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Late-Quaternary diminution and abundance of prehistoric bison (Bison sp.) in eastern Washington state, U.S.A.
Quaternary Research 62 (2004) 76 – 85 www.elsevier.com/locate/yqres
Late-Quaternary diminution and abundance of prehistoric bison (Bison sp.) in eastern Washington state, USA R. Lee Lyman * Department of Anthropology, 107 Swallow Hall, University of Missouri, Columbia, MO 65211, USA Received 17 November 2003 Available online 19 May 2004
Abstract Bison (Bison spp.) occurred in eastern Washington state during the late Quaternary. This area is considered to be peripheral to the center of this taxon’s natural range. Bison in the plains east of the Rocky Mountains, the heart of this range, underwent diminution during the late Quaternary, and apparently also did so in other, peripheral areas. A ratio diagram of measurement data derived from eight zooarchaeological collections of bison remains recovered from eastern Washington, in combination with the presence of both sexes and all age classes of individuals, indicate that local bison may have also undergone diminution there. There are, however, a relative paucity of bison remains during the middle Holocene and an apparent 2000-year absence of bison from eastern Washington at this time. As a result, the hypothesis that bison became smaller elsewhere and then immigrated to eastern Washington cannot be falsified. Both the diminution and the fluctuating abundance of bison appear to be responses to forage quality and quantity. D 2004 University of Washington. All rights reserved. Keywords: Bison sp.; Diminution; Eastern Washington state; Forage quality and quantity; North American bison; Ratio diagram
Introduction Perhaps because they are the largest extant indigenous ungulate, or because they foster a romantic image of North America, or for any of several other reasons, bison (Bison bison) are one of the most studied mammals on the continent (Shaw and Meagher, 2000). Their remains reflect aspects of paleoecology and past human behaviors (Hofman and Todd, 2001), so it is fortunate that bison remains are found across much of North America (Graham and Lundelius, 1994). Bison bones and teeth are densest in the fossil record of the Great Plains of central North America, where prehistoric humans often killed and butchered anywhere from a dozen to over one hundred animals at a time. Sites created by these hunting episodes have produced most of the extant data on the prehistory of this large grazer. Less is known about prehistoric bison occupying areas where this taxon was less abundant and hunting apparently involved taking one or two individuals at a time. The periphery of a taxon’s range is often the area where organisms are most sensitive to climatic change, competition, and the like (Brown and Lomolino, * Fax: (573)884-5450. E-mail address: [email protected].
1998). A taxon is much more likely to display responses to environmental flux, of whatever kind, that are detectable in the fossil record at the periphery of its range than in the center of that range where the response signature may be muted by high abundances of the taxon. With respect to bison, one little-studied peripheral area is eastern Washington state. Archaeologist Douglas Osborne (1953) was the first to synthesize zooarchaeological data on bison from eastern Washington and on the basis of those data to argue that modern bison (B. bison) had existed there prehistorically. Paleontological occurrences of bison remains had been known for the previous fifty years (Hay, 1927; Pope, 1952; Sternberg, 1903), but Osborne (1953, p. 261) had the first convincing evidence that bison were present in eastern Washington coincident with early humans and he was explicitly addressing those who thought bison had not been present there ‘‘in historic times’’ or during the Holocene. He listed 1 paleontological (Tipper et al., 1951) and 14 archaeological sites that at the time were known to have produced remains of bison. Schroedl’s (1973) subsequent synthesis of the zooarchaeological occurrence of bison in eastern Washington was prompted by the excavation of what at the time was the only suspected bison-kill site in the area—site
0033-5894/$ - see front matter D 2004 University of Washington. All rights reserved. doi:10.1016/j.yqres.2004.04.001
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45GA17. There are now two other purported bison-kill sites known in eastern Washington (Chatters et al., 1995; Morgan, 1993). Virtually all other known zooarchaeological remains of bison have been recovered from habitation sites. Today, more than 50 archaeological sites and several paleontological locations in eastern Washington have produced remains of bison (Fig. 1), though only a few have produced remains of more than one or two individuals. The fact that bison occurred naturally in eastern Washington during the late Quaternary is now well established. Ancient bison remains in eastern Oregon (Van Vuren and Bray, 1985) seem to confirm a suggested source of prehistoric bison in eastern Washington—they came from western Montana and Wyoming via southern Idaho and eastern Oregon. Debates as to why bison were not found in eastern Washington during the 19th and early 20th century now occupy (paleo)biologists (Laliberte and Ripple, 2003; Lyman and Wolverton, 2002; Martin and Szuter, 1999a, 1999b; Van Vuren, 1987; and references therein). What has not been discussed by zooarchaeologists or paleontologists that might shed light on these debates is the morphometry of
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the bison remains recovered from eastern Washington. That is the focus of this paper. It is well known that North American bison (Bison spp.) became smaller during the late Quaternary (Guthrie, 1970, 1990; McDonald, 1981; Wilson, 1974a, 1974b, 1978, 1980; Wyckoff and Dalquest, 1997). Originally based on decreases in the span and size of horn cores (Wilson, 1978, 1980), the chronocline of diminution of bison across the central North American grasslands of the Great Plains east of the Rocky Mountains is evident in postcranial skeletal elements as well (Bedord, 1974; Hofman and Todd, 2001; Hughes, 1978). Available data indicate that lateQuaternary diminution of bison also took place in areas peripheral to this primary range such as California (Miller and Brotherson, 1979) and Idaho (Butler et al., 1971). Few measurement data have been published for late-Quaternary bison remains from eastern Washington, an area most biologists characterize as a secondary or peripheral bison range (McDonald, 1981; Meagher, 1986; Reynolds et al., 1982; Shaw and Meagher, 2000). Tipper et al. (1951), Pope (1952), and Irwin and Moody (1978) all compare data on
Fig. 1. Map of eastern Washington showing locations of recovery of bison remains. Sites discussed in text are labeled. Dots, archaeological sites; circles, paleontological (including Holocene-age) sites. County lines shown for reference.
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local prehistoric bison remains that they collected with measurements reported by Skinner and Kaisen (1947). Tipper et al. (1951) believed that they had the cranium of a modern-sized Bison bison. Pope (1952) documented what at the time was thought to be a partial cranium of Bison antiquus, a species larger in some dimensions than modern B. bison. Irwin and Moody (1978, p. 241) reported that, based on measurements of one cranium of a mature individual, the 8700-14C-yr-old bison at Lind Coulee (45GR97) probably represented B. bison and that various postcranial bones of the 10 or so individuals represented in the collection ‘‘are comparable in size to modern bison, but a few are slightly larger.’’ They did not, however, provide any measurement data on the postcranial remains recovered from this site. Observations summarized in the preceding paragraphs beg several questions. For one, the taxonomy of late Pleistocene – early Holocene bison in eastern Washington is unclear. Because this question typically is answered with cranial remains (McDonald, 1981; Wilson, 1996), and few of these are known in the area, this question must be ignored for the present. A second question concerns whether the bison (irrespective of the taxa represented) in eastern Washington, like their congeners elsewhere, underwent diminution during the late Quaternary, and if so, why, and if not, why not. In particular, did large bison immigrate to eastern Washington sometime during the late Pleistocene, and then subsequently undergo diminution like populations elsewhere? Or did large bison first colonize eastern Washington, and then as local populations were extirpated, successively smaller bison that evolved elsewhere immigrated, thus giving the appearance that the local population underwent an independent process of diminution? In this paper, I summarize available data that shed light on the local versus extralocal diminution question and outline reasons why the available data appear as they do.
Methods and materials No single geographic location in eastern Washington has provided sufficient bison remains to allow the plotting of measurements against a span of time of several thousand years duration. In fact, every collection of bison remains from a particular site represents a single, very short span of time. Therefore, to compile a series of measurements that span the late Quaternary, it was necessary to measure bison bones collected from various sites of different ages. The few specimens per collection also demanded that any measureable skeletal element be included. Because all that was desired was an indication of whether or not local bison became progressively smaller during the late Quaternary, a diversity of geographic loci and a diversity of skeletal elements were deemed acceptable. Zoorchaeological collections recovered from eastern Washington do not often contain bison remains, and when they do, they tend to be rare relative to those of deer (Odocoileus sp.), elk (Cervus elaphus), bighorn sheep (Ovis canadensis), and pronghorn antelope (Antilocapra americana). Data from 91 zooarchaeological collections distributed throughout eastern Washington were compiled, including the number of identified specimens (NISP) of each ungulate taxon and the age of each collection (Lyman, 2004a). These data were used to determine the abundance of bison relative to other ungulates during particular time periods. Postcranial skeletal remains of bison from eight sites of varying age were measured (Table 1). Standard measurements were taken with dial calipers to the nearest 0.02 mm. Only specimens of adults with fully fused epiphyses were measured to eliminate ontogenetic variation. A ‘‘ratio diagram’’ (Simpson, 1941) is a plot of the difference between the log10 of a measurement of a dimension (e.g., length) of a comparative specimen or mean of a sample of comparative specimens—the comparative standard—and the log10 of the size of that dimension displayed
Table 1 Bison specimens measured and ages from sites in eastern Washington Site
Measured specimens
Age (14C yr BP)
Reference
45KT338 (7) 45AS80 (20) 45AD104 (4) 45WT245 (13) 45GA17 (21) 45GR97 (38) 45KT1362 (7) 45WT134 (2)
First, second, third phalanges Humerus, radius, femur, tibia, first and second phalanges First phalanges Humerus, radius, ulna, first and second phalanges Humerus, radius, tibia Radius, first, second, and third phalanges Second phalanges Humerus
1190F70 (Beta-100992)a 1330F110 (WSU-1440)b 1880F80 (Beta-60297)c 2000 14C yr B.P.d 2230F310 (WSU-465)a 8700 14C yr B.P.e 10,200 14C yr B.P.f 12,280F180 (Beta-29659)c
unpublished Lyman, 1976 Lyman, 1993 unpublished Schroedl (1973) Irwin and Moody (1978) Galm and Gough, 2000 Lyman, 1990
Note. Number in parentheses after site number designates the total number of measurements taken on specimens from the site. a Charcoal. b Burned annual-plant stems. c Bison bone collagen. d Based on temporally diagnostic artifacts. e 8600 F 65 (WSU-1422; humus); 8700 F 400 (C-827; burned bone); 8720 F 200 (WSU-1709; bone collagen). f 10,010 F 60 (Beta-133664; charcoal); 10,130 F 60 (Beta-133665; charcoal); 10,160 F 60 (Beta-133663; charcoal); 10,180 F 40 (Beta-124167; charred material); 10,680 F 190 (Beta-133650; charcoal).
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by a congeneric or conspecific prehistoric specimen. The comparative standard is set at 0.0; a negative value indicates that a prehistoric specimen is smaller than the comparative standard, and a positive value indicates that a prehistoric specimen is larger than the comparative standard. The conversion allows one to simultaneously plot multiple dimensions from multiple specimens. Traditionally, the difference in values is plotted on the x-axis and each dimension holds a unique position on the y-axis. These graphs allow one to detect allometric relationships and general evolutionary patterns. Because all dimensions are normed and plotted against a comparative standard, a ratio diagram can be constructed so as to measure change in size over time. The difference between the dimension(s) of each comparative specimen and the appropriate comparative standard is plotted on the xaxis; the y-axis is used as a chronometer to show the age of each measured specimen irrespective of the dimension measured. The resulting graph is much like a chronocline, except that multiple dimensions of multiple skeletal elements are plotted on the same graph. The dimensions can, for example, include the width of the distal condyle of the humerus, the length of the first phalanx, and the width of the proximal end of the second phalanx. Because the absolute values of the comparative standard vary across the multiple dimensions included, the resulting graph will not be interval scale if the dimensions measured are not randomly distributed across the time span considered. Even then, ordinalscale trends in size, whether stable or changing, will be indicated by the relative positions of clusters of points across time. A key step in the construction of a chronoclinal ratio diagram involves choosing the comparative standards. Many standard measurements of modern comparative bison are available in the literature (e.g., Speth, 1983; Todd, 1983), and there are also many measurements of late-Holocene prehistoric B. bison (e.g., Lorrain, 1968; McDonald, 1981).
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It is unknown if and when the late Quaternary diminution of bison stopped and their average size stabilized. Both lateHolocene data and modern data are used here as comparative standards because they are not significantly different and because the nature of the prehistoric sample demands it in order that samples of prehistoric measurements be large enough to detect general trends. Some sites produced measureable specimens for which only data from modern comparative bison are available (45GA17, 45WT134); data compiled by Todd (1983) on modern bison from Wyoming are used as the comparative standard for these materials. Other sites produced other kinds of measureable specimens for which only late-prehistoric comparative data described by Lorrain (1968) are available (45KT338, 45KT1362, 45AD104); data on bison from Texas dating to about 2650 14 C yr B.P. are used as the comparative standard for these materials. Three sites (45GR97, 45AS80, 45WT245) produced specimens for which both kinds of comparative data are available. The sites in eastern Washington that produced the collections of bison remains discussed here, the skeletal elements measured in each collection, the number of measurements taken per collection, and the age of each collection are listed in Table 1. All ages are given as uncalibrated 14C yr B.P. Only the bison remains from 45WT245 did not have a radiocarbon age directly associated with them; the age of these remains was determined on the basis of associated, temporally diagnostic artifacts.
Results The chronoclinal ratio diagram suggests that bison in eastern Washington, like those elsewhere on the continent, underwent diminution during the late Quaternary (Fig. 2). All of the 47 plotted measurements dating prior to 8500 14C yr B.P. are larger than average modern bison. Of the 65
Fig. 2. Ratio diagram of dimensions of bison bones plotted against a modern standard (0.0 vertical line). Points with positive values are larger than an average modern bison; points with negative values are smaller than an average modern bison.
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plotted measurements dating later than 2300 14C yr B.P., 41 are larger than the comparative standard (an average modern bison) and 24 are smaller. This suggests that late-Holocene bison may have been slightly larger than modern bison. Other late-Holocene collections of bison remains (e.g., Harkins, 1980) are required to reveal any trends in size change during the last several thousand years. Given that members of the genus Bison are notably sexually dimorphic, and that such dimorphism is apparent throughout the late Quaternary (Hofman and Todd, 2001), it is important to consider whether the apparent decrease in size documented in Fig. 2 is a result of a change in sex ratios. If the early collections comprise only or mostly males whereas the latter collections comprise only or mostly females, then the size trend evident in Fig. 2 could merely reflect a shift in sex ratio. This concern cannot be addressed completely because of a lack of skulls and pubic bones in available collections. Postcranial measurements of latePleistocene, early Holocene bison distinguished by sex are
not available. Such measurements are only available for modern bison. Based on comparative data for modern bison (Todd, 1983), four females and two males are represented by the 45GA17 specimens. Specimens from 45AS80 represent one female and two males. Remains of one adult female and one adult male were recovered from 45WT245. In total, there are five female and four male bison represented in late-Holocene collections measured for this study. This suggests that unless only males are represented in the collections >8500 14C yr old, differential representation of the two sexes is not creating the appearance of diminution in Fig. 2. The only other site for which sex ratios of bison are reported is the 2000 14C-yr-old 45BN161, where remains of three males and two females were recovered (Harkins, 1980). This collection does not alter my conclusion regarding sex ratios of resident late-prehistoric bison. Another consideration is whether or not the chronocline in Fig. 2 is a function of how the skeletal elements and dimensions measured are distributed across the time periods.
Fig. 3. Bivariate scatterplot of selected skeletal dimensions. In both, numbers adjacent to plotted points indicate age of the specimen in phalanx; b, distal humerus.
14
C yr. a, second
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Such a distribution does not seem to be exerting any great influence on the point scatters in Fig. 2. For example, a bivariate scatterplot of the lateromedial width of the proximal second phalanx against the lateromedial width of the distal second phalanx indicates that older specimens are, on average, larger than younger specimens (Fig. 3a). This is the only skeletal element with specimens from most represented time periods, but other elements with less complete temporal representation such as the distal humerus produce similar results (Fig. 3b). There is no evidence that temporal variation in the skeletal dimensions measured is causing the size trend evident in Fig. 2. The final and most critical issue concerns whether the bison remains plotted in Fig. 2 actually represent local bison, or immigrants from elsewhere. As noted above, available evidence suggests that bison in eastern Washington originated in western Montana and Wyoming, and that they came into the northwest through southern Idaho (Plew and Sundell, 2000) and eastern Oregon (Van Vuren and Bray, 1985). Some may have originated in Nevada (Van Vuren and Dietz, 1993) or Utah (Lupo, 1996) and moved north, though that source seems less likely given the apparent rarity of bison there relative to their abundance in Wyoming and Montana. Regardless of the source, the question of whether bison established residency and reproduced in eastern Washington is paramount to concluding that local bison underwent diminution there rather than that big bison immigrated first, and successively smaller bison immigrated as time passed. The hypothesis that resident bison became smaller suggests that remains of neonates and fetuses should be present in the eastern Washington record. Further, if the bison were reproducing locally, then individuals of all ages and both sexes should be present. Harkins (1980) reported that one near-term fetus or neonate and seven adults were represented at 45BN161. Remains of a yearling calf, along with those of two skeletally mature adults, were recovered from 45WT245. A single calf and three adults, are represented at 45AS80, and at least one calf and several adults are represented at 45GR97. Available data do not contradict the hypothesis that bison of all ages and both sexes were resident in eastern Washington during the Holocene. Nor do these data contradict the hypothesis that resident bison were reproducing in eastern Washington and underwent diminution there. But they do not contradict the hypothesis that successively smaller individuals immigrated to the area, either. The last is so because bison could have immigrated, established a small local population, reproduced a few generations, and then been extirpated, followed some years later by another immigration, reproduction, extirpation, and so on. Each successive immigrant population could have produced young that grew to be a bit smaller than their predecessors. A way out of this conundrum is to consider the abundance of bison relative to other ungulates and to consider how those abundances correspond (or not) with independent paleoenvironmental indicators and with the size of individual bison.
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This is so because individual bison size and bison abundance seem to be ecologically interrelated and covariant.
Discussion Thirty-seven years ago, Edwards (1967) said that the reason many mammalian taxa underwent diminution during the Holocene was that human predators selectively exploited the largest individuals. This argument was subsequently shown to be inapplicable on the Great Plains, where numerous bison kill sites indicate that it was not unusual for an entire herd to be killed with no regard for age or sex or size of individual bison (Frison, 1991). McDonald (1981, p. 258) puts a different twist on the human predation mechanism. He argues that throughout the late Pleistocene range of bison, human predation increased selection for faster, more agile, more rapidly reproducing organisms, all of which selected for reduced body size. Further, in McDonald’s view, the nutritionally richer, more open (grassland) environment of the terminal Pleistocene and early Holocene reduced intraspecific competition, thereby reducing selection for large individuals. Guthrie (1970, p. 10) suggested that increased nutrients would increase ‘‘the survival of larger individuals . . . allowing them to contribute more to future generations’’ but also that ‘‘increased intrapopulational competition . . . generally results in a heritable decrease in individual size.’’ A decade later he suggested that during the Holocene on the Plains of North America ‘‘bison were becoming more abundant while also decreasing in body size’’ (Guthrie, 1980, p. 56). A terminal Pleistocene decrease in interspecific competition, plus an increase in short-grass prairies favored an increase in bison numbers (Guthrie, 1980, p. 67). Almost simultaneously, there was an ‘‘opening of the winter bottleneck’’ by which Guthrie (1980, p. 68) meant that the reproductive season of bison was lengthier and there was reduced winter mortality during the Holocene relative to that during the Pleistocene. Both factors resulted in greater bison numbers which in turn increased intraspecific competition and this selected for reduced body size (Guthrie, 1980, p. 69). Butler et al. (1971) suggested that bison were subject to Bergmann’s rule, but Wilson (1974a) showed that the former had no data to substantiate their claim. Wilson (1974a) expressed preference for climatic factors being responsible for size variation in bison, probably because he noted that bison remains from Wyoming dating to the Altithermal indicated that this was the period when local bison became smaller. Thus, concluded that ‘‘a causal relationship is probable’’ between climate and bison size (Wilson, 1974b, p. 96). In light of additional data, Wilson (1978, p. 15) later concluded that ‘‘dwarfing, in fact, took place throughout the Holocene’’ and expressed favor for Geist’s (1971) and Guthrie’s (1970) suggestion that greater intraspecific competition for food ‘‘accounted for some of the Holocene dwarfing.’’
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Interestingly, and with some potential relevance for the earliest bison remains in eastern Washington, Geist (1971, p. 287) proposed a ‘‘dispersal theory’’ which held that ‘‘pioneering [colonizing] populations find a superabundance of qualitatively excellent forage in the vacant habitat that causes high birth rates, high birth weights, high milk production, rapid growth of young, early sexual maturation, [and] large body size of adults close to the genetic maximum.’’ If forage on the edge of a taxon’s range is neither abundant nor of high quality, then I suspect that when that taxon colonizes those peripheral areas individuals born there will be few and they will be small. Paleozoological evidence regarding bison abundance in Texas, near the southern edge of their range, indicates that bison only became abundant there about 1000 14C yr B.P. when grass biomass increased as a result of environmental change (Huebner, 1991). It has long been known that bison were rare prehistorically in eastern Washington relative to their abundance in the Plains. The debate over why the relative abundances should be so has a deep history. Some recent commentators, however, make it sound as if they thought up a new cause for the rarity of bison in eastern Washington when they proposed that prehistoric aboriginal exploitation depleted local populations (Kay, 1994; Martin and Szuter, 1999a, 1999b). This idea, however, has been around for at least seventy years. Kingston (1932, p. 164), for example, wrote that eastern Washington ‘‘presents a curious and rather intricate problem of biological distribution in which hunting by the Indians and physiographic difficulties explain for the most part the scarcity or absence of the buffalo.’’ The physiographic difficulties involved the fact that the likely immigration route is better characterized as a ‘‘filter’’ than as a ‘‘corridor’’ (Simpson, 1940, 1953); thus local recruitment was largely limited to reproduction and only occasionally
immigration. Fifty-five years later Van Vuren (1987) added low overall forage production in eastern Washington to the mix. The modern debate concerns whether Kingston and Van Vuren’s proposed combination of factors or only overhunting by human predators was the cause (Laliberte and Ripple, 2003; Lyman and Wolverton, 2002; Martin and Szuter, 2002). The zooarchaeological record provides no unequivocal evidence that local elk (Cervus elaphus) populations were depressed by human predation (Lyman, 2004a), despite argument to the contrary (Kay, 1994). The relative abundance of elk seems to flunctuate in concert with paleoclimatic conditions; elk remains are abundant when it is cool and moist but less abundant when it is warm and dry (Lyman, 2004b; Lyman and Wolverton, 2002). Bison abundances also fluctuate in concert with paleoclimatic conditions, and those conditions help account for the size of individual bison in eastern Washington. Bison remains dating to the mid-Holocene Altithermal climatic interval of warmth and aridity are more rare in eastern Washington than at any other time during the Holocene (Lyman, 1985; Schroedl, 1973). Based on 91 collections of zooarchaeological remains from sites throughout eastern Washington with good temporal control, bison remains are most abundant relative to other ungulates during the terminal Pleistocene, initial Holocene, and late Holocene, and least abundant during the middle Holocene (Fig. 4). Given that early, middle, and late Holocene sediments have been differentially sampled (e.g., Lyman, 1992, 2000), it is important to note that there is no correlation between the proportion of ungulate remains representing bison and either the number of radiocarbon ages per period plotted in Fig. 4 or the number of assemblages of remains per period ( p>0.5 for both). There is no evidence that changes in human subsistence pursuits or technology resulted in decreased pursuit of
Fig. 4. Abundance of bison remains relative to remains of all ungulates over the past 10,500
14
C yr in eastern Washington.
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bison between 8000 and 2500 14C yr B.P. (Lyman, 1992; Schroedl, 1973). Rather, there were few bison to pursue. As part of their response to local environmental change, local bison populations were small or nonexistant between 8000 and 2500 14C yr B.P. Thirty-two sites dating between 6000 and 8000 14C yr B.P. failed to produce remains of bison. This 2000-14C-yr gap in the local record argues against local bison undergoing in situ diminution during the last 12,000 14C yr (Fig. 2). Rather, it is more parsimonious to conclude that the local early Holocene population of large bison was extirpated about 8000 14C yr B.P. and that after about 6000 yr 14C B.P. successively smaller bison immigrated sporadically, but no resident population surivived very long or at any significantly large size until after 2500 14C yr B.P. Although any resident population established by post-6000-14C-yr-B.P. immigrants may have undergone some degree of diminution, data necessary to assess this possibility are currently unavailable. Grass (as a proxy measure of bison nutrition) was most abundant during the terminal Pleistocene, early Holocene, and late Holocene, and rare during the middle Holocene (Chatters, 1998). Bison that immigrated to eastern Washington during the terminal Pleistocene and early Holocene were of large size because they originated in herds of large individuals; they maintained their large size in eastern Washington because forage was abundant and nutritious. During the late Holocene, bison were of small size due in part to their origins in herds of small individuals. Their abundances were low due in part to intraspecific competition. The low quality and quantity of forage during the middle Holocene significantly depressed the local bison population. That depression may have been exacerbated by human predation, though evidence of such is nonexistant. At present, the most parsimonious account of available data is that the local bison population was completely extirpated about 8000 14C yr B.P. Bison sporadically immigrated between 6000 and 2500 14C yr B.P., but were unable to establish a local population because of poor forage conditions and human predation. After 2500 14C yr B.P. they were again locally abundant because climate had returned to the relatively cool – moist conditions of modern times and grass biomass had increased. Between about 4000 and 2500 14C yr B.P., climates were the coolest and most moist of the entire Holocene (Chatters, 1998), raising the question of why bison were not more abundant in eastern Washington at this time. Modern bison consume mostly graminoids, often eating higher proportions of grass than would be predicted on the basis of grass availability on the landscape; they eat very small amounts of forbs and woody species (Knapp et al., 1999, and references therein). Long-term observations indicate that bison graze preferentially in recently burned areas because of higher grass productivity and decreased forb cover (Knapp et al., 1999). The abundance of charcoal in lake sediments is known to directly correlate with fire
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history (Gardner and Whitlock, 2001, and references therein). Charcoal abundances in such sediments in eastern Washington, northern Idaho, and western Montana indicate fires were relatively infrequent between about 4000 and 2000 14C yr B.P. but were more frequent after that time (Chatters, 1998, and references therein). If bison immigrated to eastern Washington from southwestern Montana and western Wyoming, and passed through southern Idaho, then the increased incidence of (anthropogenic?) fires over the past 2000 or so 14C yr may have enhanced forage quality along that route and thus allowed larger numbers of bison to immigrate. Relatively low abundances of bison during the last 500 14C yr may be a result of increased predation brought on by the acquisition of horses about A.D. 1730 by native peoples of eastern Washington (Haines, 1938). Temporal resolution of the collections of faunal remains dating to this late prehistoric period is insufficient to determine if the depression in bison abundance is in fact coincident with the appearance of horses.
Conclusion The late Quaternary record of bison in the Great Plains— the heart of bison range during this period (Graham and Lundelius, 1994)—indicates that this taxon underwent diminution over the past 12,000 to 15,000 14C yr. Bison occurred in areas peripheral to the Plains off and on throughout this period, seeming to fluctuate in abundance more or less in concert with changes in environments, but also perhaps at least in part in response to human predation. The question that arises is whether bison in these peripheral areas also underwent diminution independently of that which took place on the Plains. Thus, the alternative hypothesis of diminution in the heartland followed by immigration to the subject peripheral area must be considered. The prehistoric record of bison in eastern Washington indicates that this taxon was present there over much of the late Quaternary, but it was often quite rare and apparently absent during significant portions of the hot – dry middle Holocene when much diminution took place among bison on the Plains. The absence of evidence for their middleHolocene presence in eastern Washington means that the diminution elsewhere followed by subsequent immigration hypothesis cannot be discounted. Only if it can be shown that bison were consistently present in eastern Washington throughout the past 10,000 or so years can the hypothesis of local diminution be considered possible.
Acknowledgments I thank D. R. Brauner, J. R. Galm, S. Gough, K. Lupo, D. Schmitt, and G. F. Schroedl for the opportunities to study the collections discussed here, and C. R. Harington and especially M. Wilson for insightful comments on an early
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draft. M. Collins (Washington State University Museum of Anthropology) provided access to the Lind Coulee and 45AS80 bison bones, and K. Lupo provided lab space. This research was partially funded by the University of Missouri—Columbia, Chancellor’s Office. This paper is dedicated to the memory of Frank C. Leonhardy, who in 1974 provided my first opportunity to study faunal remains. References Bedord, J.N., 1974. Morphological variation in bison metacarpals and metatarsals. In: Frison, G.C. (Ed.), The Casper Site: A Hell Gap Bison Kill on the High Plains. Academic Press, New York, pp. 199 – 240. Brown, J.H., Lomolino, M.V., 1998. Biogeography, second ed. Sinauer, Sunderland, MA. Butler, B.R., Gildersleeve, H., Sommers, J., 1971. The Wasden Site bison: Sources of morphological variation. In: Stryd, A.H., Smith, R.A. (Eds.), Aboriginal Man and Environments on the Plateau of Western North America. Students’ Press, Univ. of Calgary, pp. 126 – 152. Chatters, J.C., 1998. Environment. In: Walker Jr., D.E. (Ed.), Handbook of North American Indians, Vol. 12: Plateau. Smithsonian Institution, Washington, DC, pp. 29 – 49. Chatters, J.C., Campbell, S.K., Smith, G.D., Minthorn Jr., P.E., 1995. Bison procurement in the far west: A 2,100-year-old kill on the Columbia Plateau. American Antiquity 60, 751 – 763. Edwards, W.E., 1967. The late-Pleistocene extinction and diminution in size of many mammalian species. In: Martin, P.S., Wright Jr., H.E. (Eds.), Pleistocene Extinctions: The Search for a Cause. Yale Univ. Press, New Haven, CT, pp. 141 – 154. Frison, G.C., 1991. Prehistoric Hunters of the High Plains, second ed. Academic Press, New York. Galm, J.R., Gough, S., 2000. Site 45KT1362, a c. 10,000 B.P. occupation in central Washington. Current Research in the Pleistocene 17, 29 – 31. Gardner, J.J., Whitlock, C., 2001. Charcoal accumulation following a recent fire in the Cascade Range, northwestern USA, and its relevance for fire-history studies. The Holocene 11, 541 – 549. Geist, V., 1971. The relation of social evolution and dispersal in ungulates during the Pleistocene, with emphasis on the Old World deer and the genus Bison. Quaternary Research 1, 283 – 315. Graham, R.W., Lundelius Jr., E.L., 1994. Faunmap: A Database Documenting Late Quaternary Distributions of Mammal Species in the United States, Scientific Papers 25, Illinois State Museum, Springfield. Guthrie, R.D., 1970. Bison evolution and zoogeography in North America during the Pleistocene. Quarterly Review of Biology 45, 1 – 15. Guthrie, R.D., 1980. Bison and man in North America. Canadian Journal of Anthropology 1, 55 – 73. Guthrie, R.D., 1990. Frozen Fauna of the Mammoth Steppe: The Story of Blue Babe. Univ. of Chicago Press, Chicago. Haines, F., 1938. The northward spread of horses among the Plains Indians. American Anthropologist 40, 429 – 437. Harkins, S.K., 1980. The Prehistoric Occurrence and Aboriginal Utilization of Bison sp. in the Central Columbia Basin. Unpublished Master of Arts thesis. Department of Anthropology, Washington State Univ., Pullman. Hay, O.P., 1927. The Pleistocene of the Western Region of North America and Its Vertebrated Animals. Publication 322B, Carnegie Institute of Washington. Hofman, J.L., Todd, L.C., 2001. Tyranny in the archaeological record of specialized hunters. In: Gerlach, S.C., Murray, M.S. (Eds.), People and Wildlife in Northern North America: Essays in Honor of R. Dale Guthrie. British Archaeological Reports, International Series vol. 944. pp. 200 – 215. Oxford. Huebner, J.A., 1991. Late prehistoric bison populations in central and southern Texas. Plains Anthropologist 36, 343 – 358.
Hughes, S.S., 1978. Bison diminution on the Great Plains. In: Smiley, F.E., Hofman, J.L. (Eds.), Wyoming Contributions to Anthropology, vol. 1. Univ. of Wyoming Publications vol. 32(1), pp. 18 – 47. (Laramie). Irwin, A.M., Moody, U., 1978. The Lind Coulee Site (45GR97), Washington Archaeological Research Center Project Report No. 56. Pullman. Kay, C.E., 1994. Aboriginal overkill: The role of Native Americans in structuring western ecosystems. Human Nature 5, 359 – 398. Kingston, C.S., 1932. Buffalo in the Pacific Northwest. Washington Historical Quarterly 23, 163 – 172. Knapp, A.K., Blair, J.M., Briggs, J.M., Collins, S.L., Hartnett, D.C., Johnson, L.C., Towne, E.G., 1999. The keystone role of bison in North American tallgrass prairie. Bioscience 49, 39 – 50. Laliberte, A.S., Ripple, W.J., 2003. Wildlife encounters by Lewis and Clark: A spatial analysis of interactions between Native Americans and wildilfe. BioScience 53, 994 – 1003. Lorrain, D., 1968. Analysis of the bison bones. In: Dibble, D.S., Lorrain, D. (Eds.), Bonfire Shelter: A Stratified Bison Kill Site, Val Verde County, Texas, Miscellaneous Papers No. 1. Texas Tech Memorial Museum, Lubbock, TX, pp. 78 – 134. Lupo, K.D., 1996. The historical occurrence and demise of bison in northern Utah. Utah Historical Quarterly 64, 168 – 180. Lyman, R.L. 1976. A Cultural Analysis of Faunal Remains from the Alpowa Locality. Unpublished Master of Arts thesis. Department of Anthropology, Washington State Univ., Pullman. Lyman, R.L., 1985. The paleozoology of the Avey’s Orchard site. In: Galm, J.R., Masten, R.A. (Eds.), Avey’s Orchard: Archaeological Investigations of a Late Prehistoric Columbia River Community. Reports in Archaeology and History 100 – 42. Eastern Washington University, Cheney, pp. 243 – 319. Lyman, R.L., 1990. Zooarchaeology. In: Brauner, D.R. (Ed.), Archaeological Data Recovery at Hatiupuh, 45WT134, Whitman County, Washington. Report to the Walla Walla District U.S. Army Corps of Engineers. Oregon State University, Department of Anthropology, pp. 98 – 138. Lyman, R.L., 1992. Influences of mid-Holocene Altithermal climates on mammalian faunas and human subsistence in eastern Washington. Journal of Ethnobiology 12, 37 – 62. Lyman, R.L., 1993. The faunal collection. In: Morgan, V. (Ed.), Cultural Resource Investigations at 45AD104, An Upland Bison Site, Adams County, Washington. Reports in Archaeology and History 100-81. Eastern Washington University, Cheney, pp. 39 – 56. Lyman, R.L., 2000. Building cultural chronology in eastern Washington: The influence of geochronology, index fossils, and radiocarbon dating. Geoarchaeology 15, 609 – 648. Lyman, R.L., 2004a. Aboriginal overkill in the intermountain west of North America: Zooarchaeological tests and implications. Human Nature 15, 169 – 208. Lyman, R.L., 2004b. Prehistoric biogeography, abundance, and phenotypic plasticity of elk (Cervus elaphus) in Washington state. In: Lyman, R.L., Cannon, K.P. (Eds.), Zooarchaeology and Conservation Biology. Univ. of Utah Press, Salt Lake City. Lyman, R.L., Wolverton, S., 2002. The late prehistoric – early historic game sink in the northwestern United States. Conservation Biology 16, 73 – 85. Martin, P.S., Szuter, C.R., 1999a. Megafauna of the Columbia Basin, 1800 – 1840: Lewis and Clark in a game sink. In: Goble, D.D., Hart, P.W. (Eds.), Northwest lands, Northwest Peoples: Readings in Environmental History. Univ. of Washington Press, Seattle, pp. 188 – 204. Martin, P.S., Szuter, C.R., 1999b. War zones and game sinks in Lewis and Clark’s west. Conservation Biology 13, 36 – 45. Martin, P.S., Szuter, C.R., 2002. Game parks before and after Lewis and Clark: Reply to Lyman and Wolverton. Conservation Biology 16, 244 – 247. McDonald, J.N., 1981. North American Bison: Their Classification and Evolution. Univ. of California Press, Berkeley. Meagher, M., 1986. Bison bison. Mammal Species 266, 1 – 8. Miller, W.E., Brotherson, J.D., 1979. Size variation in foot elements of
R.L. Lyman / Quaternary Research 62 (2004) 76–85 Bison from Rancho La Brea. Natural History Museum of Los Angeles County, Contributions in Science 323, 1 – 19. Morgan, V., 1993. Cultural Resource Investigations at 45AD104, An Upland Bison Site, Adams County, Washington. Reports in Archaeology and History 100-81. Eastern Washington University, Cheney. Osborne, D.H., 1953. Archaeological occurrences of pronghorn antelope, bison, and horse in the Columbia Plateau. The Scientific Monthly 77, 260 – 269. Plew, M.G., Sundell, T., 2000. The archaeological occurrence of bison on the Snake River Plain. North American Archaeologist 21, 119 – 137. Pope, P.H., 1952. A fossil bison skull from Prescott, Washington. Northwest Science 26, 51 – 54. Reynolds, H.W., Glaholt, R.D., Hawley, A.W.L., 1982. Bison, Bison bison. In: Chapman, J.A., Feldhamer, G.A. (Eds.), Wild Mammals of North America: Biology, Management, Economics. Johns Hopkins Univ. Press, Baltimore, pp. 972 – 1007. Schroedl, G.F., 1973. The Archaeological Occurrence of Bison in the Southern Plateau. Reports of Investigations No. 51. Laboratory of Anthropology, Washington State University, Pullman. Shaw, J.H., Meagher, M., 2000. Bison. In: Demarais, S., Krausman, P.R. (Eds.), Ecology and Management of Large Mammals in North America. Prentice – Hall, Upper Saddle River, NJ, pp. 447 – 466. Simpson, G.G., 1940. Mammals and land bridges. Journal of the Washington Academy of Sciences 30, 137 – 163. Simpson, G.G., 1941. Large Pleistocene felines of North America. American Museum Novitates 1136, 1 – 27. Simpson, G.G., 1953. Evolution and Geography: An Essay on Historical Biogeography with Special Reference to Mammals. Univ. of Oregon Press, Eugene. Skinner, M.F., Kaisen, O.C., 1947. The fossil Bison of Alaska and preliminary revision of the genus. American Museum of Natural History Bulletin 89, 123 – 256. Speth, J.D., 1983. Bison Kills and Bone Counts. Univ. of Chicago Press, Chicago.
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Sternberg, C.H., 1903. Elephas columbi and other mammals in swamps of Whitman County, Washington. Science 17, 511 – 512. Tipper, H.W., Jizba, Z.V., Konizeski, R.L., 1951. A bison from eastern Washington. American Journal of Science 249, 318 – 322. Todd Jr., L.G., 1983. The Horner Site: Taphonomy of an Early Holocene Bison Bonebed. Unpublished Ph.D. dissertation. Univ. of New Mexico, Albuquerque. Van Vuren, D., 1987. Bison west of the Rocky Mountains: An alternative explanation. Northwest Science 61, 65 – 69. Van Vuren, D., Bray, M.P., 1985. The recent geographic distribution of Bison bison in Oregon. Murrelet 66, 56 – 58. Van Vuren, D., Dietz, F.C., 1993. Evidence of Bison bison in the Great Basin. Great Basin Naturalist 53, 318 – 319. Wilson, M., 1974a. History of the bison in Wyoming, with particular reference to early Holocene forms. In: Wilson, M. (Ed.), Applied Geology and Archaeology: The Holocene History of Wyoming. Report of Investigations, vol. 10. Geological Survey of Wyoming, Laramie, pp. 91 – 99. Wilson, M., 1974b. The Casper local fauna and its fossil bison. In: Frison, G.C. (Ed.), The Casper Site: A Hell Gap Bison Kill on the High Plains. Academic Press, New York, pp. 125 – 171. Wilson, M., 1978. Archaeological kill site populations and the Holocene evolution of the genus Bison. In: Davis, L.B., Wilson, M. (Eds.), Bison Procurement and Utilization: A Symposium. Plains Anthropologist Memoir, vol. 14, pp. 9 – 22. Wilson, M., 1980. Morphological dating of late Quaternary bison on the northern Plains. Canadian Journal of Anthropology 1, 81 – 85. Wilson, M., 1996. Late Quaternary vertebrates and the opening of the icefree corridor, with special reference to the genus Bison. Quaternary International 32, 97 – 105. Wyckoff, D.G., Dalquest, W.W., 1997. From whence they came: The paleontology of southern plains bison. In: Bement, L.C., Buehler, K.J. (Eds.), Southern Plains Bison Procurement and Utilization from Paleoindian to Historic. Plains Anthropologist Memoir, vol. 42, pp. 5 – 32.
Late-Quaternary diminution and abundance of prehistoric bison (Bison sp.) in eastern Washington state, USA R. Lee Lyman * Department of Anthropology, 107 Swallow Hall, University of Missouri, Columbia, MO 65211, USA Received 17 November 2003 Available online 19 May 2004
Abstract Bison (Bison spp.) occurred in eastern Washington state during the late Quaternary. This area is considered to be peripheral to the center of this taxon’s natural range. Bison in the plains east of the Rocky Mountains, the heart of this range, underwent diminution during the late Quaternary, and apparently also did so in other, peripheral areas. A ratio diagram of measurement data derived from eight zooarchaeological collections of bison remains recovered from eastern Washington, in combination with the presence of both sexes and all age classes of individuals, indicate that local bison may have also undergone diminution there. There are, however, a relative paucity of bison remains during the middle Holocene and an apparent 2000-year absence of bison from eastern Washington at this time. As a result, the hypothesis that bison became smaller elsewhere and then immigrated to eastern Washington cannot be falsified. Both the diminution and the fluctuating abundance of bison appear to be responses to forage quality and quantity. D 2004 University of Washington. All rights reserved. Keywords: Bison sp.; Diminution; Eastern Washington state; Forage quality and quantity; North American bison; Ratio diagram
Introduction Perhaps because they are the largest extant indigenous ungulate, or because they foster a romantic image of North America, or for any of several other reasons, bison (Bison bison) are one of the most studied mammals on the continent (Shaw and Meagher, 2000). Their remains reflect aspects of paleoecology and past human behaviors (Hofman and Todd, 2001), so it is fortunate that bison remains are found across much of North America (Graham and Lundelius, 1994). Bison bones and teeth are densest in the fossil record of the Great Plains of central North America, where prehistoric humans often killed and butchered anywhere from a dozen to over one hundred animals at a time. Sites created by these hunting episodes have produced most of the extant data on the prehistory of this large grazer. Less is known about prehistoric bison occupying areas where this taxon was less abundant and hunting apparently involved taking one or two individuals at a time. The periphery of a taxon’s range is often the area where organisms are most sensitive to climatic change, competition, and the like (Brown and Lomolino, * Fax: (573)884-5450. E-mail address: [email protected].
1998). A taxon is much more likely to display responses to environmental flux, of whatever kind, that are detectable in the fossil record at the periphery of its range than in the center of that range where the response signature may be muted by high abundances of the taxon. With respect to bison, one little-studied peripheral area is eastern Washington state. Archaeologist Douglas Osborne (1953) was the first to synthesize zooarchaeological data on bison from eastern Washington and on the basis of those data to argue that modern bison (B. bison) had existed there prehistorically. Paleontological occurrences of bison remains had been known for the previous fifty years (Hay, 1927; Pope, 1952; Sternberg, 1903), but Osborne (1953, p. 261) had the first convincing evidence that bison were present in eastern Washington coincident with early humans and he was explicitly addressing those who thought bison had not been present there ‘‘in historic times’’ or during the Holocene. He listed 1 paleontological (Tipper et al., 1951) and 14 archaeological sites that at the time were known to have produced remains of bison. Schroedl’s (1973) subsequent synthesis of the zooarchaeological occurrence of bison in eastern Washington was prompted by the excavation of what at the time was the only suspected bison-kill site in the area—site
0033-5894/$ - see front matter D 2004 University of Washington. All rights reserved. doi:10.1016/j.yqres.2004.04.001
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45GA17. There are now two other purported bison-kill sites known in eastern Washington (Chatters et al., 1995; Morgan, 1993). Virtually all other known zooarchaeological remains of bison have been recovered from habitation sites. Today, more than 50 archaeological sites and several paleontological locations in eastern Washington have produced remains of bison (Fig. 1), though only a few have produced remains of more than one or two individuals. The fact that bison occurred naturally in eastern Washington during the late Quaternary is now well established. Ancient bison remains in eastern Oregon (Van Vuren and Bray, 1985) seem to confirm a suggested source of prehistoric bison in eastern Washington—they came from western Montana and Wyoming via southern Idaho and eastern Oregon. Debates as to why bison were not found in eastern Washington during the 19th and early 20th century now occupy (paleo)biologists (Laliberte and Ripple, 2003; Lyman and Wolverton, 2002; Martin and Szuter, 1999a, 1999b; Van Vuren, 1987; and references therein). What has not been discussed by zooarchaeologists or paleontologists that might shed light on these debates is the morphometry of
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the bison remains recovered from eastern Washington. That is the focus of this paper. It is well known that North American bison (Bison spp.) became smaller during the late Quaternary (Guthrie, 1970, 1990; McDonald, 1981; Wilson, 1974a, 1974b, 1978, 1980; Wyckoff and Dalquest, 1997). Originally based on decreases in the span and size of horn cores (Wilson, 1978, 1980), the chronocline of diminution of bison across the central North American grasslands of the Great Plains east of the Rocky Mountains is evident in postcranial skeletal elements as well (Bedord, 1974; Hofman and Todd, 2001; Hughes, 1978). Available data indicate that lateQuaternary diminution of bison also took place in areas peripheral to this primary range such as California (Miller and Brotherson, 1979) and Idaho (Butler et al., 1971). Few measurement data have been published for late-Quaternary bison remains from eastern Washington, an area most biologists characterize as a secondary or peripheral bison range (McDonald, 1981; Meagher, 1986; Reynolds et al., 1982; Shaw and Meagher, 2000). Tipper et al. (1951), Pope (1952), and Irwin and Moody (1978) all compare data on
Fig. 1. Map of eastern Washington showing locations of recovery of bison remains. Sites discussed in text are labeled. Dots, archaeological sites; circles, paleontological (including Holocene-age) sites. County lines shown for reference.
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local prehistoric bison remains that they collected with measurements reported by Skinner and Kaisen (1947). Tipper et al. (1951) believed that they had the cranium of a modern-sized Bison bison. Pope (1952) documented what at the time was thought to be a partial cranium of Bison antiquus, a species larger in some dimensions than modern B. bison. Irwin and Moody (1978, p. 241) reported that, based on measurements of one cranium of a mature individual, the 8700-14C-yr-old bison at Lind Coulee (45GR97) probably represented B. bison and that various postcranial bones of the 10 or so individuals represented in the collection ‘‘are comparable in size to modern bison, but a few are slightly larger.’’ They did not, however, provide any measurement data on the postcranial remains recovered from this site. Observations summarized in the preceding paragraphs beg several questions. For one, the taxonomy of late Pleistocene – early Holocene bison in eastern Washington is unclear. Because this question typically is answered with cranial remains (McDonald, 1981; Wilson, 1996), and few of these are known in the area, this question must be ignored for the present. A second question concerns whether the bison (irrespective of the taxa represented) in eastern Washington, like their congeners elsewhere, underwent diminution during the late Quaternary, and if so, why, and if not, why not. In particular, did large bison immigrate to eastern Washington sometime during the late Pleistocene, and then subsequently undergo diminution like populations elsewhere? Or did large bison first colonize eastern Washington, and then as local populations were extirpated, successively smaller bison that evolved elsewhere immigrated, thus giving the appearance that the local population underwent an independent process of diminution? In this paper, I summarize available data that shed light on the local versus extralocal diminution question and outline reasons why the available data appear as they do.
Methods and materials No single geographic location in eastern Washington has provided sufficient bison remains to allow the plotting of measurements against a span of time of several thousand years duration. In fact, every collection of bison remains from a particular site represents a single, very short span of time. Therefore, to compile a series of measurements that span the late Quaternary, it was necessary to measure bison bones collected from various sites of different ages. The few specimens per collection also demanded that any measureable skeletal element be included. Because all that was desired was an indication of whether or not local bison became progressively smaller during the late Quaternary, a diversity of geographic loci and a diversity of skeletal elements were deemed acceptable. Zoorchaeological collections recovered from eastern Washington do not often contain bison remains, and when they do, they tend to be rare relative to those of deer (Odocoileus sp.), elk (Cervus elaphus), bighorn sheep (Ovis canadensis), and pronghorn antelope (Antilocapra americana). Data from 91 zooarchaeological collections distributed throughout eastern Washington were compiled, including the number of identified specimens (NISP) of each ungulate taxon and the age of each collection (Lyman, 2004a). These data were used to determine the abundance of bison relative to other ungulates during particular time periods. Postcranial skeletal remains of bison from eight sites of varying age were measured (Table 1). Standard measurements were taken with dial calipers to the nearest 0.02 mm. Only specimens of adults with fully fused epiphyses were measured to eliminate ontogenetic variation. A ‘‘ratio diagram’’ (Simpson, 1941) is a plot of the difference between the log10 of a measurement of a dimension (e.g., length) of a comparative specimen or mean of a sample of comparative specimens—the comparative standard—and the log10 of the size of that dimension displayed
Table 1 Bison specimens measured and ages from sites in eastern Washington Site
Measured specimens
Age (14C yr BP)
Reference
45KT338 (7) 45AS80 (20) 45AD104 (4) 45WT245 (13) 45GA17 (21) 45GR97 (38) 45KT1362 (7) 45WT134 (2)
First, second, third phalanges Humerus, radius, femur, tibia, first and second phalanges First phalanges Humerus, radius, ulna, first and second phalanges Humerus, radius, tibia Radius, first, second, and third phalanges Second phalanges Humerus
1190F70 (Beta-100992)a 1330F110 (WSU-1440)b 1880F80 (Beta-60297)c 2000 14C yr B.P.d 2230F310 (WSU-465)a 8700 14C yr B.P.e 10,200 14C yr B.P.f 12,280F180 (Beta-29659)c
unpublished Lyman, 1976 Lyman, 1993 unpublished Schroedl (1973) Irwin and Moody (1978) Galm and Gough, 2000 Lyman, 1990
Note. Number in parentheses after site number designates the total number of measurements taken on specimens from the site. a Charcoal. b Burned annual-plant stems. c Bison bone collagen. d Based on temporally diagnostic artifacts. e 8600 F 65 (WSU-1422; humus); 8700 F 400 (C-827; burned bone); 8720 F 200 (WSU-1709; bone collagen). f 10,010 F 60 (Beta-133664; charcoal); 10,130 F 60 (Beta-133665; charcoal); 10,160 F 60 (Beta-133663; charcoal); 10,180 F 40 (Beta-124167; charred material); 10,680 F 190 (Beta-133650; charcoal).
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by a congeneric or conspecific prehistoric specimen. The comparative standard is set at 0.0; a negative value indicates that a prehistoric specimen is smaller than the comparative standard, and a positive value indicates that a prehistoric specimen is larger than the comparative standard. The conversion allows one to simultaneously plot multiple dimensions from multiple specimens. Traditionally, the difference in values is plotted on the x-axis and each dimension holds a unique position on the y-axis. These graphs allow one to detect allometric relationships and general evolutionary patterns. Because all dimensions are normed and plotted against a comparative standard, a ratio diagram can be constructed so as to measure change in size over time. The difference between the dimension(s) of each comparative specimen and the appropriate comparative standard is plotted on the xaxis; the y-axis is used as a chronometer to show the age of each measured specimen irrespective of the dimension measured. The resulting graph is much like a chronocline, except that multiple dimensions of multiple skeletal elements are plotted on the same graph. The dimensions can, for example, include the width of the distal condyle of the humerus, the length of the first phalanx, and the width of the proximal end of the second phalanx. Because the absolute values of the comparative standard vary across the multiple dimensions included, the resulting graph will not be interval scale if the dimensions measured are not randomly distributed across the time span considered. Even then, ordinalscale trends in size, whether stable or changing, will be indicated by the relative positions of clusters of points across time. A key step in the construction of a chronoclinal ratio diagram involves choosing the comparative standards. Many standard measurements of modern comparative bison are available in the literature (e.g., Speth, 1983; Todd, 1983), and there are also many measurements of late-Holocene prehistoric B. bison (e.g., Lorrain, 1968; McDonald, 1981).
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It is unknown if and when the late Quaternary diminution of bison stopped and their average size stabilized. Both lateHolocene data and modern data are used here as comparative standards because they are not significantly different and because the nature of the prehistoric sample demands it in order that samples of prehistoric measurements be large enough to detect general trends. Some sites produced measureable specimens for which only data from modern comparative bison are available (45GA17, 45WT134); data compiled by Todd (1983) on modern bison from Wyoming are used as the comparative standard for these materials. Other sites produced other kinds of measureable specimens for which only late-prehistoric comparative data described by Lorrain (1968) are available (45KT338, 45KT1362, 45AD104); data on bison from Texas dating to about 2650 14 C yr B.P. are used as the comparative standard for these materials. Three sites (45GR97, 45AS80, 45WT245) produced specimens for which both kinds of comparative data are available. The sites in eastern Washington that produced the collections of bison remains discussed here, the skeletal elements measured in each collection, the number of measurements taken per collection, and the age of each collection are listed in Table 1. All ages are given as uncalibrated 14C yr B.P. Only the bison remains from 45WT245 did not have a radiocarbon age directly associated with them; the age of these remains was determined on the basis of associated, temporally diagnostic artifacts.
Results The chronoclinal ratio diagram suggests that bison in eastern Washington, like those elsewhere on the continent, underwent diminution during the late Quaternary (Fig. 2). All of the 47 plotted measurements dating prior to 8500 14C yr B.P. are larger than average modern bison. Of the 65
Fig. 2. Ratio diagram of dimensions of bison bones plotted against a modern standard (0.0 vertical line). Points with positive values are larger than an average modern bison; points with negative values are smaller than an average modern bison.
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plotted measurements dating later than 2300 14C yr B.P., 41 are larger than the comparative standard (an average modern bison) and 24 are smaller. This suggests that late-Holocene bison may have been slightly larger than modern bison. Other late-Holocene collections of bison remains (e.g., Harkins, 1980) are required to reveal any trends in size change during the last several thousand years. Given that members of the genus Bison are notably sexually dimorphic, and that such dimorphism is apparent throughout the late Quaternary (Hofman and Todd, 2001), it is important to consider whether the apparent decrease in size documented in Fig. 2 is a result of a change in sex ratios. If the early collections comprise only or mostly males whereas the latter collections comprise only or mostly females, then the size trend evident in Fig. 2 could merely reflect a shift in sex ratio. This concern cannot be addressed completely because of a lack of skulls and pubic bones in available collections. Postcranial measurements of latePleistocene, early Holocene bison distinguished by sex are
not available. Such measurements are only available for modern bison. Based on comparative data for modern bison (Todd, 1983), four females and two males are represented by the 45GA17 specimens. Specimens from 45AS80 represent one female and two males. Remains of one adult female and one adult male were recovered from 45WT245. In total, there are five female and four male bison represented in late-Holocene collections measured for this study. This suggests that unless only males are represented in the collections >8500 14C yr old, differential representation of the two sexes is not creating the appearance of diminution in Fig. 2. The only other site for which sex ratios of bison are reported is the 2000 14C-yr-old 45BN161, where remains of three males and two females were recovered (Harkins, 1980). This collection does not alter my conclusion regarding sex ratios of resident late-prehistoric bison. Another consideration is whether or not the chronocline in Fig. 2 is a function of how the skeletal elements and dimensions measured are distributed across the time periods.
Fig. 3. Bivariate scatterplot of selected skeletal dimensions. In both, numbers adjacent to plotted points indicate age of the specimen in phalanx; b, distal humerus.
14
C yr. a, second
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Such a distribution does not seem to be exerting any great influence on the point scatters in Fig. 2. For example, a bivariate scatterplot of the lateromedial width of the proximal second phalanx against the lateromedial width of the distal second phalanx indicates that older specimens are, on average, larger than younger specimens (Fig. 3a). This is the only skeletal element with specimens from most represented time periods, but other elements with less complete temporal representation such as the distal humerus produce similar results (Fig. 3b). There is no evidence that temporal variation in the skeletal dimensions measured is causing the size trend evident in Fig. 2. The final and most critical issue concerns whether the bison remains plotted in Fig. 2 actually represent local bison, or immigrants from elsewhere. As noted above, available evidence suggests that bison in eastern Washington originated in western Montana and Wyoming, and that they came into the northwest through southern Idaho (Plew and Sundell, 2000) and eastern Oregon (Van Vuren and Bray, 1985). Some may have originated in Nevada (Van Vuren and Dietz, 1993) or Utah (Lupo, 1996) and moved north, though that source seems less likely given the apparent rarity of bison there relative to their abundance in Wyoming and Montana. Regardless of the source, the question of whether bison established residency and reproduced in eastern Washington is paramount to concluding that local bison underwent diminution there rather than that big bison immigrated first, and successively smaller bison immigrated as time passed. The hypothesis that resident bison became smaller suggests that remains of neonates and fetuses should be present in the eastern Washington record. Further, if the bison were reproducing locally, then individuals of all ages and both sexes should be present. Harkins (1980) reported that one near-term fetus or neonate and seven adults were represented at 45BN161. Remains of a yearling calf, along with those of two skeletally mature adults, were recovered from 45WT245. A single calf and three adults, are represented at 45AS80, and at least one calf and several adults are represented at 45GR97. Available data do not contradict the hypothesis that bison of all ages and both sexes were resident in eastern Washington during the Holocene. Nor do these data contradict the hypothesis that resident bison were reproducing in eastern Washington and underwent diminution there. But they do not contradict the hypothesis that successively smaller individuals immigrated to the area, either. The last is so because bison could have immigrated, established a small local population, reproduced a few generations, and then been extirpated, followed some years later by another immigration, reproduction, extirpation, and so on. Each successive immigrant population could have produced young that grew to be a bit smaller than their predecessors. A way out of this conundrum is to consider the abundance of bison relative to other ungulates and to consider how those abundances correspond (or not) with independent paleoenvironmental indicators and with the size of individual bison.
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This is so because individual bison size and bison abundance seem to be ecologically interrelated and covariant.
Discussion Thirty-seven years ago, Edwards (1967) said that the reason many mammalian taxa underwent diminution during the Holocene was that human predators selectively exploited the largest individuals. This argument was subsequently shown to be inapplicable on the Great Plains, where numerous bison kill sites indicate that it was not unusual for an entire herd to be killed with no regard for age or sex or size of individual bison (Frison, 1991). McDonald (1981, p. 258) puts a different twist on the human predation mechanism. He argues that throughout the late Pleistocene range of bison, human predation increased selection for faster, more agile, more rapidly reproducing organisms, all of which selected for reduced body size. Further, in McDonald’s view, the nutritionally richer, more open (grassland) environment of the terminal Pleistocene and early Holocene reduced intraspecific competition, thereby reducing selection for large individuals. Guthrie (1970, p. 10) suggested that increased nutrients would increase ‘‘the survival of larger individuals . . . allowing them to contribute more to future generations’’ but also that ‘‘increased intrapopulational competition . . . generally results in a heritable decrease in individual size.’’ A decade later he suggested that during the Holocene on the Plains of North America ‘‘bison were becoming more abundant while also decreasing in body size’’ (Guthrie, 1980, p. 56). A terminal Pleistocene decrease in interspecific competition, plus an increase in short-grass prairies favored an increase in bison numbers (Guthrie, 1980, p. 67). Almost simultaneously, there was an ‘‘opening of the winter bottleneck’’ by which Guthrie (1980, p. 68) meant that the reproductive season of bison was lengthier and there was reduced winter mortality during the Holocene relative to that during the Pleistocene. Both factors resulted in greater bison numbers which in turn increased intraspecific competition and this selected for reduced body size (Guthrie, 1980, p. 69). Butler et al. (1971) suggested that bison were subject to Bergmann’s rule, but Wilson (1974a) showed that the former had no data to substantiate their claim. Wilson (1974a) expressed preference for climatic factors being responsible for size variation in bison, probably because he noted that bison remains from Wyoming dating to the Altithermal indicated that this was the period when local bison became smaller. Thus, concluded that ‘‘a causal relationship is probable’’ between climate and bison size (Wilson, 1974b, p. 96). In light of additional data, Wilson (1978, p. 15) later concluded that ‘‘dwarfing, in fact, took place throughout the Holocene’’ and expressed favor for Geist’s (1971) and Guthrie’s (1970) suggestion that greater intraspecific competition for food ‘‘accounted for some of the Holocene dwarfing.’’
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Interestingly, and with some potential relevance for the earliest bison remains in eastern Washington, Geist (1971, p. 287) proposed a ‘‘dispersal theory’’ which held that ‘‘pioneering [colonizing] populations find a superabundance of qualitatively excellent forage in the vacant habitat that causes high birth rates, high birth weights, high milk production, rapid growth of young, early sexual maturation, [and] large body size of adults close to the genetic maximum.’’ If forage on the edge of a taxon’s range is neither abundant nor of high quality, then I suspect that when that taxon colonizes those peripheral areas individuals born there will be few and they will be small. Paleozoological evidence regarding bison abundance in Texas, near the southern edge of their range, indicates that bison only became abundant there about 1000 14C yr B.P. when grass biomass increased as a result of environmental change (Huebner, 1991). It has long been known that bison were rare prehistorically in eastern Washington relative to their abundance in the Plains. The debate over why the relative abundances should be so has a deep history. Some recent commentators, however, make it sound as if they thought up a new cause for the rarity of bison in eastern Washington when they proposed that prehistoric aboriginal exploitation depleted local populations (Kay, 1994; Martin and Szuter, 1999a, 1999b). This idea, however, has been around for at least seventy years. Kingston (1932, p. 164), for example, wrote that eastern Washington ‘‘presents a curious and rather intricate problem of biological distribution in which hunting by the Indians and physiographic difficulties explain for the most part the scarcity or absence of the buffalo.’’ The physiographic difficulties involved the fact that the likely immigration route is better characterized as a ‘‘filter’’ than as a ‘‘corridor’’ (Simpson, 1940, 1953); thus local recruitment was largely limited to reproduction and only occasionally
immigration. Fifty-five years later Van Vuren (1987) added low overall forage production in eastern Washington to the mix. The modern debate concerns whether Kingston and Van Vuren’s proposed combination of factors or only overhunting by human predators was the cause (Laliberte and Ripple, 2003; Lyman and Wolverton, 2002; Martin and Szuter, 2002). The zooarchaeological record provides no unequivocal evidence that local elk (Cervus elaphus) populations were depressed by human predation (Lyman, 2004a), despite argument to the contrary (Kay, 1994). The relative abundance of elk seems to flunctuate in concert with paleoclimatic conditions; elk remains are abundant when it is cool and moist but less abundant when it is warm and dry (Lyman, 2004b; Lyman and Wolverton, 2002). Bison abundances also fluctuate in concert with paleoclimatic conditions, and those conditions help account for the size of individual bison in eastern Washington. Bison remains dating to the mid-Holocene Altithermal climatic interval of warmth and aridity are more rare in eastern Washington than at any other time during the Holocene (Lyman, 1985; Schroedl, 1973). Based on 91 collections of zooarchaeological remains from sites throughout eastern Washington with good temporal control, bison remains are most abundant relative to other ungulates during the terminal Pleistocene, initial Holocene, and late Holocene, and least abundant during the middle Holocene (Fig. 4). Given that early, middle, and late Holocene sediments have been differentially sampled (e.g., Lyman, 1992, 2000), it is important to note that there is no correlation between the proportion of ungulate remains representing bison and either the number of radiocarbon ages per period plotted in Fig. 4 or the number of assemblages of remains per period ( p>0.5 for both). There is no evidence that changes in human subsistence pursuits or technology resulted in decreased pursuit of
Fig. 4. Abundance of bison remains relative to remains of all ungulates over the past 10,500
14
C yr in eastern Washington.
R.L. Lyman / Quaternary Research 62 (2004) 76–85
bison between 8000 and 2500 14C yr B.P. (Lyman, 1992; Schroedl, 1973). Rather, there were few bison to pursue. As part of their response to local environmental change, local bison populations were small or nonexistant between 8000 and 2500 14C yr B.P. Thirty-two sites dating between 6000 and 8000 14C yr B.P. failed to produce remains of bison. This 2000-14C-yr gap in the local record argues against local bison undergoing in situ diminution during the last 12,000 14C yr (Fig. 2). Rather, it is more parsimonious to conclude that the local early Holocene population of large bison was extirpated about 8000 14C yr B.P. and that after about 6000 yr 14C B.P. successively smaller bison immigrated sporadically, but no resident population surivived very long or at any significantly large size until after 2500 14C yr B.P. Although any resident population established by post-6000-14C-yr-B.P. immigrants may have undergone some degree of diminution, data necessary to assess this possibility are currently unavailable. Grass (as a proxy measure of bison nutrition) was most abundant during the terminal Pleistocene, early Holocene, and late Holocene, and rare during the middle Holocene (Chatters, 1998). Bison that immigrated to eastern Washington during the terminal Pleistocene and early Holocene were of large size because they originated in herds of large individuals; they maintained their large size in eastern Washington because forage was abundant and nutritious. During the late Holocene, bison were of small size due in part to their origins in herds of small individuals. Their abundances were low due in part to intraspecific competition. The low quality and quantity of forage during the middle Holocene significantly depressed the local bison population. That depression may have been exacerbated by human predation, though evidence of such is nonexistant. At present, the most parsimonious account of available data is that the local bison population was completely extirpated about 8000 14C yr B.P. Bison sporadically immigrated between 6000 and 2500 14C yr B.P., but were unable to establish a local population because of poor forage conditions and human predation. After 2500 14C yr B.P. they were again locally abundant because climate had returned to the relatively cool – moist conditions of modern times and grass biomass had increased. Between about 4000 and 2500 14C yr B.P., climates were the coolest and most moist of the entire Holocene (Chatters, 1998), raising the question of why bison were not more abundant in eastern Washington at this time. Modern bison consume mostly graminoids, often eating higher proportions of grass than would be predicted on the basis of grass availability on the landscape; they eat very small amounts of forbs and woody species (Knapp et al., 1999, and references therein). Long-term observations indicate that bison graze preferentially in recently burned areas because of higher grass productivity and decreased forb cover (Knapp et al., 1999). The abundance of charcoal in lake sediments is known to directly correlate with fire
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history (Gardner and Whitlock, 2001, and references therein). Charcoal abundances in such sediments in eastern Washington, northern Idaho, and western Montana indicate fires were relatively infrequent between about 4000 and 2000 14C yr B.P. but were more frequent after that time (Chatters, 1998, and references therein). If bison immigrated to eastern Washington from southwestern Montana and western Wyoming, and passed through southern Idaho, then the increased incidence of (anthropogenic?) fires over the past 2000 or so 14C yr may have enhanced forage quality along that route and thus allowed larger numbers of bison to immigrate. Relatively low abundances of bison during the last 500 14C yr may be a result of increased predation brought on by the acquisition of horses about A.D. 1730 by native peoples of eastern Washington (Haines, 1938). Temporal resolution of the collections of faunal remains dating to this late prehistoric period is insufficient to determine if the depression in bison abundance is in fact coincident with the appearance of horses.
Conclusion The late Quaternary record of bison in the Great Plains— the heart of bison range during this period (Graham and Lundelius, 1994)—indicates that this taxon underwent diminution over the past 12,000 to 15,000 14C yr. Bison occurred in areas peripheral to the Plains off and on throughout this period, seeming to fluctuate in abundance more or less in concert with changes in environments, but also perhaps at least in part in response to human predation. The question that arises is whether bison in these peripheral areas also underwent diminution independently of that which took place on the Plains. Thus, the alternative hypothesis of diminution in the heartland followed by immigration to the subject peripheral area must be considered. The prehistoric record of bison in eastern Washington indicates that this taxon was present there over much of the late Quaternary, but it was often quite rare and apparently absent during significant portions of the hot – dry middle Holocene when much diminution took place among bison on the Plains. The absence of evidence for their middleHolocene presence in eastern Washington means that the diminution elsewhere followed by subsequent immigration hypothesis cannot be discounted. Only if it can be shown that bison were consistently present in eastern Washington throughout the past 10,000 or so years can the hypothesis of local diminution be considered possible.
Acknowledgments I thank D. R. Brauner, J. R. Galm, S. Gough, K. Lupo, D. Schmitt, and G. F. Schroedl for the opportunities to study the collections discussed here, and C. R. Harington and especially M. Wilson for insightful comments on an early
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Sternberg, C.H., 1903. Elephas columbi and other mammals in swamps of Whitman County, Washington. Science 17, 511 – 512. Tipper, H.W., Jizba, Z.V., Konizeski, R.L., 1951. A bison from eastern Washington. American Journal of Science 249, 318 – 322. Todd Jr., L.G., 1983. The Horner Site: Taphonomy of an Early Holocene Bison Bonebed. Unpublished Ph.D. dissertation. Univ. of New Mexico, Albuquerque. Van Vuren, D., 1987. Bison west of the Rocky Mountains: An alternative explanation. Northwest Science 61, 65 – 69. Van Vuren, D., Bray, M.P., 1985. The recent geographic distribution of Bison bison in Oregon. Murrelet 66, 56 – 58. Van Vuren, D., Dietz, F.C., 1993. Evidence of Bison bison in the Great Basin. Great Basin Naturalist 53, 318 – 319. Wilson, M., 1974a. History of the bison in Wyoming, with particular reference to early Holocene forms. In: Wilson, M. (Ed.), Applied Geology and Archaeology: The Holocene History of Wyoming. Report of Investigations, vol. 10. Geological Survey of Wyoming, Laramie, pp. 91 – 99. Wilson, M., 1974b. The Casper local fauna and its fossil bison. In: Frison, G.C. (Ed.), The Casper Site: A Hell Gap Bison Kill on the High Plains. Academic Press, New York, pp. 125 – 171. Wilson, M., 1978. Archaeological kill site populations and the Holocene evolution of the genus Bison. In: Davis, L.B., Wilson, M. (Eds.), Bison Procurement and Utilization: A Symposium. Plains Anthropologist Memoir, vol. 14, pp. 9 – 22. Wilson, M., 1980. Morphological dating of late Quaternary bison on the northern Plains. Canadian Journal of Anthropology 1, 81 – 85. Wilson, M., 1996. Late Quaternary vertebrates and the opening of the icefree corridor, with special reference to the genus Bison. Quaternary International 32, 97 – 105. Wyckoff, D.G., Dalquest, W.W., 1997. From whence they came: The paleontology of southern plains bison. In: Bement, L.C., Buehler, K.J. (Eds.), Southern Plains Bison Procurement and Utilization from Paleoindian to Historic. Plains Anthropologist Memoir, vol. 42, pp. 5 – 32.