ARCHAEOLOGICAL SEWELLEL (APLODONTIA RUFA) REMAINS ...

Journal of Mammalogy, 87(1):139–147, 2006

ARCHAEOLOGICAL SEWELLEL (APLODONTIA RUFA) REMAINS FROM DUNCAN’S POINT CAVE, SONOMA COUNTY, CALIFORNIA THOMAS A. WAKE* Zooarchaeology Laboratory, The Cotsen Institute of Archaeology at the University of California, Los Angeles, Los Angeles, CA 90095-1510, USA

Nine sewellel (Aplodontia rufa) bone specimens are identified in the Duncan’s Point Cave vertebrate archaeofauna. At least 3 individuals are represented and 3 specimens exhibit evidence of deposition in the cave as carnivore prey. The extralimital presence of A. rufa near the mouth of the Russian River demonstrates their occurrence over a wider area during the mid-Holocene and raises the possibility of the existence or recent extirpation of a 3rd isolated coastal Californian population of the species. Based on the presence of bones of A. rufa in the Duncan’s Point Cave archaeofauna and the availability of suitable habitat in the region, a detailed survey of the area surrounding the mouth of the Russian River is warranted. Key words:

Aplodontia, archaeology, California, Holocene, sewellel, taphonomy

logical perspective on local Holocene mammalian diversity. The implications of the occurrence of A. rufa in this assemblage are discussed with respect to known (and perhaps unknown) coastal California relict subspecific populations. Duncan’s Point Cave.— Recent archaeological excavations at Duncan’s Point Cave (archaeological site trinomial designation CA-SON-348/H), located on the southern Sonoma County coast of California, have recovered a diversity of artifacts, and include well-preserved invertebrate and vertebrate faunal remains (Schwaderer 1992; Wake and Simons 2000). Duncan’s Point is located approximately 8 km south of the mouth of the Russian River (Fig. 1). A large outcrop of sandstone bedrock contains several small overhangs and an ancient sea-arch cave known as Duncan’s Point Cave. This 12-m-deep natural cavern was formed during the midPleistocene, approximately 500,000 years ago, when a marine terrace about 25 m above the current shoreline was cut and wave action eroded through the exposed bedrock. The cave’s interior is almost completely filled with debris, containing a stratified archaeological deposit more than 3 m deep (Schwaderer 1992). The Duncan’s Point archaeological site is one of the largest prehistoric coastal sites located between the mouth of the Russian River and Bodega Head. Currently, it is the only known large cave site on the northern California coast between San Francisco and Cape Mendocino. Archaeological deposits are found both within and outside the cave and cover almost all of Duncan’s Point. Rae Schwaderer and a crew of California Department of Parks and Recreation personnel and volunteers excavated a 1.5-m square test unit to a depth of 317 cm during August 1989 (Schwaderer 1992).

The potential contribution of zooarchaeological study to understanding Holocene biogeography, human impacts on animal populations, and conservation biology should not be underestimated (e.g., Grayson 2001; Lyman 1996; Steadman 1995). For example, detailed analysis of Holocene vertebrate archaeofaunas from throughout the Great Basin has shed considerable light on environmental change and shifting mammalian distributions in the region (Grayson 1977, 1987, 2000; Lyman 1991a, 2004; Lyman and O’Brien 2005). Zooarchaeological analysis geared toward determining prehistoric presence or absence of mountain goats (Oreamnos americanus) in Olympic National Park, Washington, has served to spotlight the modern applicability of zooarchaeological data (Lyman 1988, 1994a, 1995, 1996, 1998; Lyman and Cannon 2004a). Results of zooarchaeological analyses have increasingly been applied in various modern conservation and wildlife management contexts throughout the United States (Lyman and Cannon 2004b, and contents). I hope this article has application in future wildlife management and conservation debates and studies concerning California’s coastal sewellel populations (Aplodontia rufa). I focus here on 9 archaeological sewellel specimens recovered from Duncan’s Point Cave and their implications. I include a brief overview of the mammalian assemblage from Duncan’s Point Cave in California to provide a zooarchaeo-

* Correspondent: [email protected]

Ó 2006 American Society of Mammalogists www.mammalogy.org 139

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Aves, and Mammalia) and sent the fish remains to J. Quinn and the bird remains to D. Simons for identification (Schwaderer 1992). I identified the amphibian, reptile, and mammal specimens (Schwaderer 1992; Wake and Simons 2000). I assigned identifiable bones to the most specific taxonomic level possible, and recorded information on skeletal element, body side, element configuration (complete, proximal, distal, medial, or fragment thereof), estimated age, and modification (burning, gnawing, fragmentation, butchering marks, and purposeful modification) for each element. I determined number of identified specimen counts for each mammalian taxon based on the number of separately identifiable specimens assignable to that taxon. I computed minimum numbers of individuals by counting the most abundant paired or unique skeletal elements for each taxonomic category, allowing for age, present within a given cultural component.

RESULTS

FIG. 1.—North-central California coast showing known historic ranges of relict subspecies of Aplodontia rufa and the location of Duncan’s Point Cave, Sonoma County, California (California archaeological site trinomial CA-SON-348).

Seven uncorrected radiocarbon dates from the cave range sequentially from 8,210 6 110 to 3,210 6 100 radiocarbon years ago (Schwaderer 1992). A possible Lake Mojave–style projectile point made of Annadel obsidian was recovered from the 240- to 250-cm level and dated to 8,210 6 110 radiocarbon years ago. Another important attribute of Duncan’s Point Cave is the presence of abundant, well-preserved vertebrate faunal remains, which sample the local biota over at least the last 8,000 years.

MATERIALS AND METHODS To summarize Schwaderer’s (1992) report on Duncan’s Cave, she placed a single 1.5-m square excavation unit in the cave and excavated it in arbitrary 10-cm levels. She passed sediment from the northeast quadrant of each level sequentially through a 6-mm (one-fourth–inch) mesh and a 3-mm (one-eighth–inch) mesh. She wet screened a 10% volumetric sample from each screen size fraction for each level. The entire excavation unit was excavated to a depth of 280 cm and in the northwest quadrant to a depth of 310 cm. Processing methods below the 270-cm level were altered because of a decrease in shell and an increase in clay. All excavated material below 270 cm was wet screened. Schwaderer sorted and separated the vertebrate remains from Duncan’s Point Cave by class (Osteichthyes, Amphibia, Reptilia,

A total of 1,438 mammal bones (73.4% of all identified vertebrate remains) were identified from Duncan’s Point Cave (Table 1), along with 203 elements from fish (10.4%), 5 from amphibians (0.3%), 1 from a reptile (0.1%), and 309 from birds (15.8%—Schwaderer 1992; Wake and Simons 2000). The 9 specimens of A. rufa are described below. Material examined.— Aplodontia rufa ssp., sewellel. Material.— Premaxilla with anterior maxilla (right), mandible (left and right); I1 (right); i1 (right); M3 (left); femur (left and right); tibia (left). Remarks.— The 9 identified specimens represent at least 3 individuals (Table 2). The right premaxilla is articulated with the anterior portion of the maxilla. The premaxilla contains the upper incisor minus the occlusal surface of the tooth. The associated anterior portion of the right maxilla includes the partial alveolus of the anterior peglike premolar. Both mandibles are nearly complete, missing only the superior, hooked portion of the coronoid process. The right mandible contains m2 and m3, and the left mandible contains m1, m2, and m3. Three isolated teeth include the right I1 and i1 and a left M3. The 2 femora are incomplete. The left femur includes the proximal portion and most of the shaft. The right femur is missing the distal articular surfaces. The tibia is incomplete, missing the proximal articular surface (Fig. 2). None of the specimens of A. rufa bear any cut marks or any sign of burning or other intentional modification by humans. The left mandible, except for the terminal portion of the coronoid process, which is easily broken, and the loose teeth do not appear modified in any way. The right mandible and the 3 incomplete long-bone specimens, 2 femora and 1 tibia, bear marks indicative of modification by carnivores. The tibia bears tiny, yet relatively wide, shallow, U-shaped grooves on the proximalmost and midshaft portions of the element. The right femur bears similar marks as well as distinctive pitting and furrowing near the broken distal end of the element (Fig. 3). The right mandible bears a series of small round punctures on the buccal surface of the angular process and a single larger puncture on the buccal surface of the alveolar portion of the element (Fig. 4).

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WAKE—DUNCAN’S POINT CAVE SEWELLEL

TABLE 1.—Number of identified mammal specimens at Duncan’s Point Cave, Sonoma County, California, by time of deposition as indicated by cultural components, with age in calibrated radiocarbon years before the present as follows: 1, undated; 2, 10,740–8,635 years ago; 3, 8,585–8,141 years ago; 4, 5,182–4,635 years ago; 5, 3,975– 3,432 years ago; 6, 3,309–2,825 years ago (Schwaderer 1992; Wake and Simons 2000). Cultural components Scientific name Sorex vagrans Scapanus latimanus Canis Canis latrans Enhydra lutris Mephitis mephitis Mustela frenata Taxidea taxus Procyon lotor Lynx rufus Carnivora Callorhinus ursinus Zalophus californianus Phoca vitulina Pinnipedia Otariidae Balaenopteridae Odocoileus hemionus Aplodontia rufa Spermophilus beecheyi Sciurus griseus

Common name

1

2

3

4

5

6

Total

Vagrant shrew Broad-footed mole

0 1

0 0

0 1

1 1

0 0

0 2

1 6

Dog Coyote Sea otter Striped skunk

0 0 0 0

0 0 3 0

1 0 8 1

1 5 4 0

0 0 4 0

1 1 14 4

3 6 33 5

Long-tailed weasel

0

0

2

0

1

1

4

American badger Raccoon Bobcat Carnivore Small carnivore Northern fur seal

0 0 0 0 0 0

4 0 0 0 1 0

1 0 2 0 2 6

0 0 0 0 2 17

0 0 0 0 0 3

0 1 0 3 0 19

5 1 2 5 3 45

California sea lion

0

1

1

2

5

2

11

Harbor seal Seals Eared seals Baleen whale Mule deer

0 0 0 0 0

0 0 1 0 0

0 2 2 0 2

18 4 2 0 5

2 0 2 0 7

2 2 6 1 2

21 13 8 1 16

Sewellel 0 1 1 2 3 2 9 California ground 0 0 0 0 0 4 4 squirrel Western gray 0 0 0 2 0 1 3 squirrel Thomomys Botta’s pocket 40 52 205 136 74 164 672 bottae gopher Neotoma Dusky-footed 0 0 0 2 4 3 9 fuscipes woodrat Peromyscus White-footed mice 0 3 0 15 9 13 42 Peromyscus North American 1 0 0 0 0 0 1 maniculatus deermouse Reithrodontomys Western harvest 0 0 0 1 3 3 7 megalotis mouse Microtus California vole 26 13 16 87 44 95 281 californicus Erethizon North American 0 0 1 4 0 0 5 dorsatum porcupine Rodentia Rodent 0 3 1 3 0 11 29 Small rodent 0 7 0 2 1 1 18 Lepus Black-tailed 0 0 0 0 0 4 4 californicus jackrabbit Sylvilagus Brush rabbit 1 15 40 21 5 21 103 bachmani Mammalia Large mammal 0 1 9 6 2 6 24 Medium mammal 0 0 8 1 3 0 12 Small mammal 0 4 8 13 0 0 25 Total 69 108 320 357 172 390 1,438

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TABLE 2.—Specimens of Aplodontia rufa from Duncan’s Point Cave, Sonoma County, California. Time of deposition indicated by cultural component (CC), as in Table 1. Side of element shown as left (L) or right (R). Accession numbera

Levelb

CC

Element

P913-6-7 P913-6-7 P913-11-4 P913-10-2 P913-13-10

2050 2050 6070 6070 7080

6 6 5 5 5

Mandible Premolar Femur Mandible Tibia

R L R L R

P913-17-5

90100

3

P913-17-5

90100

3

P913-19-6 P913-50-11

100110 240250

3 2

Lower incisor Upper incisor Femur Premaxilla

Side

Age

Portion Complete Complete Proximal Complete Distal

R

Adult Adult Adult Adult Young adult Adult

L

Adult

Complete

L R

Adult Adult

Proximal Most

Complete

a Specimens are deposited at the California Department of Parks and Recreation curation facility in West Sacramento, California. b Level ¼ cm below surface.

The relatively long, straight marks are strongly reminiscent of oblique carnivore gnaw marks (Blumenschine 1995; Blumenschine et al. 1996; Haynes 1980; Lyman 1994b; Selvaggio 1994; Shipman 1981). The pitting is equally reminiscent of those marks associated with carnivore crushing, chewing, and furrowing (Blumenschine 1995; Blumenschine et al. 1996; Brain 1981; Hill 1989; Lyman 1994b; Maguire et al. 1980; Selvaggio 1994). The punctures on the mandible do not pierce the specimen completely, are relatively small in diameter (1 mm), and strongly resemble punctures associated with carnivore modification (Lyman 1994b:209–210). The linear marks and abrasions could represent taphonomic processes not related to the modification of bone by animals, such as trampling and compaction (Olsen and Shipman 1988). The proximal left femur fragment appears to have passed through a predator’s gut. The specimen bears distinctive indications of acid etching and digestion. The broken edges of the cortical bone are smoothed and come to a gradually thinned edge (Fig. 5). Further, the articular surface of the femoral head is dissolved away, revealing the honeycombed cancellous tissue beneath (Fig. 5). Both of these characteristics, the thinned edge and exposed cancellous tissue, are strikingly similar to those described by various authors as appearing on small mammal bones passed through predator guts (Andrews 1990:64–88; Lyman 1994b; Rensberger and Krentz 1988; Schmitt and Juell 1994; Schmitt and Lupo 1995). Hill (1989) describes similar bone thinning on specimens passed through hyenas.

DISCUSSION A brief natural history of A. rufa.—Aplodontia rufa was 1st brought to the attention of European zoologists by the Lewis and Clark expedition (Godin 1964; Lewis and Clark 1814). Originally described by Rafinesque in 1817, Aplodontia is a monophyletic, monotypic genus. The most recent widely accepted classification of Aplodontia includes a single species

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FIG. 2.—Archaeological remains of Aplodontia rufa from Duncan’s Point Cave showing (top row) right femur, proximal portion of left femur, distal portion of left tibia, left M3, left and right lower incisors, above, and right premaxilla and maxilla, below, and (bottom row), left and right mandibles.

(A. rufa) and 7 subspecies, 6 of which are found in California (Carraway and Verts 1993; Hall 1981; Steele 1989). The 2 populations of A. rufa closest to Duncan’s Point Cave are isolated relict populations confined to the California coast (Fig. 1). A. rufa nigra is found in a 62-km2 area at Point Arena, Humboldt County, and A. r. phaea is found in a 285-km2 area around Point Reyes, Marin County (Carraway and Verts 1993; Steele 1989). Kurteń and Anderson (1980) note that fossil remains of Aplodontia are known only from Wisconsinan era deposits in 3 northern California caves: Samwel (sic, correct spelling is Samuel) and Potter Creek (both in Shasta County) and Hawver (Eldorado County). These sites are all well inland from the coast and within the present range of the genus. The current range of A. rufa includes the Pacific Northwest from southern British Columbia south to California along the coast, the Cascades, and most of the Sierra Nevada including Sierran western Nevada (Carraway and Verts 1993; Godin 1964; Hall 1981). The current range of A. rufa overlaps with the ranges of several other evolutionarily basal, monogeneric vertebrate families and their representative species that are confined to a similar area including Ascaphus truei (Ascaphidae: Anura: Amphibia), Dicamptodon (Dicamptodontidae: Caudata: Am-

phibia), and Rhyacotriton (Rhyacotritontidae: Caudata: Amphibia—Good and Wake 1992; Stebbins 1985). Aplodontia rufa does not range far from its burrow system entrance. Radiotelemetry studies of 10 adults showed that home ranges varied between 0.03 and 0.2 ha, averaging 0.1 ha (Martin 1971). Maximum movement from a nest burrow was 42.5 m. Individuals were recorded within 24.4 m of their nest site about 90% of the time. Martin (1971) found that subadults in Oregon traveled between 183 and 549 m from their initial capture points. Aplodontia rufa has notoriously inefficient kidneys, a condition that necessitates the constant availability of drinking water or water-rich plant material to avoid dehydration (Carraway and Verts 1993; Fisler 1965; Johnson 1971; Steele 1989). The inability of A rufa to concentrate urine and its high demand for metabolic and drinking water may account for its limited distribution, especially in California (Carraway and Verts 1993; Nungesser and Pfeiffer 1965; Steele 1989). The introduction of small mammals into Duncan’s Point Cave.— Remains of burrowing insectivores and rodents (moles, sewellel, ground squirrels, gophers, and voles) comprise a significant portion of the mammalian assemblage from Duncan’s Point Cave (n ¼ 691; 48%), and occur throughout the deposits. The abundance of burrowing insectivores and

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FIG. 3.—Close-up of broken distal end of archaeological Aplodontia rufa left femur, showing evidence of carnivore modification.

rodents most likely is a consequence of the site’s location within a grassland–coastal scrub mosaic, prime habitat for these animals, a situation also observed along the central Mendocino County coast (Simons 1990). Various archaeological studies of bioturbation at open (noncave) archaeological sites in California have noted that gopher burrowing produces stratigraphic distributions of materials within archaeological sites entirely unrelated either to prehistoric human behavior or natural alluvial, colluvial, or aeolian processes (Bocek 1986, 1992; Erlandson 1984; Erlandson and Rockwell 1987; Johnson 1989). The frequency of burrowing mammal remains at Duncan’s Point Cave suggests that bioturbation may have affected site structure, as is suggested at other sites along the northern California coast (Simons 1990). Keeping the possibility of bioturbation by humans and rodents in mind, the stratigraphic integrity of the site is excellent inside the cave itself. Well-defined stratigraphic breaks were recorded in the profile drawing of the excavation unit (Schwaderer 1992). I observed finely stratified undisturbed features such as small hearths in cross section in the walls of the single excavation unit. Only the upper levels of the site (Component 6) have been disturbed by looters and others. In fact, ongoing looting and disturbance of the site were the prime motivating factors that resulted in excavation at the cave in 1989 and its subsequent increased protection. Two observations suggest secondary introduction of many of the small mammal remains into the cave by nonhuman predators. The 1st is the fact that finely stratified features are not preserved in deposits subjected to extensive bioturbation. Second, there is little reason for small mammals to burrow into the cave sediments and cause bioturbation, especially in the area of the excavation unit—well behind the drip line in an area where no plant foods grow. The cave does provide potential roosts and shelter for owls, bobcats, coyotes, and other predators that could introduce rodent remains into the site. The condition of at least 3 of the specimens of A. rufa strongly indicates modification and consumption by predators and subsequent deposition in the deposit (Fig. 3). The possibility of the introduction of rodent remains into the cave sediments by predators and not necessarily by the

FIG. 4.—Close-up of angular process of right mandible of archaeological Aplodontia rufa, showing tooth punctures made by a small carnivore.

rodents themselves makes the exact postdepositional impact of bioturbation upon the site somewhat unclear; however, the overall effects are probably minimal. Past environments.— Erlandson and Moss (1996) point out that the archaeological record of coastal California and much of Pacific North America is complicated by sea level rise, inundation, subsidence, isostatic rebound, erosion, and tectonic activity, all of which may have affected past coastal distributions of A. rufa. When correlated with the early archaeology of the San Francisco Bay Area (Bickel 1978; Moratto 1984) and the

FIG. 5.—Close-up of proximal femur of archaeological Aplodontia rufa, showing effects of digestive action (acid etching and thinning on broken surface of shaft).

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Mendocino County coast (Simons et al. 1984), Holocene sea level rise is viewed as having profound environmental consequences. As for Duncan’s Point Cave, the shoreline may have been approximately 14 km west 15,000 years ago. By 10,000 years ago the shoreline was about 5 km west and at 5,000 years ago, about 1 km west (Atwater 1979). During the last 6,000 years shoreline retreat in California has resulted from coastal erosion ranging from 15 cm/year along much of the central California coast (Gordon 1979), 30 cm/year near Santa Cruz and on the San Mateo County coast (Griggs and Johnson 1979; Sullivan 1975), to 9 cm/year around Fort Ross (Ritter 1978). Palynological data from various North Coast localities suggest development of a more-or-less modern climatic regime occurred about 10,000–7,500 years ago on the Point Reyes Peninsula (Rypins et al. 1989). Before the terminal Pleistocene a more boreal plant community dominated by Abies and Pseudotsuga was present near Point Reyes. Studies of Pleistocene plant macrofossils from northern and central California indicate a closed-cone pine community dominated by Bishop pine (Pinus muricata) and Monterey pine (Pinus radiata) continuously distributed along much of coast from 14,500 years ago to 12,000 years ago (Axelrod 1981, 1983; Johnson 1977; Simons et al. 1984). A more complex mosaic of plant associations replaced the pine-dominated botanical communities of the terminal Pleistocene northern California coast by the mid-Holocene (Adam 1985; Adam et al. 1981; West 1990, 1993). Russell (1983) notes that during the last millennium, major vegetation changes have taken place near Wildcat Lake on the Point Reyes Peninsula. The exact causes of these changes are unknown, but likely sources include localized factors such as fire and topographic disturbances (landslides and slumps), and more general factors such as climatic change (i.e., the ‘‘Medieval Warm Epoch Drought’’—Graumlich 1993; Raab and Larson 1997; Stine 1994). Fire has been shown to have particularly devastating effects on local sewellel populations in Point Reyes National Seashore (Fellers et al. 2004, 2005). The combination of physical geographic change, general climatic change, and continuing fragmentation of expansive boreal forests at the end of the Pleistocene into the Holocene contributed to the isolation of the 2 southernmost coastal California subspecies of A. rufa (A. r. nigra and A. r. phaea). These same conditions that fragmented the California coastal range of A. rufa could easily have resulted in the isolation of other relict populations during the Early to Middle Holocene. Aplodontia rufa and the northern California coastal environment.— Because of its antiquity, Duncan’s Point Cave is an excellent location for the study of Holocene environmental and cultural change, and the effects those factors have had upon the presence of animals. Because of continuous deposition throughout much of the Holocene, the effects of local sea level rise, coastal erosion, and general climate change on the distribution of marine and terrestrial habitats are recorded in the Duncan’s Point Cave vertebrate and invertebrate assemblages (Schwaderer 1992; Wake and Simons 2000). The occurrence of A. rufa in Californian archaeofaunas is rare at best. Sewellel are not identified in several archaeofaunas from

California’s north coast and Oregon well within the current ranges of A r. humboldtiana and A. r. pacifica (Hall 1981; W. R. Hildebrandt, in litt.; Hildebrandt 1981, 1984; Hildebrandt and Levulett 1997; Jameson and Peeters 1988; V. A. Levulett and W. R. Hildebrandt, in litt.; Lyman 1991b). It is not likely that a bobcat (Lynx rufus), the most common predator of A. rufa in Washington and Oregon (Knick 1984; Nussbaum and Maser 1975; Sweeney 1978), or an American mink (Neovison vison) could or would transport a carcass very far. Therefore, the sewellel individuals represented in this assemblage were most likely captured nearby. Based on proximity alone, the Duncan’s Point Cave specimens may represent members of a broader population of A. r. phaea that once extended farther to the north of the Point Reyes region, perhaps to the lush hillsides near the mouth of the Russian River. It is also possible that the Duncan’s Point Cave A. rufa represent a southerly extension of A. r. nigra. Alternatively, the Duncan’s Point Cave specimens of A. rufa might represent another distinct, isolated coastal population that may, or may not, have disappeared some time during the late Holocene. Aplodontia rufa does not disperse far. Rather it has high site fidelity, high available water demands, and prefers lush cover. A. rufa could not disperse across the more diverse range of plant communities including the drier coastal prairie or northern coastal scrub (a xeric plant community dominated by Mimulus, Baccharis, Lupinus, Toxicodendron, and Rubus—Schoenherr 1992) environments surrounding Duncan’s Point now and for the past few thousand years (e.g., West 1990, 1993). The most likely modern location for a sewellel population near Duncan’s Point is the last north-facing escarpment near the mouth of the Russian River, across the river from the town of Jenner, largely within the boundaries of Sonoma Coast State Beach. This area is lush, shady, relatively steep and inaccessible, and covered by a large stand of coastal Douglas-fir (Pseudotsuga menziesii) and coast redwood (Sequoia sempervirens) forest. These slopes currently represent the nearest island of habitat to Duncan’s Point that is similar to the habitats found at Point Arena and Point Reyes, the locations of the 2 known relict populations of A. rufa. However, no collection or observational records are known to exist for the area (Hall 1981; Steele 1989). Given the cursory survey treatment of this general area, it is entirely possible that a population of A. rufa could remain undetected. Further research.— The Duncan’s Point Cave specimens of A. rufa are geographically closer to the Point Reyes subspecies, raising the possibility that the mid-Holocene geographic range of A. r. phaea once extended to the Russian River. If wellpreserved DNA can be extracted from the Duncan’s Point archaeological specimens of A. rufa and be sequenced, it may be possible to determine whether the Duncan’s Point Cave population is actually more closely related to A. r. phaea at Point Reyes, to A. r. nigra at Point Arena, or represents a distinct, extant or recently extirpated subspecies.

ACKNOWLEDGMENTS I thank R. Schwaderer, B. Parkman, and the California Department of Parks and Recreation for the opportunity to participate in the

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excavation of Duncan’s Point Cave and to analyze the recovered mammalian and herpetological remains. C. Campbell, J. Patton, D. Simons, D. B. and M. H. Wake, and 2 anonymous reviewers provided insightful comments that greatly improved earlier versions of this paper. Comparative osteological collections in the Museum of Vertebrate Zoology, University of California, Berkeley, and the Zooarchaeology Laboratory, University of California, Los Angeles, facilitated identification of the Duncan’s Point Cave vertebrate archaeofauna.

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Submitted 29 April 2005. Accepted 7 July 2005. Associate Editor was Carey Krajewski.