Terminal Pleistocene change in mammal communities in southeastern Washington State, USA

Quaternary Research 81 (2014) 295–304

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Terminal Pleistocene change in mammal communities in southeastern Washington State, USA R. Lee Lyman ⁎ Department of Anthropology, 107 Swallow Hall, University of Missouri, Columbia, MO 65211, USA

a r t i c l e

i n f o

Article history: Received 29 May 2013 Available online 22 December 2013 Keywords: Climate change Columbia Basin Conservation paleozoology Eastern Washington state Mammals Paleoecology Pleistocene–Holocene transition

a b s t r a c t Small mammal communities in western North America experienced declines in taxonomic richness across the late Pleistocene to Holocene transition (PHT), a recent natural global warming event. One community also experienced a decline in evenness and others replaced one species with a congener. Variability in response of small mammal communities to PHT warming is apparent. At the presently arid and xeric Marmes site in the Columbia Basin of southeastern Washington State, megafauna were absent by about 13,000 cal yr BP, evenness of small mammals declined about 11,700 cal yr BP and again about 11,400 cal yr BP whereas richness declined about 11,400 cal BP. Regional faunal turnover was, however, minimal among small-bodied taxa. Local mammal communities are depauperate as a result of megafaunal extinctions and subsequent decreases in small-mammal richness and evenness. The latter chronologically corresponds with a decrease in primary productivity driven by increasing warmth and aridity. More faunas must be studied in order to fully document the range of variability in the responses of mammalian communities to PHT warming. Documentation of patterns in those responses will facilitate understanding and enhance predictive accuracy with respect to responses of mammalian communities to modern global warming. © 2013 University of Washington. Published by Elsevier Inc. All rights reserved.

Introduction Comparisons of 50–100 yr old North American mammal samples housed in museums with recent samples of mammals collected from the same locale have shown that modern global warming typically causes up-slope shifts in range and some restructuring of community composition (e.g., Moritz et al., 2008; Rowe et al., 2010). The Pleistocene-to-Holocene transition (PHT, hereafter) centering around 11,700 cal yr BP (Walker et al., 2009) represents a period of natural warming that spanned several millennia rather than the single century represented by most natural-history collections of modern mammalian communities. Study of the long-term trends revealed by paleo-mammal communities may reveal changes of greater magnitude than those reflected by museum collections; rates of change and range of flux undocumented in the short term may be revealed by the relatively longer duration represented by temporal series of collections of prehistoric remains; and historically undocumented kinds of change may also be revealed (Barnosky et al., 2003). Further, samples of historically collected mammals may reflect modern industrial era or agricultural influences (e.g., Lyman, 2012d) that could obscure or skew the global warming signal. Samples of prehistoric remains dating to the PHT are not subject to such influences, and while North America's first human colonists (Paleoindians) may have impacted faunas (e.g., Grayson, 2006; Surovell, 2008; Haynes, 2009), this is not as yet well established ⁎ Fax: +1 573 884 5450. E-mail address: [email protected].

(Grayson and Meltzer, 2002, 2003). Whatever the case, fidelity studies indicate that prehistoric mammal faunas often provide minimally biased samples of living faunas (Terry, 2010a,b). Alternatively, close study of the accumulation, deposition, and preservation history of collections of prehistoric remains—taphonomic analysis—often reveals biases and how they can be analytically controlled or corrected (e.g., Kidwell, 2013). Modern declines in biodiversity are a major concern among conservationists, restoration ecologists, and wildlife managers (Dawson et al., 2011). A suspected cause of many of these declines is global warming, whether anthropogenically driven or not (Botkin et al., 2007; Barnosky, 2009). Much research on biodiversity loss concerns extinction of taxa, but extinction is typically preceded by decline in a taxon's population and concomitant shifts in the taxonomic composition of and taxonomic abundances comprising local communities (e.g., Guthrie, 1990; Barnosky et al., 2003; Brace et al., 2012). These shifts in turn may influence ecosystem function (Naeem et al., 1994; Wittebolle et al., 2009; Steudel et al., 2012). An increasing number of studies have demonstrated the value of collections of prehistoric animal remains for revealing the long-term effects of significant climatic change on mammalian communities (Lyman and Cannon, 2004; Dietl and Flessa, 2009; Terry et al., 2011; Lyman, 2012a; Wolverton and Lyman, 2012). As noted long ago, such collections represent the results of ecological experiments that are difficult if not simply impossible to replicate today (Deevey, 1969). Several previous studies have examined late Pleistocene and early Holocene mammalian faunas from western North America (Walker,

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1987; Grayson, 2000; Hockett, 2000; Schmitt et al., 2002; Grayson, 2005; Schmitt and Lupo, 2005; Blois et al., 2010; Schmitt and Lupo, 2012). These studies indicate local mammalian communities changed during the PHT, and importantly, they indicate that the details of change vary from locale to locale. Some studies indicate decreases in richness (Walker, 1987; Grayson, 2000; Blois et al., 2010; Schmitt and Lupo, 2012), some indicate replacement of one species by a congener (Neotoma spp. (Grayson, 2000); Thomomys spp. (Blois et al., 2010; Schmitt and Lupo, 2012); Dipodomys spp. (Schmitt and Lupo, 2005)), one indicates decreased evenness (Blois et al., 2010), and several indicate shifts in the range of some taxa (e.g., Hockett, 2000; Grayson, 2005). Variability in habitats and taxonomic composition of local communities likely account at least partially for these differences (Schmitt and Lupo, 2012). Whatever the cause of the variability, it is clear that it would be ill advised to argue that trends evident in one or two or three faunas reveal all the nuances and particularities of every instance of change. While we now recognize some of the general features of faunal change during the PHT, we are only just beginning to learn the types and range of variability in those changes. The interior Pacific Northwest, the Columbia Basin of eastern Washington state in particular, has until now not been included in discussions for want of data. Here I describe a collection of small mammal remains recovered 45 yr ago from the Marmes archaeological site in southeastern Washington, USA (Fig. 1). When excavated in 1968, the Marmes site produced the then oldest known human remains on the North American continent. The mammalian faunal collection comprises several chronometrically unique communities that span the

PHT and document the effects of warming and drying climate on local mammal community composition, richness, and evenness. The Marmes site provides another data point to facilitate pattern identification among the variability apparent in how mammalian communities responded to the PHT, and also specification of the possible variation in mammalian community response to modern global warming. Paleoenvironmental background Late Pleistocene paleoenvironmental data such as that derived from fossil pollen and plant macrofossils are not as abundant for the Columbia Basin of eastern Washington state (Fig. 1) as they are in the Great Basin (Nevada and western Utah) for two reasons. First, the northern half of the Columbia Basin was glaciated until about 15,000 yr ago. Second, much of the southern half of the Columbia Basin was scoured multiple times by catastrophic floods the waters of which originated behind glacial dams, the last flood occurring about 15,000 yr ago (references in Chatters, 1998; Wigand and Hicks, 2004). As a result, disagreement exists over some details of what all agree was a climatically dynamic period. But most agree the PHT between 15,000 and 10,000 cal yr BP (calibrated ‘calendar’ yr before present) was a time of climatic warming, with evidence for an abruptly initiated period of cold and seemingly moist conditions about 13,200 cal yr BP, and then an equally abrupt reversion back to the warming trend about 12,200 cal yr BP (Mehringer, 1996; Wigand and Hicks, 2004). This cool-moist period “corresponds to the Younger Dryas Period of northern Europe” (Wigand and Hicks, 2004:55). One of the temporally distinct faunas I describe

Figure. 1. Location of the Marmes site in eastern Washington state, USA.

R.L. Lyman / Quaternary Research 81 (2014) 295–304

here encompasses the end of this cool-moist interval and the two younger faunas represent the earliest Holocene (Fig. 2). Thus the oldest fauna (A6–16) should reflect the cool-moist event, and the youngest fauna (Marmes horizon) should reflect the subsequent warmer era. The mid-aged fauna (Harrison horizon) should reflect the transition period. Together the three faunas provide a forecast of what could happen among mammal communities as global warming continues. The shift from the cool-moist period to the warming and drying trend was abrupt, perhaps as brief as 60 yr in the North Atlantic (Steffensen et al., 2008). In the interior Pacific Northwest of North America (Fig. 1), the pollen record suggests the local shift was nearly as abrupt (Johnson et al., 1994; Mehringer, 1996). Aridity and warmth continued to increase but more gradually after the abrupt shift to warming and drying (Mehringer, 1996; Chatters, 1998; Wigand and Hicks, 2004; Huckleyberry and Fadem, 2007). During the cool–moist phase vegetation in the Marmes site area was composed of cool sagebrush steppe with scattered patches of mesic adapted conifers (haploxylon pollen); grass was relatively abundant. About 12,200 cal yr BP, the abundance of grass and sagebrush increased relative to that of conifers (absolute abundances likely decreased) and mesic-adapted conifers were replaced by xericadapted forms (diploxylon pollen) that were of limited abundance (Mehringer, 1996; Wigand and Hicks, 2004). Climate models suggest that the interior Pacific Northwest including the Columbia Basin will continue to experience the Holocene warming trend documented in paleoecological records (Christensen et al., 2007). Paleoecological data for the Columbia Basin in which the Marmes site is situated suggest the PHT not only warmed, but conditions also

Figure. 2. Chronology of Marmes site analytical units and related paleoenvironmental and geological events. U-I is within the rockshelter; A6–16, Harrison, and Marmes analytical units make up the floodplain deposits. Graphed range per analytical unit is based on the range indicated by calibrated radiocarbon ages (Table 1). The bold line through the temporal range for an analytical unit is the median age. Correspondence of calibrated and 14C age scales is approximate.

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become more xeric. The time-series of mammalian faunas at the Marmes site should reflect a trend of increasing temperature and decreasing effective moisture. Methods and materials The Marmes site (45FR50) was excavated in 1962–64 and again in 1968 (Fryxell and Daugherty, 1962; Fryxell and Keel, 1969; Rice, 1969). The site consists of two depositionally and taphonomically distinct areas—Marmes Rockshelter and the floodplain adjacent to and ~10 m below the mouth of the rockshelter (Hicks, 2004). Gustafson (1972) analyzed a sample of the mammal remains recovered from the rockshelter; the sample represented the terminal Pleistocene and the entire Holocene in a series of eight strata. Caulk (1988) studied about five percent of the mammal remains recovered from the floodplain; those remains were distributed across two strata representing the terminal Pleistocene and earliest Holocene. Gustafson and Wegener (2004) studied an additional 10% of the mammal remains from the floodplain, also representing the terminal Pleistocene and earliest Holocene. These earlier analyses found that the mammalian fauna dating to the PHT suggested climate at that time was cooler and more moist than today. Taphonomy Gustafson and Wegener (2004) compared the sample of mammalian remains from the 1962–64 excavation of the rockshelter identified by Gustafson (1972) with the mammalian sample from the 1968 excavation of the floodplain identified by Caulk (1988) combined with the sample of vertebrate remains from the floodplain they identified (Gustafson and Wegener, 2004). In particular, they compared the mammalian fauna from rockshelter strata U-I and U-II (NISP = 71), with materials from the Marmes and the Harrison horizons of the floodplain (NISP = 672) because it seemed to them that “the basal portion of U-I and U-II corresponds in time with the floodplain horizons” (Gustafson and Wegener, 2004:290). Based on more radiocarbon dates than available to Gustafson and Wegener (2004), we now know that rockshelter strata U-II was deposited after the floodplain deposits of concern here were deposited (Hicks, 2004; Huckleyberry and Fadem, 2007). Thus in the comparisons that follow, fauna from U-II are not included unless Gustafson and Wegener's (2004) comparisons are being described. Gustafson and Wegener (2004) observed that the fauna from the rockshelter includes more remains of artiodactyls (Cervidae, Bovidae) than the floodplain fauna (39.3% of NISP relative to 27.2% of NISP, respectively). A portion of this difference is explained by the fact that rockshelter sediments producing the faunal remains were screened with 0.25 in. (6.35 mm) mesh screens whereas all floodplain sediments were screened with 1 mm mesh screens; some of the rockshelter sediments were later screened with the smaller mesh. Recovery of microfauna from both was ultimately quite good (Lyman, 2012b). As a result, when all recovered and identified remains are considered rather than the small samples Gustafson and Wegener (2004) compared, the floodplain has a significantly higher proportion of small mammal remains (taxa with adult body mass ≤ 1 kg; bats excluded) than the rockshelter (91.69%, and 85.45%, respectively; arcsin transformation ts = 5.5, p b .0001). When all remains are considered, artiodactyl remains make up 4.3% of the rockshelter collection but only 0.4% of the floodplain collection (arcsin transformation ts = 8.1, p b .0001). Ascertaining why this difference occurs is beyond my scope here, but it suggests interpreting differences between the U-I fauna and the floodplain fauna in paleoecological terms is not advisable. Gustafson and Wegener (2004:294) also observed that the fauna from the rockshelter included remains of only “15 taxa” whereas remains from the floodplain represented “57 taxa.” The difference they observed is in part the result of sample size differences; they compared the taxa represented by 999 specimens from the floodplain with 112

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specimens from the rockshelter. In light of this difference in sample size, it is no surprise that the former is taxonomically richer than the latter. But Gustafson and Wegener also tallied taxa in the floodplain collection in such a way that some taxa were tallied twice; for instance, Neotoma sp. and Neotoma cinerea were tallied as two taxa rather than one. And, Gustafson and Wegener (2004) tallied amphibians, birds, and reptiles from the floodplain. Remains of 21 taxa of birds, two species of fish, a snake and an amphibian from the rockshelter and reported by Gustafson (1972) were not included in Gustafson and Wegener's (2004) tally of taxa for the rockshelter. If the total mammalian samples from rockshelter U-I and from the floodplain are considered, and the technique of tallying taxonomic richness is the same for both, the rockshelter includes remains of 25 unique mammalian taxa (genera and species) and the floodplain produced remains of 26 unique mammalian taxa (Table 2). Raptors, quadrupedal carnivores and humans accumulated and deposited many of the remains in the rockshelter; humans and overbank fluvial processes accumulated and deposited many of the remains in the floodplain sediments (Lyman, 2010). Many of the rodent remains from the floodplain appear to have been lightly charred, perhaps by wild fires that swept across the shrub-steppe vegetation believed to have occupied the floodplain. Some of the floodplain remains were fluvially deposited whereas others were accumulated and deposited by humans (Lyman, 2012c). Faunal remains from the two deposits share certain features. The Bray–Curtis Index of Dissimilarity (Magurran, 1988; see below) for the two faunas is 0.544, suggesting the two are fairly similar in terms of taxa represented and in terms of taxonomic abundances. Further, although impossible to demonstrate with available data, it appears that various remains from the rockshelter were secondarily deposited downslope on the floodplain by various agents that excavated in the rockshelter sediments (e.g., badgers [Taxidea taxus]). Humans (Homo sapiens), for example, used the rockshelter as a crematorium during the deposition of stratum U-I (Hicks, 2004), and some of

the burned human remains ended up in floodplain deposits (Krantz, 1979). Refitting and conjoining of fragments cannot be undertaken to test this suggestion because the human remains from the deposits have been repatriated. In light of the similarities and differences between the rockshelter and the floodplain deposits, analyses of change here focus on the latter. Analyses presented elsewhere of body mass distributions of both faunas combined indicate the Marmes PHT mammalian fauna represents a community that is distinct from the modern local fauna (Lyman, 2013b). Here I am concerned with determining if the fine temporal resolution provided by the floodplain strata reveals instances of change in taxonomic richness and evenness in the local mammalian community. The temporally coarse-grained rockshelter U-I fauna is included in a few analyses for comparative purposes. A potential cause of any observed faunal change evidenced by remains from an archaeological site such as the Marmes site is, of course, change in human faunal procurement strategies or technology. There is nothing in the faunal remains to suggest change in human procurement strategies. Further, one sample of human-exploited remains from the floodplain is so small that comparison of it with faunas from other floodplain strata is tenuous (Lyman, 2013a). In his study of the Windust cultural phase, which was originally defined in large part on the basis of material from the Marmes site's U-I and floodplain deposits, Rice (1972:135) noted that “there is essentially no difference between [what he termed] the Early Windust subphase [artifact] assemblage and the Late Windust subphase assemblage.” Later study of artifact material from U-I and the floodplain noted differences between the two but did not stratigraphically distinguish artifact assemblages from the floodplain (Ozbun et al., 2004). In short, there is no evidence for change in mammal procurement strategies or technology during the accumulation and deposition of the faunal remains discussed here, but that may be an artifact of how materials have typically been analyzed, that is, as a single temporal unit (e.g., Ames, 1988; Ozbun et al., 2004). This

Table 1 Chronological data for Marmes site faunal assemblages. Individual ages within analytical units are arranged chronologically (unless otherwise noted) and may be out of order stratigraphically. Analytical unit14C age BP

Laboratory ID

Dated material

Cal yr BP 2 sigmaa

Reference

Marmes horizon 9710 ± 40 9820 ± 300 9870 ± 50 9970 ± 110

Beta-156699 W-2209 Beta-120802 Y-2481

Bone collagen Mussel shell Bone collagen Mussel shell

11,227–11,081 12,223–10,478 11,396–11,198 11,827–11,206

Hicks (2004) Fryxell and Keel (1969) Hicks (2004) Fryxell and Keel (1969)

Harrison horizon 9840 ± 300b 10,130 ± 300c

W-2212 W-2218

Charcoal Charcoal

12,229–10,493 12,609–11,067

Fryxell and Keel (1969) Fryxell and Keel (1969)

A6–16 9900 ± 50d 10,270 ± 120e 10,570 ± 70f 10,830 ± 60g

Beta-296119 Beta-192237 Beta-156697 Beta-310319

Charred material Sediment Charcoal Charred material

11,409–11,208 12,427–11,601 12,667–12,378 12,883–12,590

Lyman (2013a) Huckleyberry and Fadem (2007) Hicks (2004) Lyman (2013a)

U-I 9380 ± 70 9610 ± 40 10,475 ± 270 10,750 ± 300 10,810 ± 300 11,230 ± 50h

Beta-301318 Beta-168491 WSU-366 WSU-211 WSU-363 Beta-156698

Charred material Mussel shell Mussel shell Mussel shell Mussel shell Bone collagen

10,786–10,386 11,040–10,774 12,876–11,384 13,283–11,750 13,315–11,946 13,280–12,945

Lyman (2013a) Hicks (2004) Chatters (1968), Fryxell et al. (1968) and Rice (1969) Chatters (1968), Fryxell et al. (1968) and Rice (1969) Chatters (1968), Fryxell et al. (1968) and Rice (1969) Hicks (2004)

a b c d e f g h

Ages were calibrated using Calib version 6.0 (Reimer et al., 2009). Composite sample from strata A3, A4, and A5. Composite sample from strata A3 and A4. Composite sample from strata A6, A7, and A8. Composite sample from strata A9 and A10. Composite sample from strata A10–A16. Composite sample from strata A13–A16. Dated sample lay on bedrock at the bottom of the stratigraphic column.

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precludes the detection of cultural change as a possible cause of faunal change. Chronology Analyses here focus on mammalian communities represented in floodplain sediments; the deepest deposits within the rockshelter are for the most part contemporaneous with the combined floodplain deposits (Fig. 2). Individual strata found in the rockshelter and in the floodplain deposits are lumped together into “analytical units” here because artifacts and faunal remains were not consistently recovered from individual strata during excavation. Multiple radiocarbon ages for each analytical unit (Table 1) were calibrated to calendar years using Calib version 6.0 and the IntCal09 curve (Reimer et al., 2009). Unfortunately, with few exceptions, precise vertical and stratigraphic provenience information for individual radiocarbon samples from an analytical unit is not available (Fryxell and Keel, 1969; Sheppard et al., 1984, 1987; Hicks, 2004). Thus the possibility of mixing of materials from the three floodplain analytical units (see below) cannot be evaluated using vertical or stratigraphic provenience of radiocarbon ages. Other data suggest minimal inter-analytical unit mixing. First, the numerous very detailed stratigraphic profiles of the floodplain deposits suggest minimal vertical movement of materials based on the integrity of strata boundaries (www.archaeology.wsu.edu/county/ franklin/45FR50/Appendices/AppendixB.html). Further, the four dates from the stratigraphically deepest floodplain analytical unit (A6–16, see below) are in perfect stratigraphic order (Table 1); greater ages are in deeper strata. The relative chronological order of the three floodplain analytical units is assured by their superimposed positions. For discussion purposes, the median calibrated age for each floodplain analytical unit is used, but I recognize that the duration of the deposition of the faunal materials included in an analytical unit may extend several hundred years before and after that median age (Fig. 2). Lumping of strata into analytical units was often done in the field, and was done for the deepest floodplain analytical unit (A6–16) to insure sufficient sample size for analysis. Analytical unit U-I sediments were deposited between 13,300 and 10,400 cal yr BP (Hicks, 2004) with a median age of 11,850 cal yr BP (Table 1); although they are stratified, these deposits cannot be consistently subdivided into chronologically finer subsets (Lyman, 2013b). Floodplain sediments are stratified and were deposited between 12,800 and 10,500 cal yr BP (Huckleberry et al., 2004; Huckleyberry and Fadem, 2007; Lyman, 2010) and can be divided into three chronologically distinct analytical units. The Marmes horizon consists of two incipient A pedogenic horizons designated A1 and A2; these sediments were deposited 11,800–10,500 cal yr BP; median age is 11,200 cal yr BP, placing this analytical unit in the early Holocene, but later than the Harrison horizon. The Harrison horizon consists of three incipient A pedogenic horizons designated A3, A4, and A5; these sediments were deposited 12,600–10,500 cal yr BP with a median age of 11,450 cal yr BP, placing it in the earliest Holocene. Finally, a series of eleven incipient A pedogenic horizons were identified stratigraphically beneath the Harrison horizon and designated A6 through A16 (Fryxell and Keel, 1969). Because each of these deeper and more ancient A horizons was minimally sampled, I have lumped them all together for analyses here. A6–16 sediments were deposited 12,800–11,200 cal yr BP; median age is 12,000 cal yr BP, placing it within the latest Pleistocene according to the recently sanctioned boundary between the Pleistocene and Holocene of 11,700 cal yr BP (Walker et al., 2009) (Fig. 2). A portion of this fauna likely accumulated during the end of the cool-moist interval of 13,200 to 12,200 cal yr BP given two calibrated radiocarbon ages in excess of 12,200 cal yr BP (Table 1). Some calibrated ages for an analytical unit do not overlap at the 2-sigma level (Table 1). This is particularly apparent in the A6–16 analytical unit among the three floodplain units, but it is also to be expected given that eleven incipient A horizons are represented in that analytical

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unit compared to two in the Marmes analytical unit and three in the Harrison analytical unit. Lack of overlap of some calibrated ages in analytical unit U-I is also apparent, but it too consists of multiple individual strata that when combined are as much as 2.5 m thick. Clearly, analytical unit U-I is much more coarse-grained than the any of the individual floodplain analytical units (Fig. 2). Paleozoology Previous reports described small samples of the mammal remains recovered from both the rockshelter and the floodplain (Gustafson, 1972; Caulk, 1988; Gustafson and Wegener, 2004). I recently identified the entire Marmes PHT collection from both the PHT-era sediments in the rockshelter and all of the floodplain and corrected several previously published identifications of mammal remains from the site (Lyman, 2011, 2012c, 2013b). I use the number of identified specimens (NISP) to estimate taxonomic abundances. Because NISP values tend to at best provide ordinal-scale estimates of taxonomic abundances (Grayson, 1984; Lyman, 2008), I use ordinal-scale statistics to evaluate various properties of the data. Taxonomic richness (Ntaxa) is a simple tally of the number of taxa, in this case genera or species or some combination thereof tallied in such a way that there is no chance that a taxon is counted twice. For example, if some remains are identified as the genus Neotoma sp., and other remains are identified as the species Neotoma cinerea, only one taxon was tallied for purposes of estimating taxonomic richness. Taxonomic evenness is measured two ways: as the reciprocal of Simpson's index of dominance, or 1/D, and as (1/D)/S, where S = Ntaxa. The lower the value of either index, the more an assemblage is dominated by a single taxon. In other words, as the value of these indices decrease, assemblage evenness decreases (Magurran, 1988, 2004). To quantify change from one temporally and stratigraphically bounded fauna to another, I use the Bray–Curtis dissimilarity index, sometimes called the Sorenson quantitative index (Magurran, 1988). It measures how dissimilar two faunas are by taking into account not only taxa shared by both faunas and taxa unique to each, but also taxonomic abundances. It is calculated with the formula   BCij ¼ 2Cij =Si þ S j

where Cij is the sum of the lower abundances for only those species held in common between both faunas, Si is the abundance of specimens in fauna i and Sj is the abundance of specimens in fauna j. Values of the Bray–Curtis index range from 0, indicating the faunas share no taxa, to 1, indicating the faunas are of precisely the same composition in terms of both taxa represented and taxonomic abundances (Magurran, 2004:174). Samples of identified faunal remains from the Marmes site strata vary considerably in size (Table 2). It is well known that as sample size increases, measures of central tendencies and other properties of samples change until the sample is “representative” of the population. In ecology, this property is termed the “species–area relationship” because it was initially recognized when it was noted that as more geographic area was sampled, more unique species were found until a representative sample of a community had been examined (discussion and references in Lyman and Ames, 2007). This relationship was used by paleontologists when they developed the rarefaction technique, a probabilistic technique to reduce (rarify) the size of large samples to some smaller size to allow comparison between samples of otherwise disparate sizes without fear of sample-size effects influencing results (Sanders, 1968; Raup, 1975; Simberloff, 1978; Tipper, 1979). Rarefaction should be applied to vertebrate faunas with caution because the technique assumes each specimen (bone or tooth or fragment thereof) is independent of every other specimen; such is seldom the case with

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paleozoological remains the taxonomic abundances of which are tallied as NISP (Grayson, 1984). Mammalian taxonomic abundances measured with NISP are rarified here to account for differences in sample size across the strata, but raw NISP (non-rarified) abundances are also examined. Ntaxa, 1/D, (1/D)/S, and Bray–Curtis indices were calculated both for the complete samples of all four assemblages, and for rarified samples. Each assemblage larger than the smallest of the four (A6–16, NISP = 122) was rarified 10,000 times and the mean NISP, standard deviation (SD), and 95% confidence interval (CI) determined for each taxon in the set of rarefied samples. Rarefaction curves were generated for each fauna using PAST (Hammer et al., 2001). When a genus was represented by N 1 species, all remains of a genus were lumped for purposes of rarefaction. Thus, for example, Neotoma sp. remains were lumped with those identified as N. cinerea. Mean richness, mean 1/D, mean (1/D)/S, and mean Bray–Curtis indices were determined from the rarified samples for each assemblage (Table 3). Taxa were tallied as present for tabulation of richness and calculation of 1/D, (1/D)/S, and Bray–Curtis indices only if their mean rarified NISP ≥ 0.5. Chiropterans and specimens identified only to the subfamily Microtinae were not included in analyses. Temporal change in diversity metrics (richness, evenness) was evaluated by examination of overlap in the 95% CIs, and examination of overlap between the means ± 1SD for each stratigraphically bounded assemblage.

Table 2 Taxonomic abundance data (NISP) per stratum for mammals at the Marmes site (45FR50). Taxon (common name) Sorex sp. (long-tailed shrew) Scapanus cf. orarius (coast mole) Antrozous pallidusa (pallid bat) Sylvilagus cf. nuttallii (Nuttall's cottontail) Lepus sp. (jackrabbit) Marmota cf. flaviventris (yellow-bellied marmot) Spermophilus columbianus (Columbian ground squirrel) Spermophilus washingtoni (Washington ground squirrel) Thomomys sp. (smooth-toothed pocket gopher) Thomomys talpoides (northern pocket gopher) Perognathus parvus (Great Basin pocket mouse) Castor canadensis (beaver) Reithrodontomys megalotis (western harvest mouse) Peromyscus maniculatus (deer mouse) Neotoma sp. (wood rat) Neotoma cinerea (bushy-tailed woodrat) Microtinaea Microtus sp. (meadow vole) Lemmiscus curtatus (sage vole) Ondatra zibethicus (muskrat) Vulpes vulpes (red fox) Martes sp. (marten/fisher) Martes americana nobilis (noble marten)b Mustela frenata (long-tailed weasel) Mustela vison (mink) Taxidea taxus (badger) Lutra canadensis (river otter) Lynx sp. (lynx/bobcat) Canis spp. (coyote/dog/wolf) Antilocapra americana (pronghorn) Odocoileus sp. (deer) Ovis canadensis (bighorn sheep) Cervus elaphus (elk/wapiti) Bison sp. (bison) ∑= Richness (Ntaxa) = Evenness (1/D) = Evenness [(1/D)/S] = a b

Not included in tallies. Extinct form.

U-I

A6–16 Harrison Marmes 1 1

3 3 12 99

2 3 4

4 29 42

5 8

1

5

5

8

5

35 9 187 1 1

25 2 17

255 24 114 1

359 48 158

213 90 35 447 126 10 6 10 1 1 1 1 5

12 15 12 84 21 1

36 28 12 122 41 3 1 43 1 1 8 4

65 8 5 202 64 6

10 11 33 2 11 2 921 23 6.807 0.296

3

2

1

3

1 1 1 1 4

2 1 1 1

122 13 8.026 0.617

668 20 4.474 0.224

739 15 2.748 0.183

Results Richness is not correlated with NISP or sample size in the observed assemblages (Table 2). The late Pleistocene assemblage (A6–16) is the smallest and therefore not surprisingly is less taxonomically rich than the two later floodplain assemblages. The earliest Holocene (Harrison horizon) mammal community is taxonomically richer than the later early Holocene (Marmes horizon) community (Table 2). The drop in Ntaxa from the Harrison to the Marmes horizon is particularly evident in the rarified samples (Table 3). The drop in richness from the A6–16 stratum to the Harrison horizon in the rarefied samples is very small (half a taxon) relative to that evident from the Harrison horizon to the Marmes horizon (4.06 taxa). The 95% CI for Ntaxa in the rarified Harrison horizon fauna does not overlap with the observed richness of the A6–16 horizon fauna, but the mean Ntaxa + 1SD for the latter does overlap with the A6–16 horizon fauna. This suggests there is minimal change in richness from A6–16 time to Harrison horizon time. The 95% CIs and the mean ± 1SDs for the Harrison horizon and for the Marmes horizon do not overlap, indicating a significant drop in richness from one to the other about 11,400 cal yr BP. On one hand, rarefaction curves for each assemblage indicate that the two oldest assemblages (A6–16, Harrison) are effectively identical in terms of the influence of sample size on richness (Fig. 3). The 95% confidence limits for the youngest fauna, the Marmes horizon assemblage, on the other hand, quickly diverge from those limits for the two older floodplain assemblages, suggesting there was a major decrease in the richness of the local mammalian fauna about 11,400 cal yr BP. Palynological data indicates a decrease in plant biomass about this time; less primary productivity typically corresponds with a drop in richness of herbivore taxa (Huston and Wolverton, 2011). Such also aligns chronologically with a clinal diminution in body size of some taxa (e.g., Lyman, 2010) and a decrease in the array of body sizes of the members of the mammalian community (Lyman, 2013b). Evenness, whether measured as 1/D, or (1/D)/S, drops consistently over the roughly 800–1000 cal yr represented by the three floodplain assemblages whether the observed or rarified assemblages are considered (Tables 2 and 3). Considering in particular the rarified assemblages to avoid the influences of sample size, the reciprocal of Simpson's index of dominance (1/D) drops from the A6–16 stratum to the Harrison horizon, and also from the Harrison horizon to the Marmes horizon (Table 3). Neither the 95% CI or the mean ± 1 SD for the (1/D) evenness metric overlaps for either temporally sequent pair of assemblages, indicating significant changes in evenness at both stratigraphic boundaries. The other evenness index ([1/D]/S) drops from A6–16 to the Harrison, but not from the Harrison to the Marmes horizon. The 95% CIs suggest both changes are significant, but the mean ± 1 SD for the [(1/D)/S] evenness metric indicates only the A6–16 to Harrison horizon change is significant. In short, the Marmes site floodplain faunas suggest that the drop in evenness began about 11,700 cal yr BP, a bit (~300 cal yr) before a major drop in richness. This makes sense from the perspective that each taxon that exists in an area has a minimum viable population; environmental stress would deplete that population over time until the population was locally extirpated. Thus, in ecological time (year to year), evenness should decrease prior to loss of richness. At the Marmes site, we seem to have the chronological resolution that allows us to perceive such a change in paleozoological time (century to century). The rarified assemblages indicate changes in the abundances of only a few taxa seem to drive the shift in evenness values. First, the abundance of pocket gophers (Thomomys sp.) increases markedly with the deposition of the Harrison horizon (Table 3). The abundance of Great Basin pocket mice (Perognathus parvus) increases a bit, but not nearly as much as that of pocket gophers. Second, the abundances of marmots (Marmota sp.), pack rats (Neotoma sp.), and meadow voles (Microtus sp.) all decrease. Given the habitat preferences of these three genera— they are grazers, particularly marmots and voles—it is likely that the paleoecologically documented decreases in effective moisture and

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301

Table 3 Rarified taxonomic abundance data (NISP) per stratum for mammals at the Marmes site. Values are mean of 10,000 iterations ± SD (95% CI). Taxon

U-I

Sorex sp. Scapanus cf. orarius Sylvilagus cf. nuttallii Lepus sp. Marmota cf. flaviventris Spermophilus spp. Thomomys sp. Perognathus parvus Castor canadensis Reithrodontomys megalotis Peromyscus maniculatus Neotoma spp. Microtus sp. Lemmiscus curtatus Ondatra zibethicus Vulpes vulpes Martes sp. Mustela sp. Taxidea taxus Lutra canadensis Lynx sp. Canis spp. Antilocapra americana Odocoileus sp. Ovis canadensis Cervus elaphus Bison sp. ∑= Ntaxa (NISP ≥ 0.5) = Evenness (1/D) = Evenness [(1/D)/S] =

A6–16

Harrison

Marmes

1 0.2 ± 0.4 0.4 ± 0.6 1.6 ± 1.2 13.1 ± 3.2 0.8 ± 0.8 5.8 ± 2.2 24.8 ± 4.1 0.1 ± 0.3 0.1 ± 0.3 28.2 ± 4.4 16.6 ± 3.5 16.7 ± 3.6 1.3 ± 1.1 0.8 ± 0.8 1.3 ± 1.1 0.3 ± 0.5 0.3 ± 0.5 0.7 ± 0.8

1.3 ± 1.1 1.5 ± 1.1 4.4 ± 1.9 0.3 ± 0.5 1.5 ± 1.1 0.3 ± 0.5 122 14.96 ± 1.67 (.03) 6.54 ± .54 (.01) 0.44 ± .05 (.00)

increased temperature depleted their numbers because of decreased primary productivity. In particular, it seems a decrease in the absolute abundance of grass likely drove the decrease in mammalian evenness. The increased abundance of pocket gophers at Marmes is a bit more difficult to understand. Blois et al. (2010) document a similar local loss of mammalian diversity without regional extirpation or extinction during the PHT in northern California. They document an increase in deer mice (Peromyscus sp.), a taxon with broad climatic tolerances and characterized as the most generalist of North American small mammals given its continent-wide range. The three genera of pocket gophers (Geomyidae) today found in North America also are rather generalist

2 3 4 27 17

12 27 21 1 3

0.7 ± 0.8 5.3 ± 2.0 7.6 ± 2.4 2.4 ± 1.4 50.9 ± 4.9 20.8 ± 3.8 0.2 ± 0.4 6.6 7.3 7.5 0.5 0.2 7.8 0.4 2.2

± ± ± ± ± ± ± ±

2.2 2.4 2.4 0.7 0.4 2.5 0.5 1.3

0.2 0.2 0.2 0.2 0.7

± ± ± ± ±

0.4 0.4 0.4 0.4 0.8

0.8 ± 0.8 1.3 ± 1.0 0.8 ± 0.8 67.2 ± 5.0 26.1 ± 4.1

10.8 ± 2.8 2.1 ± 1.3 10.6 ± 2.9 1.0 ± 0.9

0.3 ± 0.5

1

3

0.3 ± 0.5 0.2 ± 0.4

0.2 ± 0.4 122 13 8.026 0.617

122 12.82 ± 1.39 (.03) 4.41 ± .53 (.01) 0.35 ± .05 (.00)

122 8.76 ± 1.38 (.03) 2.73 ± .26 (.01) 0.32 ± .05 (.00)

in their adaptations, being limited in distribution only by sediment conditions. All gopher taxa prefer loose, friable, well-drained sediments with pore space for gas diffusion and thermoregulation (Chase et al., 1982; but see Marcy et al., 2013). Only one species of pocket gopher, the northern pocket gopher (Thomomys talpoides), occurs in eastern Washington today, and it is the only form that has been identified in late Quaternary sediments there (e.g., Rensberger and Barnosky, 1993; Lyman, 2013b). What might have influenced the increase in relative abundance of this species? The floodplain sediments in front of Marmes Rockshelter that comprise the stratigraphic units A6–16, Harrison horizon, and Marmes

Figure. 3. Rarefaction curves for taxonomic richness of each of four mammalian assemblages at the Marmes site. Dashed lines represent 95% confidence intervals.

302

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fauna (index = 0.232); the index is higher between the U-I fauna and the younger Harrison fauna (index = 0.453) and the Marmes horizon fauna (index = 0.451). These indices suggest that the U-I fauna is more similar to the earliest Holocene mammalian communities (Marmes and Harrison) than to the latest Pleistocene mammalian community (A6–16). Discussion

Figure. 4. Bray–Curtis index values between stratigraphically delimited mammalian communities at the Marmes site. Index values in parentheses are based on observed NISP (Table 2); index values not in parentheses are based on rarefied NISP (Table 3). U-I community is compared to the three summed floodplain communities [0.462, (0.544)], and to each individual floodplain community.

horizon largely represent fluvially deposited fine silts. The incipient A pedogenic horizons making up those stratigraphic units represent brief periods of stasis during which vegetation became established and pedogenic processes began to form soils from the overbank sediments (Huckleberry et al., 2004). Regular flooding likely would have discouraged colonization of the floodplain by pocket gophers and also depressed resident populations. The Palouse River went through an episode of downcutting about 9300 cal yr BP and subsequently ceased washing over the floodplain; 2–4 m of aeolian deposits overlay the Marmes horizon when excavations took place in the middle of the twentieth century (Huckleberry et al., 2004). Chronological resolution does not allow detection of the rate of overbank flooding below Marmes Rockshelter, but it seems plausible to conjecture that decrease in the mean annual discharge of the Palouse River began simultaneously with, or prior to, the formation of the incipient A horizons making up stratum A6–16 (there are reportedly incipient A horizons below A16, but these were only detected in soil probes and not sampled during archaeological excavations). As the floodplain accreted, and gradually became drier for longer periods of time, pocket gophers would have likely become more abundant. For the present this suggestion must remain an hypothesis that could be tested with multiple radiocarbon dates on individual A horizons and larger samples of faunal remains from each of the A horizons. Testing is precluded, however, because these A horizons are presently inundated (and have been since 1969) by an artificial reservoir. Temporal change from one stratigraphically delimited floodplain fauna to the other was fairly consistent but differed a bit in magnitude (Fig. 4). Whether the observed or rarified assemblages are considered, the temporal order of the Bray–Curtis index values are b1.0 and so indicate change from the oldest assemblage (A6–16) to the subsequent Harrison horizon. Less change occurred between the Harrison horizon and the Marmes horizon; the Bray–Curtis index values (observed and rarified) are larger than those between the A6–16 and Harrison communities, indicating greater similarity between the chronologically later pair of communities than between the chronologically earlier pair. The difference in magnitude of the two Bray–Curtis index values suggests greater change earlier (ca. 11,700 cal yr BP) than later (ca. 11,400 cal yr BP). This implies a greater degree of paleoecological change around 11,700 cal yr BP than around 11,400 cal yr BP. This aligns relatively well with local palynological data (Johnson et al., 1994; Mehringer, 1996; Weigand and Hicks, 2004) that indicate rapid major change earlier and slower minor change later. Changes in primary productivity of similar relative magnitude (major early, minor later) were likely the proximate force driving change in the mammalian community. The U-I fauna is essentially contemporaneous with the floodplain horizon faunas. Thus it would not be surprising that there would be some taxonomic similarities between the former and the latter. As noted earlier, the Bray–Curtis index between the observed U-I fauna and the summed observed floodplain faunas is 0.544 (Fig. 4). That index is lowest between the U-I fauna and the oldest A6–16 floodplain

With the exception of the noble marten (M. americana nobilis), none of the taxa identified in the Marmes site assemblages have become extinct. During the PHT at the Marmes site locale several taxa were locally extirpated; rather than become extinct these taxa shifted their ranges at the end of the Pleistocene to higher latitudes or altitudes. For example, the Columbia ground squirrel (Spermophilus columbianus) today occurs in mesic grasslands and coniferous forests at higher elevations than Marmes. The same argument might be made for red fox (Vulpes vulpes) though this canid does sometimes occur in relatively xeric settings; perhaps coyote (Canis latrans) out-competed and thus displaced them as environments warmed in this locale (Larivière and Pasitschniak-Arts, 1996; Lyman, 2012c). Whatever the case, red fox are not historically reported in the area (Johnson and Cassidy, 1997). The beaver (Castor canadensis) and muskrat (Ondatra zibethicus) require water; it seems the Palouse River reduced its flow at the end of the Pleistocene (Marshall, 1971; Huckleberry et al., 2004) and (one suspects) concomitantly riverine vegetation changed so as to be less accommodating to these two species (Wigand and Hicks, 2004). Evenness of the local mammalian community around the Marmes site dropped about 11,700 cal BP and again a few hundred years later. The later drop in evenness was accompanied by a drop in taxonomic richness, much as we might predict with respect to these two properties (evenness and richness) of a biotic community. Correspondence with the rate and magnitude of change in primary productivity, particularly the abundance of grass, suggests decreased plant biomass was the catalyst for change in the mammal community. Decrease in richness and evenness in the Marmes site mammal faunas matches that documented in nearby areas in western North America (e.g., Grayson, 2000; Blois et al., 2010; Schmitt and Lupo, 2012), suggesting that the paleoecological signal is sufficiently robust that it has not been completely obscured by the taphonomic histories of the assemblages. There was no apparent regional loss of taxa in the southern Columbia Plateau where the Marmes site is located, similar to what has been documented elsewhere (e.g., Blois et al., 2010), but there were biogeographic shifts like those documented elsewhere (e.g., Hockett, 2000). There is no evidence of mammalian turnover (taxonomic replacement, in particular) at the Marmes site such as has been documented at other sites in the west (e.g., Grayson, 2000; Blois et al., 2010; Schmitt and Lupo, 2012). Conclusions Local evidence for late Pleistocene anthropogenic impact to (now) extinct megafauna (body mass N 44 kg) is minimal (see Huckleberry et al., 2003 for mention of an as yet unpublished possible local direct association of Pleistocene megafauna and humans). The terminal Pleistocene extinction of numerous taxa of large-bodied (N44 kg) mammal is well known among both paleontologists and neozoologists (Koch and Barnosky, 2006). What is perhaps less well known is that fewer smallbodied mammalian taxa than large-bodied taxa were also lost at this time (Carrasco et al., 2009; Carrasco, 2013). Pertinent data are few in the northwestern U.S., but it seems that the Pleistocene megafauna were locally extirpated prior to the accumulation and deposition of the earliest sediments at the Marmes site (Lyman, 2013a). Terminal Pleistocene decrease in both taxonomic richness and evenness documented at the Marmes site is not a function of the loss of Pleistocene megafauna. Perhaps what we are witnessing is a sort of lag time between loss of large-bodied taxa and later loss of small-bodied

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mammalian taxa as a cascade effect (Owen-Smith, 1987). Consideration of PHT-era changes in mammalian communities documented elsewhere in western North America indicates general patterns but also nuanced variability in those changes. The Marmes site mammalian faunas add to this growing body of knowledge. We still do not fully comprehend the variability in responses of mammalian communities to global warming, whether that which took place during the PHT or that currently taking place. There is minimal evidence that the Earth's ecosystems are experiencing a sixth mass extinction in the same sense as the paleontologically documented big five events (Barnosky et al., 2011). Nevertheless, it is clear that we are losing species at an alarming rate, likely for myriad and still dimly perceived reasons. Limited research suggests that at least some of the historically documented extinctions represent the end of a process of population losses that began at the end of the Pleistocene (e.g., Grayson, 2005). Although this does not soften the reality of species losses over the past couple centuries, it does imply that anthropogenic causes are not the sole reason for those losses. Detection of causes for population loss such as decrease of grazers concomitant with warming and drying, or population gains such as with northern pocket gophers at the Marmes site, should help us better understand and predict the influences of the anthropocene and global warming. It remains to be seen if we pay heed to these lessons learned from the remote past. Acknowledgments Study of the Marmes faunal remains is courtesy of the U.S. Army Corps of Engineers, Walla Walla District and Washington State University, Museum of Anthropology. Research on the Marmes Site collection was funded by NSF grant BCS-0912851. Mary Collins and Diane Curewitz of the Washington State University Museum of Anthropology facilitated my access to the Marmes Site faunal collection. H. M. Gibb, E. M. McCarthy, D. Pierce, C. N. Rosania, and A. K. Trusler provided assistance in the lab. G. Blomquist assisted with the rarefaction analysis. Early versions of this manuscript received extremely valuable comments from Jessica Blois, J. Tyler Faith, and Gary Huckleberry. References Ames, K.M., 1988. Early Holocene forager mobility strategies on the southern Columbia Plateau. In: Willig, J.A., Aikens, C.M., Fagan, J.L. (Eds.), Early Human Occupation in Far Western North America: The Clovis–Archaic Interface. Nevada State Museum Anthropological Papers No. 21, Carson City, NV, pp. 325–360. Barnosky, A.D., 2009. Heatstroke: Nature in an Age of Global Warming. Island Press, Washington, DC. Barnosky, A.D., Hadly, E.A., Bell, C.J., 2003. Mammalian response to global warming on varied temporal scales. Journal of Mammalogy 84, 354–368. Barnosky, A.D., Matzke, N., Tomiya, S., Wogan, G.O.U., Quental, T.B., Marshall, C., McGuire, J.L., Lindsey, E.L., Maguire, K.C., Mersey, B., Ferrer, E.A., 2011. Has the Earth's sixth mass extinction already arrived? Nature 471, 51–57. Blois, J.L., McGuire, J.L., Hadly, E.A., 2010. Small mammal diversity loss in response to latePleistocene climatic change. Nature 465, 771–774. Botkin, D.B., Saxe, H., Araújo, M.B., Betts, R., Bradshaw, R.H.W., Cedhagen, T., Chesson, P., Dawson, T.P., Etterson, J.R., Faith, D.P., Ferrier, S., Guisan, A., Hansen, A.S., Hilbert, D.W., Loehle, C., Margules, C., New, M., Sobel, M.J., Stockwell, D.R.B., 2007. Forecasting the effects of global warming on biodiversity. Bioscience 57, 227–236. Brace, S., Palkopoulou, E., Dalén, L., Lister, A.M., Miller, R., Otte, M., Germonpré, M., Blockley, S.P.E., Stewart, J.R., Barnes, I., 2012. Serial population extinctions in a small mammal indicate Late Pleistocene ecosystem instability. Proceedings of the National Academy of Sciences of the United States of America 109, 20532–20536. Carrasco, M.A., 2013. The impact of taxonomic bias when comparing past and present species diversity. Palaeogeography, Palaeoclimatology, Palaeoecology 372, 130–137. Carrasco, M.A., Barnosky, A.D., Graham, R.A., 2009. Quantifying the extent of North American mammal extinction relative to the pre-anthropogenic baseline. PLoS One 4, e8331. Caulk, G.H., 1988. Examination of Some Faunal Remains from the Marmes Rockshelter Floodplain. Unpublished Master of Arts thesis. Department of Anthropology, Washington State University, Pullman. Chase, J.D., Howard, W.E., Roseberry, J.T., 1982. Pocket gophers (Geomyidae). In: Chapman, J.A., Feldhamer, G.A. (Eds.), Wild Mammals of North America: Biology, Management, and Economics. Johns Hopkins University Press, Baltimore, MD, pp. 239–256. Chatters, R.M., 1968. Washington State University natural radiocarbon measurements I. Radiocarbon 10, 479–498.

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