elaphus Odocoileus virginianus Mephitis mephitis* Clethrionomys gapperi* Zapus princeps* Glaucomys sabrinus*

6. Late Pleistocene refugium in Great Basin of eastern Oregon, post Wisconsin entry to eastern Washington Lagurus curtatus Neotoma cinerea Spermophilus townsendii Mustela frenata Mephitis mephitis* Bison bison Spilogale putorius Dipodomys ordii

Marmota flaviventris Onychomys leucogaster Sylvilagus nuttallii* Lynx rufus Antilocapra americana Lepus californicus Eutamias minimus Reithrodontomys megalotis

n Asterisks indicate that species occurs in more than one zoogeographic

certain FRs had served as Pleistocene refugia for particular taxa (Table 1). Dalquest (1948, pp. 52-53) also defined three modern “faunas” in Washington, two

group.

of which are relevant here. Each fauna occurs more or less distinctly in a particular and limited habitat and range (Table 2). Dalquest believed that the modern taxo-

364

LYMAN

TABLE

2. SUMMARY

OF DALQUEST’Y

AND

LIVINGSTON

MAMMALIAN

Great Basin faunab Taxidea taxus (11’ Marmotaflaviventris (6) Spermophilus townsendii (6) Spermophilus washingtoni (I) Eutamias minimus (6) Sylvilagus nuttallii (1, 6) Perognathus parvus (1)

FAUNAS

OF EASTERN

WASHINGIUN

Dipodomys ordii (6) Onychomys leucogaster (6) Reithrodontomys megalotis (6) Lagurus cm-tutus (6) Lepus californicus (6) Sy1lGlagu.s idahoensis

Rocky Mountain fatmad Lynx canadensis (2, 4, 5) Marmota caligata (5) Spermophilus lateralis (3) Spermophilus columbianus (3) Eutamias amoenus (3, 4, 5) Tamiasciurrcs hudsonicus (3, 4, 5) Glaucomys sabrinus (3, 4, 5)

Synaptomys borealis (2) Phenacomys intermedius (2) Microtus richardsoni (3. 4) Ochotona princeps (4) Lepus americana (3, 4, 5) Rangifer tarandus Oreamnos americanus (4)

0 Dalquest, 1948 (pp. 52-53). b In general, members of this group occupy the Columbia Plateau, SE Washington. and Yakima Valley FRs and NE Oregon at the present time. c Number(s) in parentheses indicate the zoogeographic group to which the particular taxa belong (Table I). d In general, members of this group occupy the Cascade, NE Washington, and Blue Mountain FRs at the present time.

nomic composition of each of these faunas was attained as a result of environmental fluctuations during the last 20,000 years affecting zoogeographic shifts in the ranges of particular taxa (compare Tables 1 and 2). By tracing the zoogeographic history of the taxa in Tables 1 and 2, paleoenvironmental conditions might be inferred. There are, however, problems with such an approach (Grayson (1981) and references therein). In order to detect problems like stratigraphic mixing and long distance transport of skeletal remains, the paleoenvironmental history of eastern Washington as inferred from botanical remains will be employed as a control for environmental implications derived from the fauna1 record. Phytogeographic data suggest that an altitudinal lowering of habitats occurred (Daubenmire, 1975) and palynological data indicate that a cool, mesic steppe habitat existed in the Columbia Plateau FR during the late Pleistocene (Mack et al., 1976; Martin et al., 1982). The available geological evidence indicates the last Wisconsin advance began ca. 17,500 yr B.P. and that the recession was taking place ca. 13,000 yr

B.P. (Clague et al., 1980; Mullineaux et al., 1978). These dates may reflect the timing of fluctuations in habitat ranges as well. Palynological data further indicate a period of less-effective precipitation between ca. 8000 and 4000 yr B.P. (Mack et al., 1976, 1979; Nickman, 1979), and, in general, a warm, dry steppe habitat in the Columbia Plateau FR is indicated. Finally, there is limited evidence for a brief period of cooler, moister environments, centering around ca. 3500 yr B.P. (Mack et al., 1978), before essentially modern environmental conditions prevailed. RESULTS AND DISCUSSION

Dalquest’s (1948) study, as summarized in Tables 1 and 2, holds specific implications for late Quaternary environmental conditions in eastern Washington as reflected by mammalian zoogeography. The implications concern the zoogeographic histories of what we here term “Borealmesic steppe taxa” and “xeric steppe taxa.” The environmental significance of the members of each group of taxa will be summarized, the relevant zoogeographic data will be reviewed, and the similarities

MAMMALIAN

ZOOGEOGRAPHY

and differences between the environmental implications of the botanical and mammalian records will be described. Boreal-Mesic

Steppe Taxa

If the late Quaternary variation in habitats indicated by the botanical data is correct, taxa with preferences for cooler and moister habitats should be found in late Pleistocene sediments (ca. 18,000 to 10,000 yr B.P.) and in sediments dating to ca. 3500 yr B.P. in areas where they are not found today and which today are warm and dry. The mammalian taxa involved minimally include those listed in Zoogeographic Groups 2, 3, 4, and 5 (Table 1) and in the Rocky Mountain fauna (Table 2). Bones of Kangifer sp. of unclear but probably Blancan age (Strand and Hough, 1952; Gustafson, 1978) have been found near Wallula Gap (Fig. 1). This taxon may have been present here during the Pleistocene, as well. The nearest extant population of Rangifer is in the Selkirk Mountains, some 250 km northeast of Wallula Gap (Hall, 1981, p. 1105). Bones of Alopex lagopus and Martes americana dating to ca. 10,000 yr B.P. have been found along the Snake River at Marmes Rockshelter (Fig. 1) (Gustafson, 1972). The nearest extant population of A. lagopus is over 1000 km north of the Snake River (Hall, 1981, p. 936) and the nearest extant population of M. americana is found in the Blue Mountains FR, 50 km south of the Snake River (Hall, 1981, p. 983). Bones of Bootherium sp., a possible woodland form (Martin and Guilday, 1967), have been found at the Umatilla Mammoth Site west of Wallula Gap (Gilbow, 1981), however, the status of this genus is currently in doubt (Kurten and Anderson, 1980). Stratigraphic data suggest that the Umatilla Mammoth Bootherium specimens of this now extinct musk-ox date to shortly after 13,000 yr B.P. Thus far, all Lynx sp. bones have been identified with confidence only to the genus level. While it is unknown when Castor

OF

WASHINGTON

canadensis first

365

entered eastern Washington, it apparently occupied a wide range during the early Holocene. Records for this taxon are reported at the Lind Coulee Site in the center of the Columbia Plateau FR, dating to ca. 8700 yr B.P. (Irwin and Moody, 1978), and at the Road Cut Site along the farthest downstream reaches of the Columbia River within the study area, dating to 8000 to 10,000 yr B.P. (Cressman et al., 1960). The earliest records for the taxa in Zoogeographic Groups 3, 4, and 5 (Table 1) are all younger than 11,000 yr BP. However, the available data indicate that some of these taxa were in areas other than their postulated Pleistocene refugia in early Holocene times (Tables 3 and 4). Aubrey (1983) suggests that the eastern slopes of the Cascade Range are now occupied by an indigenous population of Vulpes vulpes. Individuals of this taxon found in the northernmost Yakima Valley, in extreme northeastern Washington, and in the Columbia Plateau, Southeast Washington, and Blue Mountains, FRs are believed by Aubrey to represent descendants of populations introduced to eastern Washington during the historic period. The prehistoric data (Table 4) suggest indigenous groups of V. vulpes may have been present in the Columbia Plateau and Southwest Washington FRs during the middle and late Holocene, respectively, areas that are regularly believed to be too arid for this taxon (Hall, 1981, p. 938). The indigenous form tends to be adapted to more mesic and forested conditions, while the introduced form tends to be adapted to the drier conditions of lower elevations in eastern Washington (Aubrey, 1983). Specimens of Oreamnos americanus are reported from two sites, both in late Holocene (~3000 yr B.P.) contexts. One is from 45FE24 (Collier et al., 1942) in the northeast (NE) Washington FR, and the other is from Coulee Creek Rockshelter (Grabert, 1974), in the northern half of the Cascade FR (Fig. 1). Today, 0. americanus

LYMAN

366 TABLE

3. EARLIEST HOLOCENE

RECORDS

AND LIVINGSTON FOR TAXA

IN ZOOGEOGRAPHIC

GROUPS

3.4, AND Y

Age

Taxon Felis concolor Mephitis mephitis

Cervus cf. efuphus

Ovis canadensis

Spermophitus

Ursus

columbianus

americanus

Reference

Site

(yr B.P.)

Road Cut Road Cut Lind Coulee Road Cut Marmes Rockshelter Lind Coulee Ksunku 45WT2 Cold Springs Cox’s Pond Granite Point Tucannon Cold Springs Granite Point TUcannon 45WT7 Road Cut Ksunku Ksunku 45GA17

lO,OOO-8000 lO,OOO-8000 8700 lO,OOO-8000 lO,OOO-7500 8700 8000-6000 >7300 >6700 6700-3000 8000-6700 8000-5000 >6700 8000-6700 8000-5000 8000-5000 8000-4000 6000-4400 6000-4400 2330

Cressman ef al., 1960 Cressman et al., 1960 Irwin and Moody, 1978 Cressman er al., 1960 Gustafson, 1972 Irwin and Moody, 1978 Grayson, 1977 Gustafson, 1972 Shiner, 1961 Hartmann, 1975 Gustafson. 1912 Bense. 1972 Shiner, 1961 Gustafson, 1972 Bense, 1972 Bense, 1972 Cressman et al., 1960 Grayson, 1977 Grayson, 1977 Sprague ef al., 1968

a Table 1: see also Table 4.

occupies much of the higher Cascade Range and the Blue Mountains of northeast Oregon (Hall, 1981, p. 1112). Dalquest (1948, p. 409) reported that a single individual from northeast Washington probably represented a “wanderer” from northern Idaho. The archaeological specimens were recovered from sites outside of the present range of 0. americanus, and could be representative of extralimital populations. The specimens may, however, have been transported to these lowland riverine sites by people. Eight fragments of mountain goat horn were recovered from 45FE24; horn is TABLE 4. HOLOCENE

a commonly used tool material ethnographically and prehistorically in eastern Washington. Several tooth fragments identified as 0. americanus were recovered from Coulee Creek Rockshelter; there is no good evidence suggesting that teeth were transported long distances by people. While Cervus elaphus is reported at the Lind Coulee Site in the central Columbia Plateau FR at 8700 yr B.P. (Irwin and Moody, 1978), this taxon is not present in faunas in the central Columbia Plateau FR that are less than about 7000 yr old (Meyer Cave (Bryan, 1955); 45AD2 (Deaver and

RECORDS

FOR Vulpes

vulpes

As

Site

(yr B.P.)

Reference

Road Cut Lind Coulee Granite Point 45AD2 Timothy’s Village Pig Farm Benian Island

lO,oOO-4000 8700 8000-6700 3500-1000 5000-2500 3000-100 200

Cressman et al., 1960 Daugherty, 1956; Irwin and Moody, 1978 Gustafson , 1972 Deaver and Greene, 1978 Lyman, 1976 Lyman, 1976 Osborne, 1953, 1957

MAMMALIAN

ZOOGEOGRAPHY

Greene, 1978)). Ovis canadensis has not been reported in late Pleistocene or Holocene contexts in the central Columbia Plateau FR. The available data concerning Borealmesic steppe taxa indicate that two taxa, Rangifer and ,4. lagopus, should be added to Zoogeographic Group 2 (Table 1). The presence of Alopex, M. americana, C. canadensis, V. vulpes, and C. cf. elaphus in the central Columbia Plateau FR, Bootherium in the Yakima Valley FR, and C. cf. elaphus and 0. canadensis in NE Oregon between ca. 13,000 and 8000 yr B.P. and the subsequent general disappearance of many of these taxa from those FRs is to be expected given the trend toward warmer and drier environmental conditions suggested by the palynological record at this time. Possible responses of Boreal-mesic steppe taxa to the suggested cooler and wetter environment ca. 3500 yr B .P. are not readily apparent. Perhaps the records for V. vulpes in the Columbia Plateau FR and the extralimital records for 0. americanus dating to ca. 3000 yr B.P. are reflecting this fluctuation. The apparent late Holocene disappearance of indigenous populations of V. vulpes from these arid lowlands of the Columbia Plateau (Aubrey, 1983) may reflect a shift in environmental conditions from cooler and moister than present toward those of the present, a suggestion corroborated by some palynological data. Not all of the mammalian zoogeographic data readily fit the environmental history suggested by the palynological record. In particular, the presence of Spermophilus columbianus in the center of the Snake River Canyon (Table 3), an edaphically arid environment, during a period indicated by the pollen record to have been a time of less effective precipitation, seems curious. While S. columbianus can occupy a varied habitat, it tends to occupy “more humid habitats than s’. townsendii and S. washingtoni” (Dalquest, 1948, p. 272). When S. columbianus is sympatric with the more xericadapted S. beldingi in northeastern Oregon,

OF

WASHINGTON

367

the former taxon tends to occupy more mesic microhabitats. When xeric-adapted competitors are absent and streams are nearby, however, S. columbianus occupies more xeric environments (Turner, 1972). What we may be seeing, then, is an instance of competitive release: perhaps only S. columbianus is reported in the archaeological sites listed in Table 3 and in sites to the east because S. washingtoni was not present. Gustafson (1972, pp. 60-61) suggested that “there appears to be no overlap of the ranges of S. washingtoni and S. columbianus” and that “the range boundary between these two species must lie between 45GA17 and Granite Point” (Fig. 1). Dalquest (1948, pp. 268, 274) and Hall (1981, pp. 384, 389) indicate, however, that the modern ranges of these two species do overlap. The possibility of prehistoric sympatry is suggested by two sites on opposite sides of the Snake River and a 2-km distance from one another. Because these two occurrences are separated by 3000 yr in time, the evidence for sympatry is weak; possibly we may be dealing with an instance of shifting ranges. S. columbianus has been found at 45WT7 (8000-5000 yr B.P.; Bense, 1972) and S. washingtoni at 45GA17 (2330 yr B.P.; Gustafson, 1972). Further research may eventually suggest whether shifting ranges and competitive release between S. washingtoni and S. columbianus are responsible for the presence of the latter taxon in the prehistoric record of the Snake River Canyon. Documentation of sympatry (bones of both species in association in a site) or competitive release would have implications for the environmental history of eastern Washington. Xeric Steppe Taxa

Dalquest’s (1948) zoogeographic history was developed mainly to account for those taxa he considered indigenous to Washington prior to the last glacial age. He believed many of these taxa may have occupied different ranges than at present during

368

LYMAN

AND

the Wisconsin glacial maxima (Table 1). The modern array of taxa, however, also contained species whose xeric adaptations and taxonomic affinities led him to suggest these taxa were Holocene immigrants from the Great Basin (Table 1, Zoogeographic Group 6; Table 2, Great Basin fauna). He proposed a four-stage sequence for the dispersal of these “Great Basin” mammals into eastern Washington. This sequence involved (1) the dispersal of Great Basin mammals into the southeast (SE) Washington FR from the north-central (NC) Oregon FR, (2) crossing the Snake and/or Columbia rivers into the Yakima Valley FR from the SE Washington FR, (3) crossing the Snake River and dispersing throughout the Columbia Plateau FR from the SE Washington FR, and (4) crossing the Columbia and/or Spokane rivers and entering the NE Washington FR from the Columbia Plateau FR. This dispersal sequence was postulated on the bases of the modern distributions of taxa, the taxonomic affinities of Thomomys populations in the southern Cascade, Yakima Valley, SE Washington, and Blue Mountain FRs (Dalquest and Scheffer, 1944), the potential of major rivers like the Snake and Columbia to act as barriers to dispersal (Davis, 1936), and the 20th-century dispersal of members of the Great Basin fauna into eastern Washington (Broadbooks, 1958, 1969; Couch, 1927). In addition to the post Wisconsin immigrants from the Great Basin, Dalquest (1948) considered that some taxa which are adapted to relatively xeric conditions may have been in eastern Washington during the last glacial age (Table 1, Zoogeographic Group 1). As climate tended to become warmer and drier at the end of the Pleistocene, these relatively indigenous members of the Great Basin fauna were joined by more recent arrivals from eastern Oregon. The botanical record indicates a general warming and drying trend since the last glacial age. Mammalian taxa listed in Zoogeo-

LIVINGSTON

graphic Groups 1 and 6 (Table 1) and as members of the Great Basin fauna (Table 2) should, therefore, appear in the prehistoric record throughout the late Quaternary of eastern Washington. The middle Holocene warming and drying phase and the ca. 3500 yr B.P. cooler and moister climatic phase may also have affected the ranges occupied by these taxa. Concerning the taxa in Zoogeographic Group 1 (Table l), Procyon and Taxidea are reported in “Irvingtonian” or later contexts at the Delight Site (Fry and Gustafson, 1974), and Taxidea is recorded in “late Pleistocene” sediments at the Willow Creek Site (Tipper ef al., 1951) (Fig. 1). Two genera, Perognathus and Thomomys. have been recovered from the ca. 13,000 yr B.P. Jeppson Locality (Martin et al., 1982). Early Holocene records for these taxa are summarized in Table 5. Concerning members of Zoogeographic Group 6 (Table I), two reports for late Pleistocene Bison sp. are available. One is from the Willow Creek Site (Tipper et al., 1951) and possibly dates to the “late Pleistocene. ” The other, from the Prescott Site (Pope, 1952), may date between 20,000 and 10,000 yr B.P. Onychomys leucogaster and S. townsendii are reported at the ca. 13,000 yr B.P. Jeppson Locality (Martin et al., 1982), and Antilocapra americana is reported at the ca. 13,000 yr B.P. Umatilla Mammoth Site (Gilbow, 1981). These records are located in the NC Oregon, SE Washington, and Yakima Valley FRs, areas involved in the first stages of Dalquest’s hypothesized four-stage dispersal scheme for these taxa. Records for taxa in Zoogeographic Group 6 (Table 5) indicate many members of this group were fairly widespread through the southern half of the study area by the early Holocene. We suggest adding two new members to the taxa listed in Zoogeographic Group 6 (Table 1). Urocyon cinereoargenteus was not considered by Dalquest (1948), but a single record for this taxon has been reported from sediments dating to ca. 2000 yr

MAMMALIAN TABLE

5.

EARLIEST

HOLOCENE

ZOOGEOGRAPHY RECORDS

FOR TAXA

OF WASHINGTON

IN ZOOGEM;RAPHIC

369

GROUPS

1 AND

6 (TABLE

1)

Age

Taxon Taxidea

taxus

cf. cin,erea

Neotoma

Sylvilagus

Antilocapra

Bison

cf. nuttallii

americana

sp.

Marmota

Perognathus

Onychomys

cf. j7aviventris

parvus

leucogaster

Site

(yr BP.)

Reference

Lind Coulee Granite Point Meyer Cave Road Cut Cox’s Pond Timothy’s Village Marmes Rockshelter Lind Coulee 45KT241 Granite Point Meyer Cave Marmes Rockshelter Lind Coulee Granite Point Meyer Cave Cold Springs Marmes Rockshelter Granite Point Meyer Cave lbcannon Lind Coulee Meyer Cave 45WT7 45WT2 Marmes Rockshelter

8700 8000-6700 8000-6000 80004000 6700-3000 SOCO-2500

Daugherty, 1956; Irwin and Moody, 1978 Gustafson, 1972 Bryan, 1955 Cressman et al., 1960 Hartmann, 1975 Lyman, 1976

10,000-7500 8700 8000-7000 8000-6700 8000-6000 lO,OOO-7500 8700 8000-6700 8000-6000 >6700 lO,OOO-7500 8000-6700 8000-6000 8000-5000 8700 8000-6000 8000-5000 6700-1300 lO,OOO7500 8700 8000-7000 8000-6700 8000-6000 8700 8000-7000 7000-5000 6000-4000 5000-2500

Gustafson, 1972 Irwin and Moody, 1978 Chatters, 1979 Gustafson, 1972 Bryan, 1955 Gustafson, 1972 Irwin and Moody, 1978 Gustafson, 1972 Bryan, 1955 Shiner, 1961 Gustafson, 1972 Gustafson, 1972 Bryan, 1955 Bense, 1972 Daugherty, 1956; Irwin and Moody, 1978 Bryan, 1955 Sprague e? af., 1968 Gustafson, 1972 Gustafson. 1972

Lind Coulee 45KT241 Granite Point Meyer Cave Lind Coulee 45KT241 Timothy’s Village Indian Dan Timothy’s Village

B.P. at 45AD2 in the center of the Columbia Plateau FR (Deaver and Greene, 1978). This taxon may have entered eastern Washington along a route similar to that proposed for other members of Zoogeographic Group 6. The nearest extant population of U. cinereoargenteus is 200 km to the southwest of 45AD2 (Hall, 1981, p. 944). The second additional taxon is Sylvilagus idahoensis. It has been suggested elsewhere that the apparently disjunct extant population of S. idahoensis (Dalquest, 1941; Hall, 1981, p. 295) in eastern Wash-

Irwin and Moody, 1978 Chatters, 1979 Gustafson, 1972 Bryan, 1955 Irwin and Moody, 1978 Chatters, 1979 Lyman, 1976 Browman, 1966 Lyman, 1976

ington became isolated from populations to the south at the onset of the middle Holocene as a result of decreased effective precipitation (Grayson, 1982). Two records for S. idahoensis are available, both in the central Columbia Plateau FR. One is from 45AD2 and dates to ca. 1500 yr B.P. (Deaver and Greene, 1978). The other record is from Lind Coulee and dates to ca. 8700 yr B.P. (Irwin and Moody, 1978). Apparent Holocene extralimital records also exist for two taxa in Zoogeographic Group 6. Bones of Spermophilus town-

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sendii have been reported from 45AD2 in contexts dating from 3000 to 1000 yr B.P. (Deaver and Greene, 1978). The nearest extant population of this species is 100 km to the southwest of this site (Hall, 1981, p. 383). Second, the earliest record for Lagurus curtatus in eastern Washington dates to about 8000 yr B.P. It is from the Granite Point Site (Gustafson, 1972) and represents an extralimital record. The nearest known extant population is about 50 km to the west of the site (Hall, 1981, p. 822). Gustafson (1972, p. 65) hypothesized that these specimens represented recent intrusive elements and reflected isolated edaphic conditions. Additional late Holocene extralimital records for this taxon are now known and include specimens from 50 km (Wawawai (Yent, 1976)) and 75 km (Pig Farm, 45AS80, Timothy’s Village (Lyman, 1976)) east of the nearest known extant population. All of these records for L. curtatus come from the canyon of the lower Snake River, an area that might be considered an edaphic, eastward extention of more arid and warmer environmental conditions and habitats similar to the Wallula Gap area (Daubenmire, 1970). During the last 3000 yr the relative abundance of L. curtatus seems to have remained stable in the Columbia Plateau FR (data from 45AD2 (Deaver and Greene, 1978)) while its abundance seems to have decreased in the eastern portion of the lower Snake River region at this time (data from Granite Point (Gustafson, 1972) and Timothy’s Village (Lyman, 1976)). This may reflect recent local extirpation of L. curtatus in the latter area. Recent mammal surveys in the lower Snake River Canyon failed to produce a record of L. curtatus in the vicinity of Granite Point or Timothy’s Village. Other members of Dalquest’s Great Basin fauna (Perognathus parvus and Reithrodontomys megalotis) were collected near Granite Point but not Timothy’s Village (Lewke and Buss, 1977). This suggests that late Holocene local extirpations of Great Basin taxa

LIVINGSTON

in the eastern lower Snake River Canyon may have been selective. Other recent mammal surveys (e.g., Rickard, 1960) indicate that L. curtatus is presently restricted to areas west of Granite Point, and that P. parvus, while widespread west of Granite Point, may only be found in restricted areas east of Granite Point. Finally, it can be noted that the earliest Antilocapra americana in the central Columbia Plateau FR dates to ca. 7500 yr B.P. (Meyer Cave; Bryan, 1955). This taxon is present in faunas in this FR until early historic times. The earliest reported Bison sp. in the central Columbia Plateau FR is from the Lind Coulee Site and dates to 8700 yr B.P. (Daugherty, 1956; Irwin and Moody, 1978). Bison are also present in faunas in this FR until early historic times. The available data concerning the xeric steppe taxa are insufficient to assess the accuracy of Dalquest’s (1948) suspicions concerning which taxa were present in eastern Washington prior to the last glacial age (Zoogeographic Group 1). Of the taxa that he inferred to have entered eastern Washington from the Great Basin after the last glacial age (Zoogeographic Group 6), three (0. leucogaster, S. townsendii, and Antilocapra americana) were present there ca. 13,000 yr B.P., and another (Neotoma cinerea) was present by 10,000 yr B.P. Bison might have been present throughout the last 20,000 yr. If so, this taxon should be moved from Zoogeographic Group 6 to Zoogeographic Group 1 (Table 1). The earliest radiametrically dated specimen of this taxon is 8700 yr old (Table 5). The zoogeographic records for S. idahoensis, U. cinereoargenteus, and L. curtatus are suggestive of a period of relatively warmer and drier environments during the middle Holocene. In particular, the apparent local extirpation of the latter two taxa in the easternmost portions of the SE Washington and Columbia Plateau FRs may be taken as evidence of the return to cooler and moister conditions inferred from the palynological record.

MAMMALIAN

ZOOGEOGRAPHY

The apparent zoogeographic extension of the ranges of Bison sp., A. americana, and S. townsendii from NE Oregon, the Yakima Valley FR, and the western part of the SE Washington FR during the late Pleistocene to the central Columbia Plateau FR during the early (bison and antelope) and late (S. townsendii) Holocene, respectively, may also reflect the warming and drying trend of the first part of the Holocene. No late Pleistocene faunas are recorded, however, in the central Columbia Plateau FR, so what we may, instead, be seeing is a reflection of sampling. SUMMARY

AND CONCLUSION

The late Quaternary mammalian zoogeographic history of eastern Washington in light of Dalquest’s (1948) postulated zoogeographic history and his environmentally distinctive faunas essentially conforms to the scenario of past environmental conditions inferred from botanical data. Some Boreal-me& steppe taxa like C. eluphus, 0. canadensis and M. americana had distributions different than their modern ones during the late Pleistocene, indicating cooler, moister environmental conditions than at present. The data for the Borealmesic steppe taxa possibly reflect more mesic conditions ca. 3500 yr B.P. as also indicated by the botanical data. Some of the xeric steppe taxa, like S. townsendii, Bison sp., and A. americana, also occupied ranges different than their modern ones during the late Quaternary. The zoogeographic histories of these taxa tend to reflect the general warming and drying trend of the early Holocene and hint at the return during the late Holocene to more mesic and cooler conditions as suggested by their disappearance from eastern Washington in general and the Columbia Plateau FR in particular. The mammalian zoogeographic history postulated by Dalquest (1948) can be refined on the basis of the available data. Taxa which Dalquest originally suggested were forced southward into eastern Wash-

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ington by expansion of glaciers must now include Rangifer and A. lagopus. The taxa which he suggested were present in eastern Washington during the last glacial age must now include Bison sp. Taxa which he suggested entered eastern Washington during the Holocene must now include U. cinereoargenteus, and, with the qualification that the taxon entered the Columbia Plateau FR, S. townsendii as well. ACKNOWLEDGMENTS Many people helped in the compilation of the data discussed here. In particular, Hal Kennedy helped gain access to obscure reports. An earlier. much different version of this paper was commented on by A. D. Bamosky, V. Butler, W. W. Dalquest, P. Ford, D. K. Grayson, G. T. Jones, R. Leonard, S. D. West, and two anonymous reviewers. We thank Dr. D. K. Grayson for his encouragement to complete this project and for reading several drafts of the paper.

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