Great Basin Naturalist
Volume 59 Number 2 Article 6
4-30-1999
An Irvingtonian species of Brachylagus (Mammalia: Lagomorpha) from Porcupine Cave, Park County, Colorado
Colleen N. Ramos Denver Museum of Natural History, Denver, Colorado and University of Colorado at Boulder
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Recommended Citation Ramos, Colleen N. (1999) "An Irvingtonian species of Brachylagus (Mammalia: Lagomorpha) from Porcupine Cave, Park County, Colorado," Great Basin Naturalist: Vol. 59 : No. 2 , Article 6. Available at: https://scholarsarchive.byu.edu/gbn/vol59/iss2/6
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AN IRVINGTONIAN SPECIES OF BRACHYIAGUS (MAMMALIA: LAGOMORPHA) FROM PORCUPINE CAVE, PARK COUNTY, COLORADO
Colleen N. Ramos]
AI3STHACT.-Brachylflgus is currently a monotypie genus of UJlcertain origins and known only fi'om Holocene and late Raneholabrean. A new species ofleporid is described from the early and middle Pleistocene (Irvin~tonian) deposits of Porcupine Cave, Park County, Colorado. Stratified deposits of the Pit and the Velvet Room, 2 localities within Porcu pine Cave, have been dated biochronologically and paleomagnetically from the middle Irvingtonian and early to middle Irvingtonian, respectively. Brachylagu8 coloradoensis, sp. nov., is characterized hy its conserved p3 enamel patterns which are intermediate between B. idahoensis and Hypolagus, and its si7.e which is slightly larger than that of B. idflhoen::,;is. This suggests a possible ancestral relationship belwecn I-Iypolagus [md Brachylag11.s.
Key words: Brachylagus, Irvingtonian, BraehyJagus idahoensis, pygmy rabbit, Leporidae, Lagomorpha, Pleistocene.
Bmchylagus idahoensis (pygmy rabbit), the Brachylagus idahoensis, being the smallest sole living species of Brachylagus, is a small leporid in North America, is easily identified rabbit that is restricted to dense stands of by its unique p3 enamel patterns and its small sagebrush (Artemisia spp.) in the Great Basin. size. Even so, few paIeontological.sites include An isolated population also survives in eastern this species in their faunal lists; thus, it has a \Vashington and is currently the subject of limited fossil record that extends only to the attention by conservationists. Because of its late Rancholabrean (Kurten and Anderson highly specific habitat needs (as reviewed by 1980). However, material from Cathedral Dobler and Dixon 1990), the species is vulner Cave, an Irvingtonian site in eastern Nevada, able to local extirpation as its natural habitat is has yielded isolated teeth easily referable to B. modified and/or destroyed by agriculture and idahoensis (C.J. Bell personal communication). cattle grazing. The species has apparently Despite its limited geologic presence, this existed in the Great Basin area for much of its genus has been hypothesized to have origi evolutionary history as only a single paleoIlto~ nated either in the Miocene and descended logical site is known from outside its current from the genus Alilepus (Hibbard 1963), or range (Walker 1987). later in the Pliocene and again derived from The genus has a long history of taxonomic Alilepus (White 1991b). These hypothetical uncertainty but is now generally considered to ancestries are based upon formation of the p3 be valid. The species now known as B. ida~ enamel pattern found in Brachylagus, which is hoensis was first described by Merriam (1891) completely distinct from other North Ameri but ascrihed to the genus Lepus. Miller (1900) can leporids. Given the limited fossil record of formally proposed the subgenus Brachylagus, Brachylagus and extensive radiation of the which Lyon (1904) elevated to full generic sta genus Hypolagu8 during the Pliocene, a rea tus, a usage maintained by Nelson (1909). Later sonahle, alternative hypothesis is that Bmchy authors placed the species in the genus of lagus arose directly from Hypolagus in the late Sylvilagus (Grinnell et al. 1930, Orr 1940, Dur Blancan or early Irvingtonian. These hypothe rant 1952). More recent morphological work ses are summarized in Figure 1 and evaluated has supported the generic status of Brachyla below. gus (Kenner 1965), with recent genetic work The species described here occurs in the (reviewed hy Chapman and Ceballos 1990) Irvingtonian deposits of Porcupine Cave, Park further suhstantiating this. County, Colorado. The cave contains several
1lknver \1"",,,mu of I\"attlml History, 200l COlOlWlo lII\"d., DcnVel; CO 80205·5'ro8, ~md University ofColorado at Boulder, Campus IkJx 334, BOllldeJ; CO 80300
151 152 GREAT BASIN NATURALIST [Volume 59
A
Hypolagus Alilepus Brachylagus
B ---~
Alilepus Brachylagus
c A Hypolagus Brachylagus
Fig. 1. Summary ofhypotheses concerning evolution ofp3 form seen in Brachylagus: A, Brachylagus descending from Alilept~s', posterior-external reentrant (PER) fi.lses with posterior internal reentrant to form separate lobes as seen in Brachylagus (summarized from Hibbard 1963); B, Brachylagus descending from AlileptlS (\vhich arose from Hypolagus) during late Pliocene or early Pleistocene (after \iVhite 1991b); and C, Brachylagus descending directly from Hypolagus, FEH eventually extends completely across \vidth of the tooth to separate the trigonid from the talonid. rooms from which fossils have been collected apparently undisturbed. These lower levels are (Barnosky and Rasmussen 1988, Wood and also yielding mammal species, such as Hypola Barnosky 1994, Anderson 1995). These rooms gus, that are older than those found anywhere and areas represent localities within Porcu else in the cave. Although other rooms with pine Cave and include the Pit, Velvet Room, fossilized remains exist and have been sam and Mark's Sink. Fossil deposits range in age pled, the 3 rooms described above have pro from early to late Irvingtonian. The Pit, a duced most fossil specimens to date and may, small room in the cave, was excavated by the collectively, span more than one million years. Carnegie Musenm (C~1) in the 1980s and was In the 1980s certain specimens from the Pit originally thought to date between approxi "vere identified as Brachylagus idahoensis by mately 0.38 my to roughly 0.8 my (Barnosky Barnosky and Rasmussen (1988). While ana and Rasmnssen 1988, Wood and Barnosky lyzing the leporid remains from the Velvet 1994). More recent analyses date the majority Room and Mark's Sink, I discovered several ofthe Pit m'ound 850,000 y.b.p. (Barnosky per teeth and mandibles similar in size to those of sonal communication). Paleomagnetic studies B. idahoensis but lacking the diagnostic p3 of the Velvet Room have fonnd that the npper enamel pattern of this species, When I reex 5 levels correlate to the present normal polar amined the Pit specimens, I found that they ity while the majority of the lower levels were too lacked the B. idahoensis form ofp3. Previ deposited during the Matuyama reversed polar ous morphometric work (Ramos in press) pro ity chron (B. Raynolds and J. Friedman per vided a large sample of extant B. idahoensis (n sonal communication) and are therefore at = 85) as well as an understanding of variation least 0.78 million years in age. Mark's Sink, an in dental characters for this species. \Vhen isolated section of the Velvet Room, contains used for comparison with the Porcupine Cave bioturbated material in its upper sections, fossils, these data demonstrated that although while its lower portions are stratified and the fossils were similar in several ways to B, I999J IHVINCTONIAN SPECIES OF BRACIlYIAGUS 153 i
HYPODIGM.-Pit: CM #66408-66409, p3; than the average B. idalwensis although some CM #66431, edentulous right maxilla; CM specimens fall within the range of individual #66432, edentulous left maxilla; CM #65486, variation for B. idahoensis (Tables 1, 2). Results left dentary with p4-m2; CM #66604, p3; CM of the 2-sample t test indicate that p3 of B. #65607, p3. Velvet Room: DMNH #28901, ida/wensis and p3 of B. co/oradoensis differ in 33250, 33255, p3. Mark's Sink: DMNH #28902, length (FSl19, = 4.5, P < 0.0001) and width left humerus; DM H #33257, left dentary (FS1 ,19 = 2.66, P < 0.001). These results, and with p3-p4; DMNH #33260, three isolated the differences between enamel patterns, indi p3s; DMNH #33272, four isolated p3s; cate that the 2 samples are not drawn from the DMI\'H #33271, left dentary with p3-m2; same population and support the designation DMNH #33270, left dentary with p3-ml. of B. coloradoensis as a species separate from TYPE LOCALITY AND AGE.-Middle Pleis B. idahoensis. The PER of B. coloradoensis does tocene, early to middle Irvingtonian, DMNH not extend completely across the tooth width Locality No. 1349, Porcupine Cave, 2900 m, as in B. idahoensis. No complete skulls of B. Park Co., Colorado ( 'WI/4 of SW1/4 Sec. 23, cokn'adoensis are known from this site; how Tl5S, R76W, Lat. 38°43'45"N, Long. 105° ever, portions of dentaries and edentulous 51'41"W, Gribbles Park 7.5' Quad). maxillae are present Disarticulated postcra ETYMOLOGY.-Named for the locality and nial material has been tentatively assigned to follOWing the precedent set in this genus. At this species due to the unusually small size present, this species is known only from a sin which allows no other confident identification gle site in Colorado. within the Leporidae. Several uncataloged DIAGNOSJS.-The enamel pattern of the p3 humeri, ulnae, and podial remains have thus is distinct from that of B. idahoensis in having been identified to B. coloradoensis and are a posteroexternal reentrant (PER) that extends similar in size to corresponding elements ofB. between 1/3 and 2/3 the width of the tooth idahoensis. (Fig. 3). The posterointernal reentrant (PIR), when present, is only a slight indentation. DJSCUSSIOI\ Dimensions of postcranial material strongly resemble those of modern B. idahoensis but Brachylagus coloradoe'11sis, sp. nov., shows are slightly more robust some affinity to the extinct genus Hypolagus COMPARISONS.-B1"achylagus coloradoensis that radiated profusely in the Pliocene (Fig. 3). may be distinguished from Hypolagus species Although some p3s ofB. coloradoensis display by its smaller size. Also, the PER ofB. colora patterns with clear PIRs as seen in Alilepus, doensis p3 extends farther across the tooth none of the upper molars found and ascribed width, generally greater than 1/2 the distance to E. coloradoensis display the characteristic (Fig. 3). Although B. idahoensis is of similar enamel lake found in Alilepus. This lake is also size, B. coloradoensis, sp. nov" is generally absent in Hypolagus. Hypolagus is easily dis more robust. In comparing the size of p3, B. tinguished by its simpler p3 enamel patterns coloradoensis is approximately 12--15% larger and larger size.
A B c D E F
Fig. J. Comparison ofpJ enamel patterns for Hypo[agUJi, B. cOWradoensis, and B. idahoensis. A is Hypolagw (redrawn from White 1987:439), B-E (DMNH # unknown, DMNH #33257, DMNH #33255, and DMNH #33260) represent the range of variation seen in specimens of B. coloradoensis. B predominates slightly in earlier sediments whereas C becomes somewhat more plentiful in younger sediments at Porcupine Cave. F is that of B. idahoensis (UCM #5840). Nole also that some specimens ofB. coloradoensis have a slight posterior-internal reentrant (FIR). 1999J IRVINGTONIAN SPECIES OF BRACHl'LAGUS 155
TABLE 1. Dimensions and descriptive statistics of dentary and lower dentition of Brachylagus colo1"udf}{:nsis and B. idahoellsis (in par~nt:hescs). Valuc.... for B. idahoensis are for left measurements only.
M~urement x ±s Range , Alveolar I.cnglh oftooth row 10.48 (n/a) n/a 2 (9.1 ± 0.34) (8.24-10.26) (85)
Length of diastema 10.9 (,,/a) n/a 1 (9.0 ± 0.43) (8.4f~l1.92) (85)
Depth ofdenbtry at p4 8.86 ± 0.30 7.74-9.56 4 (7.54 ± 0.42) (6.60-8.26) (85)
Length ofp3 2.02 + 0.28 1.74-2.56 19 (1.78 ± O.Il) (1.62-2.28) (85)
W;d,h ofp3 1.80 ± 0.22 J.40-2.12 19 (1.6 ± O.ll) (1.42-2.18) (85)
Width ofp4 2.34 ± 0.12 2.12-2.54 5 (1.96 ± 0.1) (1.68-2.99) (85)
Width of ml 2.35 + 0.12 2.24-2.46 4 (1.98 + 0.1) (1.78-2.26) (85)
Width of m2 2.22 ±0.16 2.08-2.40 3 (1.94 ± 0.1) (1.74-2.20) (85)
Width of013 1.70 (ofa) nfa 2 (1.02 ± 0.(9) (0.82-1.30) (85)
TABLE 2. Dimensions and descriptive statistics of maxilla and upper dentition of Brachylagus coluradoensis and B. idahoensi~' (in parentheses). Values for B. idalwensis are for left measurements only. Measurement x ±s Range n Alveolar I~ngth of tooth row 10.08 (ofa) nfa 2 (9.13 ±0.35) (8.32-10.1) (&5)
Depth of zygomatic proce:o;s 3.49 (,,/a) n/. 2 (3.49 ± 0.31) (2.88-4.24) (RS)
Width ofl'2 2.09 (ofa) n/a 2 (1.73±0.14) (1.52-2.36) (85)
\iVidth of P3 3.19 (n/a) n/a 2 (2.96 ± 0.2) (2.58-3.38) (85)
'Width ofP4 3.19 (n/a) n/a 2 (2.98±0.17) (2.52-3.64) (&5)
Width orMl 3.05 (n/a) n/a 2 (2.87± 0.18) (2.3fhl.26) (85)
Width of:M2 2.6 (o/a) n/o 2 (2.6 ± 0.15) (2.32-2.96) (85)
Width ofM3 1.15 (o/a) n/a 2 (1.07± 0.18) (0.6--1.84) (85) 156 CHEAT BASIN NATURALIST [Volume 59
The lower third premolar (P3) is considered Of the hypotheses concerning the ancestral somewhat diagnostic among leporids and has stock of this genus (Fig. 1), it appears just as been used in numerous species descriptions, likely that Brachylaglls arose directly from especially ofextinct leporids (White 1984, 1987, Hypolagus as Ii-om Alilepus. Although B. col 1991a, 1991b, Hibbard 1963). In B. colora oradoensis exhibits p3s ofboth Hypolaglls and doensis there is some variation in this charac~ Alilepus form, there are no enamel lakes pre tor which, based upon the limited number of sent in any of the upper molars examined. specimens available, seems to be present re Enamel lakcs in upper cheek teeth are absent ganlless of geologic age. In short, no particu in Ilypolagus, but present in P3 of AlileptlS, lar evolutionary trend is seen in this character. and may represent plesiornorpbie characters Although the simplest form is slightly more retained from Paleolaginae from which Archae prevalent in older strata and the more derived olaginae arose. Also, the time trames hypothe fonn seems more prevalent in later strata, sized by Hibbard (1963) and White (1991a) both extremes of the continuum are present in are unsubstantiated by the fossil record as oldest and youngest strata. The range of varia there is no evidence that Brachylagus existed tion in the p3 enamel pattern of B. colora earlier than the very latest Blancan to earliest doeHsis is depicted in Figure 3. The P2 con Irvingtonian. tains a single anterior reentrant as in B. -ida Genera such as Lepus and Sylvilagus are hoemis and Hypol"gus. not considered to have originated directly The size of thc Mark's Sink B. coloradoen from f1ypowgus. One or more potential inter sis, which is presumably the oldest in the cave, mediates have been hypothesized over the is somewhat larger than that of either the Vel years, including Alilepus, Sel'engetilag"s (Plio pentalngus), and others (Hihbard 1963, White vet Room or Pit specimens of B. coloradoel1sis 199Ib), and fossil evidence does not dispute (Fig. 4) while retaining more eonscrvative p3 these hypotheses. However, B. coloradocnsis enamel patterns. This is an intriguing trend appears to indicate a more direct transition and may indicate a time frame for the clado from Hypolagus to Brachylagus. 11ms, the living genic event that gave risc to the genus B1"achy pygmy rabbit may he the only direct descen lagus. Unfo,-tunately, the strata from which dent of an extinct genus and may be more dis these earliest specimens come are bracketed tantly related to Syluilagus t1,an previously only with upper and lower time limits and thought. Although Halanych and Robinson there is little refinement in the age estimates (1997) found some molecular evidence placing of the levels themselves. Therefore, a more BrachyuJgus as sister taxon to SylvikJgllS based accurate picture of the rate at which the varia upon mitochondrial 12S rDNA data, other tion narrowed is not possible at tl1i5 time. aspects of their analysis did not support this However, the wide range of size variation strongly. 1 find that the high degree of mor noted for specimens of B. colomdoensis in the phological differences between Brachylagus early Irvingtonian of Porcupine Cave (Fig. 4) and SylvilagllS, as well as the paleontolcgical may indicate that the cladogenetic event evidence described above, present a strong occurred shortly (in geologic terms) before thc case against a close relationship between them Porcupine Cave record. At least 2 species of (see also Kenner 1965, Creen and Flindcrs Hypolagus, ,L"i yet unidentified, are present in 1980). the older sections of Mark's Sink with the B. Given the high degree ofhabitat specificity colonuloensis material. One of these species is exhihited by the extant species of Brach!llagfls, very small, though still somewhat larger than it would be interesting to cletennine whether the lar~est specimens of B. coloradoensis. The its extinct species was also highly habitat spe relationship between B. coun-adnensis and this cific. Unfortunately, paleoecological evidence small Hypolagus is undear at present. A more at Porcupine Cave is scanty and restricted detailed phylogenetic analysis utilizing several mostly to hmnal remains. Such pustcranial re characters is currently in progress and should mains as have been assigned to B. culomdoen provide stronger evidence concerning the ori sis resemhle B. iduhoensis in almost all details. gins of Brachylagus and its rdationships to This indicates a similar degree of locomotor extinct and extant genera (C.N. Ramos in adaptation as seen in the extant spt."'Cies. Brachy preparation). lagtlS coloradoe1lSis was probably also closely 1999J IRVINGTONIAN SPECIES OF BRACHYLAGUS 157
3 • 2.5 • • ••• -e 2 -e • • • • • • • ...C- • • •• o 1.5 i j 1
0.5
o o 5 10 15 20 Approximate chronological order (1-6 from Mark's Sink, 7-18 from Velvet Room)
Fig. 4. Change in p3lengt11 over time. Size data plotted over time for the different rooms in Porcupine Cave. Length of p3 has been used to illustrate the general trend, that of wide size range in older sediments, then loss of upper size limits and closer approximation to size range of B. idahoensis. Line indicates average p3length for B. idahoensis.
allied to brush and dense cover and avoided Although complete species associations can open areas, although the plant biota compris not be reconstructed at this time for Porcu ing its habitat is impossible to ascertain at pine Cave, it is notable that this new species this time. However, Barnosky and Rasmussen of leporid was associated with extinctleporids (1988) reported the consistent concurrence of such as AztlonologWi and HypologWi in the old sage vole (Lagums curtattls) "ith B. colora est sections of Porcupine Cave, yet found with doensis (reported as Sylvilagus ida/lOensis). extant species such as LepWi townsendii and This arvicoline rodent is generally found in SylvilogLls audubonii in younger sediments at stands ofArtemisia (Armstrong 1972) and cur tltis site (Ramos 1998). These associations do rently has a distribution overlapping tbat of B. not necessarily indicate sympatry but do indi ·idahoens1s, Both mammal species are absent cate a temporal coexistence. Sympatry is rare from the Wyoming Basin, as is B. idahoe""is, among species of rabbits and hares, and the despite the apparent presence of suitable habi remains at Porcupine Cave are the result of tat (Walker 1987). The presence of an Irving predation by raptors and mammals, snme of tonian species of Brachylagu8 on the eastern which have large ranges. Prey remains are side of the Cnntinental Divide and the persis thus sampled from a pntentially wide radius tence of extant Brachylogus in the Great Basin and do not represent close species associations. may indicate tbat the·2 populations became It is still intriguing to lind that this little lep isolated on opposite sides of tbe Rocky Moun orid was able to survive while nther leporid tains after dispersing through tbe Wyoming species, even genera, were going extinct. Basin. Wbere the genus originated is unclear; Unfortunately, the fossil record at Porcupine and although I nnd the presence of a small Cave truncates several hundred thousand HypologWi associated witb B. coloradoensis in years ago and we cannot l'11ow, at this time, the older strata of M3J·k's Sink to be sugges how long B. colomdoensis survived in associa· tive, it certainly is not conclusive. lion with modern fauna. 158 GREAT BASIN NATURALIST [Volume 59
Porcupine Cave is continuing to be exca BENSLEY, B.A. 1926. Practical anatomy of the rabbit. P. vated and specimens of this new species are Blakiston's Son & Co., Philadelphia. 298 pp. CHAPMAN, J. A, AND G. CEBALLOS. 1990. The cottontails. being added to the collections at DMNH. In Pages 95-110 in J.A. Chapman and J.E.c. Flux, edi addition, temporal limits of localities within tors, Rabbits, hares, and pikas: status survey and the cave are being clarified, which will allow conservation action plan. International Unian for more refined analysis ofmorphological change, Conservation of Nature and Natural Resources speciation rate, and faunal associations. As fur (IUCN), Gland, Switzerland. ther analysis of this important Irvingtonian DOBLER, RG., AND K.R. DIXON. 1990. The pygmy rabbit. Pages 111-115 in J.A. Chapman and J.E.G. Flux, site continues, our understanding ofthis extinct editors, Rabbits, hares, and pikas: status survey and species will continue to grow, hopefully pro COnservation action plan. International Union for viding new insights into the evolutionary history Conservation of Nature and Natural Resources ofthe single living species of Brachylagus. (IUCN), Gland, Switzerland. DURRANT, S.D. 1952. Mammals of Utah, taxonomy and distribution. University ofKansas Publications, Muse ACKNOWLEDGMENTS um of Natural History, Lawrence 6:1-549. FINDLEY, J.S., A.H. HARRIS, D.E. WILSON, AND C. JONES. David Daitch of the University of Colorado 1975. Mammals of New Mexico. University of New at Boulder provided the excellent drawings for Mexico Press, Albuquerque. 360 pp. the figures. I also thank the following individ GREEN, J.S., AND J.T FLINDERS. 1980. Brachylagus ida hoensis. Mammalian Species 125:1-4. uals for their help with this work: Cheri A. GRINNELL, J., J. DIXON, AND J.M. LINSDALE. 1930. Verte Jones and the Denver Museum of Natural brate natural history of a section ofnorthern Califor History; Dave Armstrong of the University of nia through the Lassen Peak region. University of Colorado at Boulder; Elaine Anderson, Russ California Publications in Zoology 35:1-594. Graham, Kathy Honda, and Kirk Johnson of HALANYCH, KM., AND TJ. ROBINSON. 1997. Phylogenetic relationships ofcottontails (Sylvilagus, Lagomorpha): the Denver Museum of Natural History; Jane congruence of 12S rDNA and cytogenetic data. Mol Bock, Herbert Covert, Carol Wessman, Greg ecular Phylogenetics and Evolution 7:294-302. Carey, and Jill Skarstad of the University of HIBBARD, C.W. 1963. The origin ofthe P3 pattern of Sylvi Colorado at Boulder; Christopher J. Bell, lagus, Caprolagus, Oryctolagus, and Lepus. Joumal Anthony D. Barnosky, Julio Friedman, C.A. of Mammalogy 44: 1-15. KENNER, G.H. 1965. Comparative osteology of rabbits of Repenning, Bob Raynolds, Don and Jerry the genera Brachylagus Miller and Sylvilagus Gray. Rasmussen, Lou Taylor, and all the Porcupine Unpublished master's thesis, University of Utah, Cave volunteers. Thanks to the following insti Salt Lake City. 125 pp. tutions for the generous use of specimens: KURTEN, B., AND E. ANDERSON. 1980. Pleistocene mam Denver Museum of Natural History, Carnegie mals of North America. Columbia University Press, New York 443 pp. Museum of Natural History, Idaho Museum of LYo~, M.W 1904. Classification of the hares and their Natural History, Field Museum, National Mu allies. Smithsonian Miscellaneous Collections seum of Natural History, University of Colo 45(1456i321--447. rado MuseUffi, and Cowan Vertebrate Museum. MERRIAM, C.H. 1891. :Mammals of Idaho. North Ameri William Akerston, Anthony D. Barnosky, and can Fauna 5:75-78. MILLER, G.S. 1900. A new subgenus for Lepus idalwensis. John White acted as reviewers and their com Proceedings of the Biological Society, Washington ments helped improve the original draft. 13,157. NELSON, E.W 1909. The rabbits of North America. North LITERATURE CITED American Fauna 29:1-314. ORR, RT. 1940. The rabbits of California. Occasional A~DERSO~, E. 1995. Preliminary report on the carnivores Papers of the California Academy of Sciences of Porcupine Cave, Park County, Colorado. Pages m1-227. 259-282 in K. Stewart and K. SeymoUl; editors, RAMOS, C.N. 1998. Evolution and biogeography of North Palaeoecology and palaeoenvironments of Late Ceno American Leporidae. Doctoral dissertation, Univer~ zoic mammals: tributes to the career of C.S. (Rufus) sity ofColorado at Boulder. 192 pp. Churcher. University ofTaronto Press, Toronto. . IN PRESS. Morphometric variation among leporids ARMSTRONG, D.M. 1972. Distribution of mammals in Col --("Mammalia: Lagomorpha) of the Interior \i\/est. Pro orado. Monograph, University of Kansas Museum of ceedings ofthe Denver Museum of Natural History. Natural History 3:1-415. WALKER, D.N. 1987. Late Pleistocene/Holocene environ BARNOSKY, A.D., AND D.L. RASMUSSEN. 1988. Middle Pleis mental changes in \Vyoming: the mammalian record. tocene arvicoline rodents and environmental change Pages 334-392 in R\'Z Graham, H.A. Semken, and at 2900-meters elevation at Porcupine Cave, South M.A. Graham, editors, Late Quaternary mammalian Park, Colorado. Annals of the Carnegie Museum 57: biogeography and environments of the Great Plains 267-292. and prairies. Illinois State Museum, Springfield. 1999] IRVINGTONIAN SPECIES Of BRACHYLAGUS 159
WHITE, J.A. 1984. Late Cenozoic Leporidae (Mammalia, _---". 1991b. North American Leporinae (Mammalia: Lagomorpha) from the Anza- Borrego Desert, South Lagomorpha) from late Miocene (Clarendonian) to ern Califomia. Special Publication, Carnegie Museum latest Pliocene (Blancan). Journal ofVertebrate Pale· ofN"atural History 9:41-57. ontology 11:67-89. __,;" 1987. The ArchaeoLaginae (~1ammaJia; Lagomor WOOD, D.L., AND A.D. BARNOSKY. 1994. Middle Pleisto pha) of North America., excluding Archaeolagus and cene climate change in the Colorado Rocky .\1000 Panola:c. Journal of Vertebrate Paleontology 7:425 taiDS indicated by fossil mammals from Porcupine 450, Cave. Quaternary Research 41:366-375. . 1991a, A new 5ylvilagus (Mammalia: Lagomor --p';ba) from the Blancan (Pliocene) and Irvingtonian Received 21 NOfJ6TI1h.er 1997 (Pleistocene) of Florida. Journal of Vertebrate Pale Accepted 22 June 1998 ontology 11:243-246.