Bison antiquus from Kenora, Ontario, and Notes on the Evolution of North American Holocene Bison Jerry N. McDonald and George E. Lammers Bison antiquus occidentalis ,an extinct taxon that was the evo¬ ABSTRACT lutionary link between the late Pleistocene North American An associated skeleton o fan adul tmale Bison antiquu socciden- steppe bison. Bison antiquus ,and the extant North American talis from Kenora ,Ontario ,is radiocarbon dated at 4270±65 yrs bison. Bison bison. Sediment from within the cranium of the BP ,making i tthe younges tunequivoca lrecord for the species .It Kenora bison was radiocarbon dated at 4850±60 years before also extends the range 280 km north and 65 km east of the previ¬ present (yrs BP) (Beta-3779) and ribs of the bison were radio¬ ously documented limits fo rthe genus in the western Grea tLakes area. The Kenora bison died in a shallow pond in an oak-pine carbon dated at 4270±65 yrs BP (DIC-3381). Sediment from woodland ;i twas experiencing nutritiona lstress a tthe time o fits the site yielded pollen ,plant macrofossils ,and mollusks ,which death .Both halves o fthe mandible had been fractured by trauma provided information about the regional environment at the earlie ir nlife. time the bison lived. A review o fmorphologica lchange in bison during the Holocene shows tha tlate rbison ,Bison bison ,were absolutely smaller ,had The Kenora bison is significant for several reasons. Associ¬ absolutely shorter limbs ,and had more robus tupper limbs and ated skeletons of Bison antiquus occidentalis are uncommon more gracile lower limbs ,relative to length ,than did the earlier from othe rthan archaeologica lcontexts ,and descriptions o fthe bison .Bison antiquus .Compared with tha to fBison antiquus ,the postcrania lbones of such associated specimens are rare .The thoracic limb o fBison bison became elongated relative to the pel¬ Kenora bison was found farther north and east than any bison vic limb ,and limb length increased relative to skul lsize .Relative to the norm in Bison antiquus ,these traits diverged further in the previously known in the western Great Lakes region ,and it is a wood bison. Bison bison athabascae ,than in the plains bison. rare record of bison in Ontario. The radiocarbon age of 4270± Bison bison bison .The pattern so fpelage developmen tand social 65 yrs BP is the youngest known for Bison antiquus. Patholo¬ behavio rin Bison bison bison h, owever a, re more distan tfrom par¬ gies of the face and teeth provide information about the life his¬ alle pl attern spostulated fo rBison antiquu santiquu sthan are pat¬ terns o fpelage developmen tand socia lbehavior in Bison bison tory of this animal and its biophysical condition at the time of athabascae. death ,and contribute information abou tpatterns o fpathology in the taxon. Although most characters of the Kenora bison skul lare o faverage to greater-than-average size for Bison anti¬ Introduction quus occidentalis ,the postcrania lbones are smaller than might be expected ,a circumstance tha traises questions abou tthe pat¬ The skull ,mandible ,14 teeth ,and 58 postcrania lbones of an tern of coevolution of the postcranial skeleton and skull in bi¬ adult male bison were collected late in the 1970s during dredg¬ son during th eHolocene. ing operations in a peat bog near Kenora, Ontario. This bison, In this paper we (I) describe the partial bison skeleton, pol¬ now in the collections of the Manitoba Museum of Man and len spectrum ,and invertebrate remains from the Kenora site; Nature ,and cataloged as MMMN V-1914 ,has been referred to (2) review the zoogeographical, paleoecological, and evolu¬ tionary context of the Kenora bison; (3) document the pattern Jerr Ny M. cDonald R,esearc hAssociate D, epartmen ot Pfaleobiology, of skull and limb coevolution seen in North American Ho¬ Nationa Ml useum o Nf atura Hl istor yS,mithsonia nInstitution W, ash¬ ington ,D.C .20560-0121 ;and McDonald & Woodward Publishing locene Bison, and, on the basis of the observed pattern of co¬ Company 4, 31-B Eas tColleg eStreet G, ranville O, hio 43023 G. eorge evolution, (4) comment upon the position of the wood bison. LEa.mme (rdseceased). Bison bison athabascae ,in the evolutionary sequence of North 83 84 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY Figu r .Ie_Top T:h Keenor aO, ntario a,re sahowin gth leocatio no tfh Keenor baiso nsite B.ottom S:outhwestern Ontari oan dadjacen rtegion showin gth eKenor abiso nsit ean dothe prlace ms entione di nth etex i tnregional context. American Bison. All dates are given in uncalibrated radiocar¬ MMM NManitob Ma useum o Mf a nan dNatur eW, innipeg M, anitobaPIN Paleontologica Ilnstitute R, ussia nAcadem yo Sfciences M, os¬ bo nyear sbefor epresent. Acknowledgments. —We extend our appreciation to Nick cow. Serduletz of Kenora, Ontario, for salvaging the Kenora bison The Kenora Bison skeleton and donating it to the Manitoba Museum of Man and Nature. Gary Haynes and Clayton E. Ray reviewed this paper Location and Description of the Kenora Site and provided many helpfu lcomments ,but the authors retain full responsibility for the facts and opinions as finally pre¬ The bison skeleton was recovered during a period of several sented. years by Nick Serduletz while dredging a peat bog located in Abbreviations .—The following museum abbreviations are the watershed of Laurensons Creek 3.0 km east of Kenora .The used: dredging extended to a depth of approximately 4.5 m ,at which NUMB E9R3 85 0 20 40 0 0 0 0 20 40 60 0 0 0 0 0 0 0 0 0 0 0 20 0 0 0 0 20 400 0 0 0 0 0 0 Percent | Sum Figur e2 —. Polle nan dspor ediagram sfo Hr aye sLak ean dth eKenor abiso nsite O, ntario s,implifie dfrom McAnd (r1e9w8s2). 86 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY Figur e3.—Kenor abiso nsku li lndorsa vliew (.Scal ebar= 5cm.) covered pond one or two meters deep that was bordered with a covered. The skull is nearly complete, lacking only the nasals, marsh dominated by shrubs; wetland conifers (spruce ,larch) most of the right premaxilla ,and most of the palatine process grew nearby” (McAndrews ,1982:47^8). of the left maxilla (Figures 3-5). Minor fragments are missing Five species of mollusks were collected from within the cra¬ from other bones of the skull. Both halves of the mandible are nium of the bison. These included the small spire (Amnicola complete (Figures 6-9). Teeth present include RP4-M3, walkeri) ,three-keeled valve {Valvata tricarinata) ,ribbed valve LM2-M3 ,Rp4-m3 ,and Lp4-m3 .Postcrania lelements present (Valvata sincera sincera) ,modest gyraulus {Gyraulusparvus), include the atlas; axis; cervical vertebrae 3, 5, and 7; thoracic and shiny pea clam {Pisidium nitidum). Although the pond of vertebrae (1), 2, 4, 5, 11, 9, and ?10; lumbar vertebrae (2), (3), 4270 yrs BP probably was intimately associated with upper (4), and (5); sacrum; ribs Rl, (R?2), (R3), (R4), (?R5), R6, R7, Laurensons Creek ,similar to the situation o fthe depression to¬ (R8), Rll, (R12), LI, (L2), (L3), (L4), (L5), L6, L8, L9, LIO, day ,the gastropods from the bison skul lsuggest that the body Lll, LI3 or LI4; manubrium; (stemebrae); R scapula; R radi¬ of water containing the skeleton was relatively slow moving oulna; R radial carpal; R metacarpal; L scapula; L radioulna; L and pond-like. The Kenora site is within the present range of fused 2+3 carpal; proximal thoracic sesamoid; proximal tho¬ the five mollusks (Clarke, 1981), and their presence 4270 yrs racic first phalanx; medial thoracic phalanx; bony pelvis ,R fe¬ BP does not require ecologica lconditions substantially differ¬ mur, R tibia, R calcaneum; L femur, L tibia, two proximal pel¬ ent than exist today .At the same time ,these mollusks are wide¬ vic phalanges .Specimens in parentheses are incomplete ;those spread today ,and their sympatry 4270 yrs BP would have been no tin parentheses are either complete or nearly complete .Cra¬ possible even under environmenta lconditions somewha tdif¬ nial, vertebral, and rib measurements are given in Tables 1-3; feren tthan those o ftoday. those for other elements are given in the Appendix. The bones and teeth of the Kenora bison are well preserved SiCELETAL AND DENTAL MATERIAL COLLECTED and exhibit no evidence of pre-entombment or diagenetic weathering a, brasion t,rampling c, rushing o, rchew-gnaw dam¬ The skull, mandible, 14 teeth, and 58 complete or partial age. The bones do vary in color, ranging from light to dark postcrania lskeleta lelements o fthe Kenora bison have been re¬ brown ,which suggests exposure during diagenesis to different NUMB 9E3R 87 Figur e4.—Kenor abiso nsku li lnlatera vliew l,e fstide (.Scal ebar= 5cm.) concentrations o fdecaying organic matter or to groundwater Females of al lspecies of North American bison have relatively with differing concentrations of oxygen. All breakage of the gracile skulls .Typically ,among the short-homed North Ameri¬ bone scan be attributed to damage during excavation. can bison ,the fronta land fronto-parieta lsutures are unfused Age and Sex. —The Kenora bison was approaching senes¬ over most or all of their length, the supraorbital foramina are cence at the time of its death. All teeth were in full to advanced not bridged by bone at the level of the orbits, the hom cores do wear. The RMl and both mis either had lost or were losing, by not possess burrs at the base, the hom cores grow straight or wear ,the ename lfossettes within the dentine field. spira lanteriorly around the longitudina laxis ,and the antero¬ Severa lmorphologica lfeatures identify this specimen as a posterior plane of the bases of the hom cores is either rotated male .The skul lis robust ,the fronta land fronto-parieta lsutures forward relative to the plane of the frontals or paralle lwith the are obseured by fusion over most of the dorsal surface of the plane o fthe frontals (McDonald ,1981). cranium ,the supraorbita lforamina are bridged by bone at the Taxonomic Identity. —The Kenora bison is referred to Bi¬ level of the orbits, the horn cores possess distinct burrs at the son antiquus occidentalis on the basis of diagnostic criteria base, the horn core growth is spiraled posteriorly around the given in McDonald (1981). The hom cores of the Kenora bison longitudina laxis ,and the anteroposterior plane of the base of are triangular in cross-section and symmetrica labout the dor- the hom cores is rotated rearward relative to the fronta lplane. soventral axis at the base. The tips of the hom cores are ellipti- Figur e5.—Kenor abiso nsku li lncauda vliew (.Scal ebar= 5cm.) 88 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY Tab l1—e. Sku (llinea mr easuremen t imsn m a,ngula mr easuremen t idsnegree s oK)fenor baiso {nBiso anntiquus occidentali sc)ompare wdi tth meean ossifmil amreasuremen tf osrth esrhorter-home Ndor tAhmerica bnison. (Dat ar ferom McDonal d1,98 1e,xcep mt easuremen tfso Bran fbfiso nw, hic har ferom Haringto n1,98 4v;alue isn sing lpearenthese asr estimate dimension fso trh ceomple tceharacte rv;alue idsnoub lpearenthese asr feo inr ¬ comple tceharacte rasn ad rperovide sdole l tydoescrib teh aemou n obtfon peresent.) Measurement N' umbe o rsfpecimen ms easure dvarie fsrom t1 o41. N^ umbe o rsfpecimen ms easure dvarie fsrom 2 t5 o91. N^ umbe o rsfpecimen ms easure dvarie fsrom 5 t6 o142. Numbe o rsfpecimen ms easure vdarie fsro m t 71o1. *Denot esasmp cleontaini no gsn 1plyecimen. ca lin cross-section .Growth along the longitudina laxis is mod¬ arched longitudinal axis of the core, the posterior margin is estly spiraled posteriorly, the posterior margins of the horn straight ,and the anteroposterior plane o fthe horn cores is par¬ cores are slightly concave in dorsa lview ,and the anteroposte¬ alle lwith the fronta lplane .The horn cores of Bison bison grow rior plane of the horn core is rotated rearward relative to the straigh talong the longitudina laxis ,they are straigh talong their plane of the frontals .The length along the upper surface of the posterior margins ,and they are less slender (i.e. ,length :basal horn core ,tip to burr ,is approximately 315 mm ,wel lwithin the circumference ratio is less )than in Bison antiquus occidentalis. range of 186-392 mm known for the taxon. In Bison antiquus Pathologies.— Pathologica lconditions are apparent in the antiquus, the tips of the horn cores are typically cordiform to bones and teeth of the skul land in both halves of the mandible. triangular in cross-section and possess a distinct groove on the The RP2 and P3 were missing during at least the later part of dorsal surface, growth is straight (not spiraled) around the the life of the Kenora bison ,as evidenced by the ossified alveo- Tab l 2e—. Measuremen t(si nmm o )vfertebra (eexclusiv oe aftla asn daxi so )Kfenor baison (.Measurements confor mt tohos deescribe fdo brovid ivsno dne Dnriesc h1,97 6M. easuremen t ipsnarenthese asr estimate di¬ mensio fnto hcsroemple cthearacter.) Measurement NUMB 9E3R 89 Figur 6e.—Kenor baiso nmandib li endors avliew (.Sca lbear= c5m.) lar trough that would have accommodated the roots of these teeth (Figure 6). A hole is present in the dentine fields of LM3. The insertion tubercle for the masseteric muscle on the right mandible is conspicuously larger than the same tubercle on the Figur e7.— Le f(ttop a)n drigh (tbottom h) alve so Kf enor abiso nmandibl ei nlatera vliew (.Scal ebar= 5cm.) SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY Figur e8.— Le f(ttop a)n drigh (tbottom h) alve os Kf enor abiso nmandibl ei nmedia vliew (.Scal ebar= 5cm.) NUMB 9E3R 91 Tab l 3e—. Measuremen t(si nmm o )rfib o sKfenor baison (.Measuremen tisn doub lpearenthese as rfeo inr comple telemen tasn ad rperovide sdole l tydoe¬ scrib teh aemoun o tfh elemen ttha ipstresent.) nRuibmber Figur e10.—Th evolutionar fyol do 5f000^00 0radiocarbo nyr BsP w, hen th evolutio on Bfiso bniso fnrom Biso anntiquu ws a rsapid lcyomplete (dfrom McDo n1a9l8d1,). Discussion The Kenora bison lived at the time when the nucleus of the North American bison range was anchored on the northern Great Plains and when the final, rapid steps in the evolution of Bison bison from Bison antiquus were taking place (Figure 10) (McDonald, 1981). The radiocarbon age of 4270±65 yrs BP is the youngest such date of which we are aware for a specimen that can be assigned with confidence to Bison antiquus occi- dentalis. Harington (1984) reported a partial cranium from bone on the lateral ,ventral ,and media lsurfaces of the healed Banff, Alberta, dated at 3240±90 yrs BP (1-11638), which he fracture, and a subcircular hole leading to the interior of the referred to Bison bison occidentalis ,equivalent to Bison anti¬ bone opens through the ventra lsurface of the fractured area. quus occidentalis as used in this paper. Having examined the Rostral to the level of Rml, the body of the right ramus of the Banff cranium ,we believe that this specimen is referable to Bi¬ mandible is conspicuously wider than that of the left, and the son bison. All measurements that can be obtained from the diastema is displaced ventral to its normal position. Root ca¬ skull of the specimen are within the range of Bison bison (Ta¬ nals for the missing Rp2 and Rp3 are present in the alveolar ble 1), and the qualitative aspects of characters still visible on trough ,indicating that these teeth were present at ,or at least the specimen also are consistent with those typica lof Bison bi¬ shortly before, the time of death. The medial side of Rm3 was son.Measurements of the Kenora bison skull are all within the shorn away in life, and the dentine surface thus exposed was range documented for Bison antiquus occidentalis ,but some worn smooth by use. The surface of the healed left body of the are outside the observed ranges for Bison antiquus antiquus mandible is much smoother than that of the right ,and no dis¬ and Bison bison (Table 1; cf tables 21, 25, 29, and 34 in Mc¬ placement of the diastema occurred. A circular opening is lo¬ Donald, 1981). Most features of the Kenora bison skull con¬ cated between and below Lp4-Lml; both of these teeth are form harmoniously with both qualitative and quantitative at¬ loosely rooted in the alveolus ,and an abnormally long gap sep¬ tributes of the skul ltypica lof Bison antiquus occidentalis. The arates the two teeth .The bone surrounding the opening is dense horn cores are longer and more slender (i.e. ,long relative to di¬ and does not appear to have been affected by infection at the ameter at base) ,and the dorsa lpart of the cranium is narrower time of death. Root canals for the missing Lp2 and Lp3 are than typica lspecimens ,however .The breadth o fthe base o fthe present ,bu tthey are shallow—a condition tha tsuggests either cranium is near the mean for the taxon. displacement of the teeth or modification of the roots by the Al lmeasured dimensions of the long bones of the Kenora bi¬ trauma to the mandible or by its subsequen thealing .Ename lon son are smaller than the observed means for the limbs of Bison the medial surfaces of Lp4, Lml, and Lm3 had been chipped, antiquus occidentalis except the anteroposterior diameter of part of the medial wall of the rostral cusp of Lm3 is missing, the femur ,which is slightly larger (within one standard devia¬ and holes are present in the dentine fields of Lm3 and Lp4. The tion ( 5 )) than the observed mean for the taxon (Table 4). The tooth rows of both halves of the mandible are unusually sinu¬ length of the radius is >ls, and that of the femur and tibia are ous ,probably reflecting modification o fthe tooth alignmen tby between l.s and 2s ,of the mean for Bison antiquus occidentalis th efracture san dsubsequen ht ealing. long bone lengths ,whereas the length of the metacarpa lin the No pathologic conditions were noted in any of the postcra- Kenora bison is smalle rthan the heretofore-documented mini¬ n ibaol nes. mum for the taxon. The transverse diameters of all four limb 92 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY Tab l 4e—. Lim bmeasuremen t(si nmm f)o srhorter-horne dNort hAmerica nbison m, ale osnl y(.Fo erac htaxon, to rpo wmis ea mn easuremen bt,otto mro wsistandar deviatio nD.a tfaro mMcDonal d1,98 1A.P=anteroposterior diamete TrR; =transver sdeiamete Lr^;^rotation laelngt *h=; samp wle iot hsn 1plyecimen.) Species bones ,and the anteroposterior diameter of the femur ,are near bones of the thoracic limb are shorter relative to those of the the mean for Bison antiquus occidentalis ,whereas the antero¬ pelvic limb ,a condition resembling more nearly the limb pro¬ posterior diameters of the shaft of the radius ,metacarpal ,and portions of Bison antiquus antiquus than Bison bison bison. tibia are >1 ̂from the observed mean for Bison antiquus occi¬ (The thoracic limb:pelvic limb ratio was lowest (0.81) in Bison dentalis. At the same time, however, the dimensions of the antiquus antiquus, greater (0.83) in Bison bison athabascae, Kenora bison long bones are also within the ranges observed still greater (0.846) in Bison bison bison, and greatest (0.854) for Bison bison bison. The two thoracic limb bones of the in Bison antiquus occidentalis (i.e., the thoracic limb, relative Kenora bison are >1 ̂shorter than the mean, whereas the two pelvic limb bones are very near the mean for Bison bison bison. to the pelvic limb ,was shortest in Bison antiquus antiquus and The diameters of al lfour bones are near the mean for Bison bi¬ longest in Bison bison occidentalis). Bison bison occidentalis son bison. All of the long bones of the Kenora bison are also demonstrated the greatest variation in limb dimensions, smalle rthan the documented range fo requivalen tcharacters in which is not unexpected given its dynamic role as a decana- male Bison bison athabascae. lized evolutionary bridge between two relatively canalized spe¬ The fact that the Kenora bison lived near the end of the Bison cies (Tables 5 ,6). antiquus lineage ,as Bison antiquus was changing rapidly into ZOOGEOGRAPHIC SIGNIFICANCE. —The Kenora bison is a Bison bison (Figure 10) ,suggests that the Kenora bison should range record fo rthe subspecies ,species ,and genus .The nearest be relatively small bodied compared with the mean for Bison known remains of bison from natura lcontexts are of Bison an¬ antiquus occidentalis .Except for the horn cores ,which are rel¬ tiquus antiquus from Betula Lake ,Whiteshel lProvincia lPark, atively long for the subspecies, the cranium of the Kenora bi¬ Manitoba, and Bison antiquus occidentalis from Nicollet son generally conforms to this expectation of small size. The Creek, Itasca State Park, Minnesota, respectively 65 km west long bones are shorter than migh tbe expected when compared and 280 km south of Kenora (Figure 1) (Cleland, 1966; Shay, with the size of the skull. If the evolutionary reduction in all 1971 ;McDonald ,1981) .Prehistoric Bison bison remains have skeleta lelements were taken to be isometric ,these long bones been recovered from the Hungry Hal larchaeologica lsite along would probably be identified casually as those of Bison bison the east side of Rainy River near where it discharges into Lake bison had they been found without the associated skull .In gen¬ of the Woods, Ontario (Moore, 1975). Bison from Hungry Hall eral, however, the long bones are broad and robust relative to are north and east of the known natural range of Bison bison; their length ,a condition suggesting that the Kenora bison was they probably were imported by Indians from the nearby bison more heavily built than Bison bison bison. In addition, the range. Tabl e5.— Ratio so afnteroposterio arn dtransvers ediameter st olength so lfon glim b ones (.Dat afrom Mc¬ Donal d1,98 1a p,sresente idTnab l4Ae.P^anteroposterio driamete Tr;R=transvers deiamete Lr;=rotational lengt h*;=samp lwei tohn s1lpyecimen.) Species NUMB E9R3 93 Tab l6—e. Ratio bsetwee snelecte cdrani aln ldim biometri cfso srhorter-home Ndor tAhmerica Bniso nm, ales onl y(D. a tfaro mMcDonal d1,98 1F.W=lea sfrtont ablreadt hN;-Pm=nasio tnporemaxill aB;W=bread tahcross bas oe sfku a lmlt astoi dprocesse sT;L=combine drotation alel ngt ho hfumeru sr,adiu sa,n dmetacarpa Pl;L= combine rdotation alelng t ofhefmu tri,bi a,n md etatarsa Hl;CL^ho mco rlength.) Species Paleoecological Significance. —During the Atlantic cli¬ tral parts but not the dorsolateral part. The incompletely sev¬ matic episode (ca. 8500-5000 yrs BP) the regional vegetation ered left body could have functioned as the splint that held the of extreme southwestern Ontario was dominated by a mixed rostral parts of both halves of the mandible in place while they forest. Early in the episode, when conditions were cooler and healed. Healing occurred with slight ventral displacement of more mesic, conifers dominated. By the end of the episode, the diastema o fthe righ tbody. when the warming and drying trend reached its maximum ,de¬ In addition to the pathologic conditions discussed above ,the ciduous broadlea fspecies were more prominen tthan conifers broken teeth conform to a pattern o ftooth pathology seen in the in the regiona lforest ,and open areas supporting grasses and dentition of numerous musk oxen (Ovibos moschatus) from other herbs presumably reached their maximum .The onset of Greenland, the Arctic islands of Canada, and Alaska (Henrich- the Sub-Boreal climatic episode (ca. 5000-2750 yrs BP) sen, 1981, 1982; Anne Gunn, pers. comm., 1987; McDonald, brought a cooler ,more mesic climate ,which led to a closed for¬ unpub. data). The usual pattern of this pathology is that holes es twith conifers increasing in importance relative to deciduous exist in the dentine field in correlation with broken teeth. In broadl esapfecies. musk oxen ,the enamel walls of teeth are often conspicuously The parkland vegetation tha tcharacterized the Kenora region thin in individuals exhibiting the condition .The etiology of this around 5000 yrs BP, with the deciduous browse and sparse condition is not known, but it is believed to result from inade¬ grasses ,would have afforded the bes tloca lenvironmenta lcon¬ quate or imbalanced intake of (= access to) essentia lnutrients, ditions available during the Holocene for supporting popula¬ such as calcium or phosphorus. The hypoplastic condition of tions of bison .This does not mean that the population of bison the dentine and ename lappears to weaken the teeth to the ex¬ in the Kenora region was either large or permanent. The tent that norma luse results in tooth breakage .In musk oxen ex¬ Kenora bison does no tpossess morphologica lcharacteristics hibiting hypoplasia and subsequent breakage ,irritation o fthe suggesting inbreeding depression in the population (Mc¬ alveolar bone is common, and teeth frequently are loosely Donald, 1981), but it does show evidence of extended nutri¬ rooted and hav enodulated roots. tion satrless. In the Kenora bison, the medial wall of the Rm3 was broken McAndrews’s (1982) interpretation of the death of the to below the gum line (Figure 8), but no irritation of the alveo¬ Kenora bison—that it died in a shallow pond—is consistent lar bone is apparent. Indeed, the alveolar bone is dense and with taphonomic evidence as well as paleobotanic and sedi¬ most of the teeth, except those near the fractures, are well mentary evidence .Similarly ,the bones recovered consist of a rooted. The fact that some teeth exhibit hypoplasia and are random array of elements, not the patterned collection that chipped or more extensively broken suggests that the Kenora would be expected had the bison been killed by predators or its bison was experiencing modest long-term nutritiona lstress. carcass scavenged, or had flowing water sorted the bones This stress could have been caused by the inadequate supplies (Voorhies, 1969; Haynes, 1980, 1982; Shipman, 1981). All of the requisite nutrients in the formerly glaciated, shallow- breakage in the bones appears to be the result of excavation by soile dKenor area. heavy machinery rather than the result of battering in a high- energy fluvial environment. Some of the elements appear to hav eweathere dafte erxcavation. Gross Allometry in Holocene Bison Evolution Interpretation of Pathologies. —Two pathologic condi¬ The evolution of limb length and proportions in North Amer¬ tions appear to be represented in the Kenora bison .A trauma to ican Holocene bison is summarized in Tables 4 and 5. The most the face appears to have broken the bodies of both halves of the general tendency was toward absolutely shorter limbs, with mandible and dislodged the R/L? P2 and P3. A sharp blow to more robust (i.e. ,greater shaft:length ratio) upper limb bones the right half of the mandible, along or immediately caudal to and more gracile (i.e., lesser shaft:length ratio) metapodials, the diastema ,could have produced the fractures that broke the relative to length .Superimposed on this pattern was a trend to¬ bone completely through ,allowing i tto become displaced ,and ward the evolution of longer thoracic limbs relative to pelvic bent the body of the left side of the mandible to the left, pro¬ limbs .Both of these observations apply to the derivation of the ducing a green-stick fracture tha tseparated the media land ven¬ late Holocene Bison bison from the late Wisconsinan Bison an- 94 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY tiquus. In other ways ,the pattern of limb evolution in Bison bi¬ The Evolutionary Place of Bison bison athabascae son athabascae diverged from that in Bison bison bison. The latter evolved to absolutely smaller size ,with the greatest rela¬ The evolutionary position of Bison bison athabascae has tive reduction of length taking place in the upper limb bones been a subject of scholarly and practica lconcern for a century.The existence of a form of extant bison in north-centra lCanada and the least occurring in the metapodials .In Bison bison atha¬ bascae, the femur and humerus evolved to be only slightly different from the plains bison .Bison bison ,has been known to (l%-3%) shorter than in Bison antiquus antiquus, the radius science since at least 1829 (Richardson, 1829; Hind, 1859). and tibia remained abou tthe same size ,and the metapodials be¬ Most early writers regarded the northern bison as little more cam loenger. than a phenotypic variant ,or merely a margina lpopulation ,of The skul lof bison became smaller during the Holocene ,and the plains bison .Rhoads (1897) ,however ,regarded the north¬ the proportions of some characters altered, which produced ern ,or wood ,bison to be taxonomically distinct and gave it the changes in shape. The most obvious changes were those in¬ name Bison bison athabascae .Scientific and politica lopinion volving the horn cores, which became smaller, directed more on the taxonomic status of the wood bison has vacillated fre¬ caudally, and rotated caudally. The cranium became more quently during the last century .There is no question ,however, domed on the dorsal surface, and the occipital condyles pro¬ that the bison native to the parklands of north-centra lCanada jected less far from the occiput. The skull became shorter and, during the nineteenth and twentieth centurie swere demonstra¬ on the dorsa lsurface ,narrower in nearly equa lproportions ;but bly larger than the contemporary plains bison .Here we follow the rate of shortening exceeded the rate at which the base of the McDonald’s (1981) premise that it is scientifically useful and cranium narrowed ,leaving the ventra lside of the cranium pro¬ valid to designate the wood bison as taxonomically distinct, a portionately wider in Bison bison than it had been in Bison an¬ position recently reinforced by van Zyll de Jong and others tiquus. (1995). When the evolutionary changes in the limbs and skull of bi¬ The wood bison became protected by law in 1891, and two son are correlated ,the pattern that emerges is a trend toward a years later a preserve was created as a refuge for the remaining relatively smaller head (with conspicuously allometrically populations. The total population of wood bison was about shorter horns) set relatively lower on a body whose limbs, es¬ 1500-2000 when, between 1925 and 1928—at a time when pecially the thoracic limbs, were becoming relatively longer wood bison were considered by responsible authorities as not (Table 6). Bamforth (1988), drawing upon data in McDonald distinct taxonomically from the plains bison—6673 plains bi¬ (1981) ,observed that limb length in North American bison in¬ son were introduced into Wood Buffalo Nationa lPark .Today, creased relative to overal lbody size during the early Holocene. most zoologists familiar with North American bison recognize The relative lengthening of the thoracic limbs and the only a single extant North American species— Bison bison — changes in the robustness of the limb bones are probably asso¬ containing the two subspecies Bison bison bison (the plains bi¬ ciated with the evolution of relatively more cursorial habits in son) and Bison bison athabascae (the wood bison) (e.g. ,Skin¬ Bison bison than characterized Bison antiquus antiquus. The ner and Kaisen, 1947; McDonald, 1981; Nowak and Paradise, musculature of the upper limbs was exaggerated, relative to 1983; Meagher, 1986; van Zyll de Jong, 1986; van Zyll de Jong that oiBison antiquus antiquus ,paralleling the increased rela¬ et al., 1995). Flerow (1971) considered the wood bison to be¬ tive robustness of the upper limb bones, and the metapodials long to a separate species .Bison priscus ,and he identified it as developed into support structures carrying absolutely less Bison priscus athabascae .Geist (1990) ,however ,recently ar¬ weigh tand bette requipped fo rmore frequen tand versatile mo¬ gued that the wood and plains bison are taxonomically identi¬ bility than had typified Bison antiquus antiquus. The length of cal. the thoracic limb, relative to the pelvic limb, in Bison bison bi¬ Interpretations of the evolutionary origins of Bison bison son is greater than in Bison bison athabascae, probably be¬ athabascae have fallen generally into two categories .The first cause o fselection for greater cursoriality ,more frequen tor spe¬ interpretation is that Bison bison athabascae represents a de¬ cialized agonistic encounters during the rut (which selected for scendan t(either changed or unchanged )form o fbison derived greater forebody strength ,focused on the thoracic limb ,for de¬ from ancestra lpopulations tha toriginated in eastern Siberia or livering and enduring head-to-head clashes and various head- Beringia and dispersed southward. Bison bison athabascae thrusting movements) ,and the essentially obligatory grazing then represents eithe r(1 )the evolutionary and geographic end¬ feeding behavior (requiring regular raising and lowering o fthe point of that parent dispersal or (2) an arrested phase of the relatively low-slung but massive head) of the former ,al las dis¬ evolutionary trajectory that continued as the parent dispersal cussed at greater length in McDonald (1981) .Bison bison atha¬ continued southward to produce ,eithe ralone o rwith introgres- bascae, on the other hand, is less cursorial, probably has less sion with more southerly bison, the plains bison (Skinner and frequen tor less specialized agonistic encounters during the rut, Kaisen, 1947; Guthrie, 1970; Flerow, 1971; Harington, 1984; and has greater opportunity to browse at higher than ground van Zyll de Jong, 1986). A second interpretation is XhdX Bison level with its relatively high-placed head; it has therefore de¬ bison athabascae represents a (changed) descendant form of veloped a relatively lower hump and shorter thoracic limb. bison derived from ancestral populations that originated in NUMB E9R3 95 midlatitude North America and dispersed northward during the of Bison bison athabascae. Lastly, bison collected from along early and middle parts of the Holocene (McDonald ,1981). the Bolshaya Chukochya River span nearly 1.7 million years, Elsewhere, McDonald (1981) has argued against the Berin- and mos tspecimens have been collected withou tstratigraphic gian origin of Bison bison athabascae and Bison bison bison context and have not been radiocarbon dated (Sher, 1974, and for the common origin of both of these subspecies from Bi¬ 1986, pers. comm., 1992; McDonald et al., 1991). Thirty years son antiquus .McDonald relied upon the dense and continuous ago ,Sher cautioned against the assignment of bison to a given factual record documenting both the spatial, temporal, and geological age based upon the size and shape of horn cores, morphologica lcontinuity among midlatitude North American noting that this practice “is risky to say the least” (Sher, bison throughout the Holocene and the absence of any such 1974:207). spatia land morphologica lcontinuity among bison in ,or those Our position is that the weight of factual evidence supports coming from, Beringia. Quite simply, the vast body of factual the view that Bison bison athabascae originated in North evidence stil ldemonstrates tha tBison bison athabascae is mor¬ America from the same lineage that gave rise to Bison bison bi¬ phologically ,temporally ,and spatially nearer to Bison bison bi¬ son. McDonald (1981:260-261) was of the opinion that Bison son than to the late Pleistocene bison of eastern Siberia .Opin¬ bison athabascae could have attained its morphologica lchar¬ ions to the contrary are based upon antiquated concepts of acteristics by either “(1) suspended equilibrium ,in which case either (1) progressive diminution of body size or (2) periodic B. b. athabascae retained large body size as it evolved charac¬ radiation of new taxa into North America from Siberia ,or both ters from B. antiquus adapted to the new r-type forest/wood¬ (e.g., Allen, 1876; Gromova, 1935; Schultz and Frankforter, land selection regime ;or (2) more recent adaptive differentia¬ 1946; Skinner and Kaisen, 1947; Guthrie, 1970; Wilson, 1975, tion, in which case B. b. athabascae has increased in body size 1996 H; arington 1,986). from B. b. bison _Elements of both alternatives could have Van Zyll de Jong (1986) argued the case for a Beringian ori¬ operated in the origin and continuation of B .b .athabascae ,but gin for Bison bison athabascae and supported his thesis with a the second has probably been more important over time....” well-preserved skull (PIN 835-624-39) from the Bolshaya Considering the evolutionary trends in the size, shape, and Chukochya River in Siberia .He considered this skul lto be rep¬ proportions o fskeleta lelements ,as documented for bison dur¬ resentative o f“athabascae-\\V.Q bison in eastern Siberia a tthe ing the Holocene, one is left with the realization that Bison bi¬ close of the last glaciation.” “The similarity of this bison to son athabascae may be considered the most “advanced” or North American occidentalis and to athabascae is strong evi¬ “furthes tevolved” form o fHolocene bison in severa limportant dence o fthe former existence o fan ancestra lBeringian popula¬ respects. When the skull and limb morphology typical of the tion” (van Zyll de Jong, 1986:53). Van Zyll de Jong did not ex¬ ancestra lBison antiquus antiquus is compared with that of the amine the specimen personally ,and he provided no absolute derivative taxa .Bison bison bison and Bison bison athabascae, age for the specimen or the population it represented .Nor did the integrated skeleton of Bison bison athabascae is less like he figure the specimen ,describe any diagnostic characteristics that of Bison antiquus antiquus than is Bison bison bison .Spe¬ that would allow it to be accepted unequivocally as resembling cifically, in Bison bison athabascae, the skull is smaller, the Bison bison athabascae or Bison antiquus occidentalis more upper limb bones are relatively more robust ,and the metapodi- nearly than a Eurasian form of Bison, or provide measure¬ als are relatively more gracile than in Bison bison bison. Rela¬ ments .To diagnose this specimen ,he relied upon linear mea¬ tive to head size. Bison bison athabascae is the most long- surements alone (which were sunk in his quantitative summa¬ limbed of short homed North American bison ,whereas Bison ries and never presented, in the 1986 paper, as raw data); but antiquu santiquu si sthe shortes tlimbed. measurements alone frequently are insufficient to diagnose In other respects .Bison bison athabascae is probably not as mos tbison specimens ,especially those representing taxa that distant from Bison antiquus antiquus as is Bison bison bison. obviously overlap others in size. It is quite possible to obtain The evolutionary lengthening of the front limb relative to the identica lmeasurements from two o rmore bison specimens that hind limb has progressed further in Bison bison bison than in posses ssubstantiall ydifferen tqualitative (and geneticall ycon¬ Bison bison athabascae. Also ,the forebody pelage used to ef¬ trolled) characteristics .It also is theoretically possible for con¬ fect social rank and breeding success during the mt is less ex¬ vergent evolution to produce similar phenotypes from gener¬ aggerated and seasonally pronounced in Bison bison athabas¬ ally similar ancestral stock even in the absence of recent cae than in Bison bison bison (Geist and Karsten, 1977; Lott, genetic unity .The morphological ,behavioral ,and ecological 1979) .The horns therefore perhaps have relatively greater im¬ similarities between Bison bison and the Eurasian bison or portance for establishing socia lrank or dominance in Bison bi¬ wisent. Bison bonasus, is a case in point. Although these spe¬ son athabascae than in Bison bison bison. cies differ in morphological detail, they evolved generally via Th einterpretation o fskeleta ml easurement spresented herein pathways that resulted in convergence of size ,form ,and func¬ suggests that the evolution of the skeleton of Bison bison atha¬ tion; it is obvious from the fossi lrecord that Bison bonasus had bascae has ,collectively ,proceeded further than has that of Bi¬ a larger ancestor tha tcould have passed through a stage during son bison bison, whereas the opposite is tme for evolution of which linear features of its skul lmorphology resembled those selected morphologica lfeatures and behavior of socia lsignifi- 96 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY cance. Claims that Bison bison athabascae is more or less far ual organisms also change. The skeleton of Bison bison atha¬ evolved from its ancestra lmorph should ,therefore ,indicate ex¬ bascae appears to have differed allometrically from Bison anti- actly which parameter of evolution is being considered. It is quus to a greater extent than has the skeleton of Bison bison not necessary that al lparameters evolve the same distance ,at bison ,but behaviorally and ecologically Bison bison athabas¬ the same time, or in the same direction. As environments cae might differ less from Bison antiquus than from Bison bi¬ change ,the multitude o fselective pressures tha tac ton individ¬ so bnison. Appendix Measurements (in mm) of limb bones of Kenora Bison (Measuremen tasn adbbreviation fsollo wthos geive fno brovid ivsno dne Dnriesc h1,976) 1. Atlas: GB=199.9; GL=129.5; BFcd=111.3; BFcr= anteroposterio rdiameter d, ista el piphysis=40.3 B; p=77.5. 123.8 ;H=102.1. 11. Proximal sesamoid, thoracic limb: length=26.1; trans¬ 2. Axis: LCDe=122; H=162.1; LAPa=122.6; BFcd=67.7; vers ebreadth=17.8 a;nteroposterio drepth=15.8. BPtr=132.8; BPacd=88.5; BFcr=113.6; SBV=75.3. 12. Proximal phalanx, thoracic limb: GLpe=65.3; Bp=39.7; 3. Manubrium: Dorsoventral depth=92.3; length= 104.5; SD=35.8;Bd=39.6. breadth at articular surfaces=47.2; minimum breadth= 13. Medial phalanx, thoracic limb: GLpe=43.3; Bp=38.1; 17.5. SD=31.3; Bd=35.9. 4. Sacrum: BFcr=72.0; HFcr=35.3. 14. Bony pelvis: GL=526; GBTi=279; SBI=180; GBA= 5. Scapula (right): Ld=276; HS=504; DHA=487; SLC= 268; SB=37.0; GBTc=503; LS=208.6; LA=89.6. 79.2; GLP=95.4; LG=75.0; BG=69.8; transverse breadth 15. Femur (right): GL=436; GLC=405; rotational length= of neck= 37.9; breadth, lateral border of spine to medial borde ro fneck=76.7. 383; Bp=154.1; SD=48.8; Bd=119.3; DC=56.4; antero¬ 6. Radius (left): Approximate rotational length=309; GL posterio rdiamete ro fdiaphysis=51.8. (radius)=333 ;GL (radioulna)=452 ;minimum anteropos¬ 16. Tibia (right): Rotational length = 370; GL=403; Bp= terio rdiamete ro fdiaphysis=32.0 ;minimum transverse di¬ 127.4 ;Bd= 103.4 ;SD=51.7 ;minimum anteroposterior di¬ amete ro df iaphysis=57.8. ameter of shaft=35.0; Bd=75.7; Dd=53.0. 7. Ulna (left): GL-425; DPA = 98.9; SDO=68.5; LD = 17. Calcaneum (right): GL=158.9; GB = 56.7; transverse 145.7 ;minimum transverse breadth o fdiaphysis=21.3. breadth o nf eck=23.0. 8. Radial carpal (right): GB=48.2. 18. Proximal phalanx 1, pelvic limb: GLpe=69.7; Bp=34.2; 9. Fused 2+3 carpal (left): GB=47.4. SD=31.6; Bd=35.2. 10. Metacarpal (right): GL= 190.0; SD==48.3; DD=28.2; BD 19. Proximal phalanx 2, pelvic limb: GLpe=70.3; Bp=34.7; =76.8 a; nteroposterio rdiameter p, roxima el piphysis=46.0; SD=31.4; Bd=35.6. 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"Bison antiquus from Kenora, Ontario, and Notes on the Evolution of North American Holocene Bison." Cenozoic mammals of land and sea : tributes to the career of Clayton E. Ray 93, 83–97. View This Item Online: https://www.biodiversitylibrary.org/item/266341 Permalink: https://www.biodiversitylibrary.org/partpdf/352089 Holding Institution Smithsonian Libraries Sponsored by Smithsonian Institution Copyright & Reuse Copyright Status: In copyright. Digitized with the permission of the rights holder. Rights Holder: Smithsonian Institution License: http://creativecommons.org/licenses/by-nc-sa/4.0/ Rights: http://biodiversitylibrary.org/permissions This document was created from content at the Biodiversity Heritage Library, the world's largest open access digital library for biodiversity literature and archives. Visit BHL at https://www.biodiversitylibrary.org. This file was generated 27 May 2023 at 22:33 UTC