White (1988) - The Archaeolagine of NA

The Archaeolaginae (Mammalia, Lagomorpha) of North America, Excluding Archaeolagus
and Panolax
Author(s): John A. White
Source: Journal of Vertebrate Paleontology, Vol. 7, No. 4 (Jan. 22, 1988), pp. 425-450
Published by: Taylor & Francis, Ltd. on behalf of The Society of Vertebrate Paleontology
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Journal of Vertebrate Paleontology 7(4):425-450, December 1987
THE ARCHAEOLAGINAE (MAMMALIA, LAGOMORPHA) OF NORTH AMERICA,

EXCLUDING ARCHAEOLA GUS AND PANOLAX
JOHN A. WHITE
Florida State Museum, University of Florida, Gainesville, Florida 32611
ABSTRACT-The Archaeolaginae of North America, excluding Archaeolagus and Panolax, include

the following genera and species: Hypolagus vetus, H. parviplicatus, H. gidleyi n. sp., H. fontinalis, H.
tedfordi n. sp., H. furlongi, H. edensis, H. oregonensis, H. ringoldensis, H. regalis, H. voorhiesi n. sp.,
H. arizonensis, Lepoides lepoides n. gen. and sp., Pewelagus dawsonae, and ?Pewelagus mexicanus.
?Hypolagus apachensis is referred to the Leporinae. The genus Notolagus is referred to the Leporinae
and Paranotolagus is tentatively referred to the Leporinae. Lepoides lepoides n. gen. and sp. is tentatively
thought to be a jack rabbit ecomorph, while some species of Hypolagus possibly cottontail ecomorphs.
The Archaeolaginae of North America became extinct by the end of the Blancan land mammal age
(Pliocene).
INTRODUCTION
above valid taxa, Hypolagus apachensis (in part), No-
The subfamily Archaeolaginae of Northtolagus Americavelox, in-
N. lepusculus, and Paranotolagus com-
clude five leporid genera: Archaeolagus, Panolax, plicatus, areHy-
referred to the Leporinae in this paper.
polagus, Lepoides n. gen., and Pewelagus.The purpose of this research is to describe newly
Since Dawson's (1958) excellent review of available specimens, to bring up to date the systematics
the later
Tertiary Leporidae of North America, a large of the Archaeolaginae
number (excluding Archaeolagus and
of specimens ofleporids have been obtained Panolax), to document the stratigraphic range of each
by several
institutions from new and previously known taxon, localities
and to provide phylogenetic interpretations for
in North America. The following are thethe subfamily.
species de-
scribed and referred to the Archaeolaginae since the
publication of Dawson's monograph: Downey (1962) METHODS AND MATERIALS
described Hypolagus arizonensis from the Blancan of
Arizona; Gustafson (1978) described H. ringoldensis All specimens examined in this study are li
from the Blancan of Washington; Miller and Carranza- each species in the section on Systematic Pal
Castaiieda (1982) described H. mexicanus and Para- In most North American localities where leporids
notolagus complicatus from the Blancan of Guanajua- have been collected, the elements most useful for taxo-
to, Mexico; Alvarez (1963) described Archaeolagus nomic diagnoses are the P3 and P2 if the latter are
sonoranus from Tertiary sediments in Sonora, Mexico; associated with the former. Although P4-M3 and P3-
Barnosky (1986) described A. emeraldensis from the M3 may some day prove to be diagnostic, they have
Hemingfordian of Wyoming; Stevens et al. (1969) de- not been adequately described and, where described,
scribed A. buangulus from the early Miocene of Texas; are usually not diagnostic. With a few exceptions, most
and White (1984) described Pewelagus dawsonae from systematic work on Neogene leporids has been based
the Blancan of California. on mandibles and especially the morphology of the P3.
To date, the valid taxa included in the subfamily Thirty-two per cent (22 out of 68) of the localities from
Archaeolaginae are: Archaeolagus ennisianus (Cope), which archaeolagine specimens were examined for this
A. acaricolis Dawson, A. premigenius (Matthew), A. research yielded only isolated P3's.
megacephalus (Matthew), A. sonoranus Alvarez, A. Camera lucida drawings were made of the occlusal
buangulus Stevens, Stevens, and Dawson, A. emeral- enamel pattern of P3 for each of 466 P3's of archaeo-
densis Barnosky, Panolax sanctaefidei Cope, Hypola- lagines using the same Wild microscope and camera
gus parviplicatus Dawson, H. apachensis Gazin, H. lucida attachment. It was arbitrarily decided to place
fontinalis Dawson, H. limnetus Gazin, H. vetus (Kel- all drawings on 3 x 5 inch index cards. Small-sized
logg), H. regalis Hibbard, H. edensis Frick, H. orego- specimens, such as those in Hypolagus edensis, were
nensis Shotwell, H. arizonensis Downey, H. mexicanus drawn at a magnification of x 26.8, and those of larger-
Miller and Carranza-Castafieda, H. ringoldensis Gus- sized specimens, such as H. regalis, were drawn at a
tafson, Pewelagus dawsonae White, Notolagus velox magnification of x 13.4.
Wilson, N. lepusculus (Hibbard), and Paranotolagus Lines were marked on each of the drawings and mea-
complicatus Miller and Carranza-Castafieda. Of the surements were made between these lines (Fig. 1). The
© 1987 by the Society of Vertebrate Paleontology 425
measurements were: anteroposterior length, width,
depth of anteroexternal reentrant, and depth of pos-
teroexternal reentrant. The margin of error is ±0.2 mm
G A
for specimens drawn at x 13.4 magnification, and ±0.12
mm for specimens drawn at x 26.8 magnification. Cal-
ipers calibrated to 0.1 mm were used to measure most
specimens in addition to the above method, and to
measure skull parts.
The deflection of the posteroexternal reentrant of P3
was measured in degrees on each one of the camera
lucida drawings. This was accomplished by extending
a line from the center of TN (see abbreviations) at the
point immediately after it has completed its mediad
turn, to the medial-most extension of TH (Fig. 2).
The use of camera lucida drawings of occlusal pat-
terns in morphological analyses of small teeth has the
advantage of enabling one to compare all the drawings
at the same time rather than examining one or a few -D
teeth under the microscope at the same time.
Camera lucida drawings were repeated on specimens - -E
that had been figured previously. Some small details a
were slightly different on comparable drawings of the

same specimen but not sufficient to invalidate the re- FIGURE 1. Diagram of the occlusal surface of an archaeo-
liability of these drawings. lagine P3 indicating how measurements were made. The mea-
Very little is known about age variation in the skull surements are: anteroposterior length F, G; width A-E; depth
morphology of lagomorphs. It is often difficult to de- of posteroexternal reentrant (PER) B-E; depth of anteroex-
cide if a small-sized specimen is a young individual of ternal reentrant (AER) C-E; line of orientation D.
a large-sized species or the adult of a small-sized species.
In some specimens of P3, if the anteroposterior length
is measured near the occlusal surface and again at the
base of the tooth crown, the former measurement in across occipital condyles; least postorbital constriction
young specimens is significantly smaller than the latter. (constriction posterior to supraorbital ridges); depth
There are no known data to suggest the ontogenetic anterior end zygomatic arch (from ventral side of an-
age when P3 becomes parallel-sided. terior end of zygomatic arch to anteroventral surface
In some unworn specimens of P3 thickness of the of orbit); width of cranium at anterior end of zygomatic
enamel on the periphery of the tooth seems unusually arches (across anterior prominences at anterior end of
great; however, enamel thickness, even in adult spec- zygomatic arches); length of I2_p2 diastema at alveoli;
imens, is variable and does not seem to be significant alveolar length of P2_M3; anteroposterior length of P3;
for age-determination or for systematic purposes. width of P3; length of I-P3 diastema at alveoli; man-
For each sample of five or more specimens, the mean, dibular depth below M,; alveolar length of P3-M3.
standard deviation, and observed ranges was calculat- Of the latter measurements, the length of I-P3 dia-
ed. When the word "significant" is used in statistical stema was plotted against the anteroposterior length
comparisons, it indicates a difference at the 95 per cent of P3 using 421 skulls of extant Sylvilagus and Lepus.
level or greater. When possible, twenty-five skull mea- The resulting dots, representing each genus, were then
surements were made as follows: greatest length of cra- circumscribed and the dots erased (Fig. 9). Comparable
nium (from anterior surface of front incisor to inion); dots were then superimposed using comparable mea-
condylopremaxillary length of cranium (from anterior surements of extinct archaeolagines.
alveolus of front incisor to posterior edge of occipital The local faunas mentioned in this paper are listed
condyles); length of palatine foramen; width of palatine alphabetically by states or provinces in Table 4.
foramen (at widest part of foramen near palate); palatal
length; from posterior edge of palate to suture at an- Abbreviations
terior end of basioccipital; width of choana; length of Institutional- AMNH(FAM) = American Museum
basioccipital (from ventral notch of foramen magnum of Natural History; BUNM = Museum of Southwest
to anterior end of basioccipital); anteroposterior di- Biology, University of New Mexico; FHSM = Stern-
ameter of bulla; width of tympanic bulla (from the berg Memorial Museum, Fort Hays Kansas State Uni-
most lateral extent of suture between bulla and basi- versity; UF = Florida State Museum, University of
occipital to most lateral extension of bulla); distance Florida; IGCU = Instituto de Geologia, Universidad
between bullae (from most medial suture between Aut6noma bulla de Mexico; IMNH = Idaho Museum of
and basioccipital to same point on other side); width Natural History; IVCM = Imperial Valley College Mu-
426 JVP 7(4), December 1987
Hypolagus vetus (57) __
H. porv/plicatus(l)
H. g/d/ey/ (3 6)
I H. regolis(35)
H. fonttin olis ( 17)
H. tedfordi(38)
H. fur/ongi (29) _ _ _ _ _ 9
H. voorhiesi(32)
H. e densis( 3 4) ___
H. oregonensis(20)
H. ringoldensis (59)
H orizonensis(2)
Lepoides /epoides(21)
Pewe/ogus dowsonoe(16)
P. mexiconus (2)
PH. opachensis ( 15)
S......
200 150.. .I100
.........I........
50 00 5" 100
deflection of PER
15
- anterior posterior-
FIGURE 2. Dice-Leraas diagrams of deflections in degrees of the posteroexternal reentrant of P3 from a line perp
to the line of orientation (see Fig. 1) in species of Hypolagus, Lepoides n. gen. and Pewelagus. For abbreviations
seum; KU = Museum of Natural History, University Geological Survey, Branch of Stratigraphy and Pa-
of Kansas; FMNH = Field Museum of Natural His- leontology, Menlo Park, California; USNM = United
tory; LACM = Los Angeles County Museum of Nat- States National Museum of Natural History;
ural History; LACM(CIT) = specimens from the Cal- UWBM = Burke Museum, University of Washington
ifornia Institute of Technology, now in LACM UWMG = University of Wyoming, Museum of Ge-
collections; MUBD = Biology Department, Midwest- ology; WTSU = Geology Department, West Texas State
ern University; SDSM = Museum of Geology, South University.
Dakota School of Mines and Technology; UALP = Dental Morphology (Fig. 3)-P3: AER = anteroex-
University of Arizona Laboratory of Paleontology; ternal reentrant; AIR = anterointernal reentrant; AR =
UCMP = Museum of Paleontology, University of Cal- anterior reentrant; PER = posteroexternal reentrant;
ifornia, Berkeley; UMMP = Museum of Paleontology, PIR = posterointernal reentrant; TH = thick enamel
University of Michigan; UNSM = University of Ne- on anterior edge of PER; TN = thin enamel on pos-
braska State Museum; OU = University of Oregon terior edge of PER. P2: EAR = external anterior reen-
State Museum of Anthropology; TTU = The Museum, trant; IAR = internal anterior reentrant; MAR = main
Texas Tech University; USGS-M = United States anterior reentrant.
JVP 7(4), December 1987 427
IAR
MAR
EAR
AR TH TN
anterior AER PER

labial
FIGURE 3. Diagrams of the occlusal surfaces of P2 and P3 with labels indicating tooth structures. The abbreviations are as
follows: EAR = external anterior reentrant; MAR = main anterior reentrant; IAR = internal anterior reentrant; AR = anterior
reentrant; AIR = anterointernal reentrant; PIR = posterointernal reentrant; PER = posteroextemal reentrant; AER = antero-
external reentrant; TH = thick enamel in PER; and TN = thin enamel in PER.
SYSTEMATIC PALEONTOLOGY zygomatic root more anterior to cheek teeth in the

Palaeolaginae (Dawson, 1958, in part).
Family LEPORIDAE Gray, 1821
From the Leporinae the Archaeolaginae can be dis-
Subfamily ARCHAEOLAGINAE Dice, 1929
tinguished by the absence on P3 of AIR and PIR in
adults. leporids
Emended Diagnosis--Small to large-sized
with P3 devoid of lingual reentrants (AIR All and archaeolagines
PIR) in lack internal reentrants except in
adult specimens; anteroexternal reentrantArchaeolagus (AER) slight and in Hypolagusfontinalis in which there
to deep, cement-filled or not, extendingisfrom a PIR that does42
1 to not extend to the base of the crown
per cent of occlusal width; posteroexternal in reentrant
P3. This concept of the archaeolagines agrees with
(PER) extending from 36 to 68 per cent across occlusal
that of Dice (1929) and sets this subfamily apart from
surface, and filled with cement. the more advanced leporines in which AIR and/or PIR
Remarks--Five genera of the Archaeolaginae are presentand or modified. Although AIR is absent and
their temporal ranges for North America PIR are occurs in some palaeolagines, the latter, as men-
as follows:
Archaeolagus, Arikareean to Hemphillian (late Oli- tioned above, have primitive traits that distinguish
gocene to late Miocene); Hypolagus, Barstovian to them from the archaeolagines.
Blancan (mid Miocene to late Pliocene); Pewelagus, The transfer of Notolagus to the Leporinae from the
Blancan (middle to late Pliocene); Panolax, Claren- Archaeolaginae seems reasonable to the writer because
donian and/or Hemphillian (middle to late Miocene); P3 in this genus retains AIR. Wilson (1937:101) allied
and Lepoides n. gen., Hemphillian to Blancan (Late Notolagus to Hypolagus and wrote: "Presence, in No-
Miocene to middle Pliocene). tolagus of a short posteroexternal inflection [PER] and
The Archaeolaginae can be distinguished from the absence of a corresponding inflection from the postero-
Palaeolaginae by the following characters: cheek teeth internal side of the tooth ally this genus with Hypo-
hypsodont and ever-growing; upper molariform pre- lagus.'" Dawson (1958) concurred with Wilson and at-
molars usually more molariform; two external reen- tributed the presence of AIR in Notolagus to, possibly,
trants on P3, anterior one shallower than posterior one the deepening of a variant AIR in Hypolagus apach-
and filled with cement; anterointernal (AIR) and pos- ensis.
terointernal (PIR) reentrants missing except in juve- Hibbard (1963) wrote that: "The posterior [PIR] and
niles; basicranial portion of the basicranial-basifacial anterior [AIR] internal reentrant angles, or enamel lake,
axis with a stronger ventral tilt; palatine forms pro- can be considered as ancestral characters [of the Le-
portionally shorter part of bony palate; and anterior porinae] in common with 'Alilepus' dietrichi, Pental-
~T·'· '· '·.'
~·/''' ·~·' -L· '''.··

·
·
·.. .r
/"
t· r
r. --
·
~·
'~ ' '.··
'
''
'
.-..\ r I.r
'
·"
·
·'
.~·
·. ~ - ·. . ·. . ·. r·.
''
'
'
·''~
·~~·
...,.. .. ..
..
.
·
·
··
: ..
'' r·.
A B
FIGURE 4. Diagra
MUBD 12001 and B, MUBD 11989.
agus, and Pronolagus.'" From the latter it can be fairlythat happens, in each case, to be situated exactly where
assumed that the P3 in primitive leporines had both one would expect a PIR to be? I believe the former to
AIR and PIR with equal frequency and, presumably,be the case, and that both Notolagus and Paranotolagus
of equal importance. Two P3's of Notolagus lepusculus fit well into Hibbard's (1963) scheme for the evolution
from Beck Ranch (MUBD 11989 and 12001, Fig. 4) of the P3 in leporines; I have therefore referred them
and one of N. velox from Yep6mera (IGCU 2734), to the Leporinae.
have distinct PIR's. Of the 86 specimens of Notolagus
that were examined, three (3.5 per cent) have PIR's Genus ARCHAEOLAGUS Dice, 1917
that extend to the bases of the tooth crowns. Notolagus
Archaeolagus Dice, University of California Publica-
is known only from the Blancan (N. lepusculus from
tions, Bulletin Department of Geology 10(12): 180,
Rexroad loc. 3, Truth or Consequences, and Beck
23 March 1917. Type, Lepus ennisianus Cope.
Ranch, and N. velox from Yepomera and Rancho Vie-
jo), and the genus, in the writer's opinion, represents Diagnosis--Single, narrow anterior reentrant on P2
a stage of development comparable to that in the de-(MAR) in adult specimens; internal reentrants on P3
velopment of the Lepus pattern (of Hibbard, 1963) inM2 usually straight-walled; p3_p4 less molariform than
Sylvilagus and Lepus, in which AIR is present 1.3 permolars; P3 with two external reentrants, anterior one
cent in Sylvilagus and 5 per cent in Lepus. Thus in shallow and devoid of cement, and posterior reentrant
Notolagus AIR is emphasized and in Sylvilagus and deep and filled with cement (Dawson, 1958).
Lepus PIR is emphasized.
Although AIR's are unknown in P3's of Paranoto- Genus PANOLAX Cope, 1874
lagus, the genus is known from only six specimens. If
Panolax Cope, Academy of Natural Sciences, Phila-
my interpretation is correct, one would expect to find
delphia, Proceedings 1874:151, 20 October 1874.
PIR's when additional specimens of P3's of this genus
are found. Diagnosis - "Single anterior reentrant on P2; on up-
AIR is unknown in the P3 ofpalaeolagines. It appears per molariform teeth internal reentrant straight walled,
for the first time in North America in the Clarendonian cement filled and crossing half or less of occlusal sur-
of Apache Canyon in specimens referred to Hypolagus face" (Dawson, 1958:59).
apachensis (specimens of this species having AIR's and Geologic Age and Distribution -?Clarendonian,
PIR's, in my opinion, exclude this species from the ?Hemphillian (Miocene) of New Mexico and possibly
genus Hypolagus. See "Remarks" under Hypolagus Clarendonian of southern California.
tedfordi). If such is the case, then the presence of AIR Remarks-Panolax is a monotypic genus (P. sanc-
in the ancestral pattern of the leporines distinguishes taefidei, USNM 1095) from the "Loup Fork forma-
the latter from the palaeolagines and from the archaeo- tion," New Mexico. Perhaps the occurrence in New
lagines. Mexico is in the Santa F6 group, either in the Tesuque
Assuming PIR to be a basic structure in an ancestral formation (Clarendonian) or in the Chamita formation
leporine, how does one handle its presence in Noto- (Hemphillian) (Galusha and Blick, 1971; McFadden
lagus? Is it in fact PIR, or a non-homologous variant et al., 1979).
Hypologus vetus(54)
H. porvip/icotus(1)
H. gidleyi (3 9)
H. regolis(39)
H. fonfinolis(17)
H. tedfordi(42) i 7
H. furlongi( 26) I_ _ _
H. voorhiesi(23)
H. edensis(36)
H. oregonensis (18)
H. r/ngo/densis (5 9) _
H. arizonensis (2)
Lepoides lepoides(22)
Pewelogus dowsonae (16)
Pp mexiconus(2)
PH. cpachensis( l)
0 1 AER5 20 5 30 35 40 45100
AER/P3 X 100
FIGURE 5. Dice-Leraas diagrams of the degree to which the anteroexternal reentrant (AER) extends across the width of the
occlusal surface of P3, expressed in percentages.
Wilson (1939) tentatively referred an upper left ?P4 to deep AER and deep PER, both filled with cement,
from the Avawatz local fauna (Clarendonian) of south- and usually without AR; crenulations on enamel of
ern California to Panolax. anterior surfaces of reentrants on p3-M2 decreases pro-
Dice (1929) and Dawson (1958) pointed out that in gressively posteriorly.
Panolax the upper molariform teeth have simple in- Geologic Age and Distribution - Barstovian (middle
ternal reentrants and there is a single anterior reentrant Miocene) to Blancan (late Pliocene) of North America,
on P2 that resembles the comparable structure in Ar- Pliocene of Eurasia.
chaeolagus more so than in Hypolagus, and that Pan- Remarks- Three species which up to now have been
referred to Hypolagus should be transferred from th
olax may represent a late survivor of a primitive stock.
It may be that Archaeolagus is referable to Panolax Archaeolaginae to the Leporinae.
but this cannot be determined until lower and upper Examination of specimens of Hypolagus browni re-
dentitions of Panolax are found in association. vealed that this species should be referred to Aluralagu
because there is, in adult specimens, an enamel lake
Genus HYPOLAGUS Dice on P3 formed by a fossettid enclosing a PER.
I agree with Miller (1980) that Dice's (1929) referral
Hypolagus Dice, University of California Publications,
of Lepus giganteus Brown, from the Conard Fissure
Bulletin Department of Geology 10(12):181, 23
deposits, to Hypolagus is erroneous. The specimen (ho-
March 1917. Type species: Lepus vetus Kellogg.
lotype) representing this species is a cranial fragment
Emended Diagnosis - Size small to medium; P2 with with P4-M' of a large leporid. It seems best to refer
deep MAR and shallow or absent EAR; P3 with shallow this specimen to Lepus as was done originally by Brown
(1908). The known characters of this specimen fall proximal end of tibia, LACM 7434; cranial fragments,
within the limits of variation of comparable characters LACM 24754; cranial fragments, four metatarsals,
in Lepus. other fragments, LACM 23431; cranium lacking zy-
The retainment of AIR and PIR in the P3 of adult gomatic arches, with P2, humeral and ulnar fragments,
Hypolagus apachensis Gazin suggests that this species LACM 13729; skull lacking zygomatic arches, LACM
is referable to the Leporinae and that it may represent 20923; cranial fragments, LACM 24581, 24585; skull
a new genus; the species is here designated as ?Hypo- lacking zygomatic arches with posterior ends of man-
lagus apachensis (see Figs. 2, 5-7, 9, and 12). The dibles and fragmentary skeletal parts, LACM 24921;
holotype and topotypes of Hypolagus tedfordi newskull lacking zygomatic arches, with basicranium, right
species were previously referred to ?Hypolagus apach-mandible, fragments of scapula, right and left proximal
ensis Gazin (Gazin, 1930; Wood, 1937; Dawson, 1958), radii and ulnae articulating with right and left humeri,
but differ from the latter species in being significantly LACM 25669; mandible with P3-P4, LACM 24965;
larger, having PER and AER more deeply incised, and mandible with P3-M3, LACM 40139; immature man-
lacking internal reentrants that extend to the base of dible, LACM 11510; mandible with P3-P4, LACM
the tooth crown in adults. 16859; mandible with I-M2, LACM 24579; mandible
with P3-M , LACM 24971; cranium lacking zygomatic
HYPOLAGUS VETUS (Kellogg) arches, with atlas in articulation, LACM 23447; P3,
(Fig. 7F) LACM 13742; femoral and tibial fragments, LACM
24670; astragalus and calcaneum, LACM 23417; distal
Lepus vetus Kellogg, University of California Publi-
humeral fragment, LACM 23422; articulated pes and
cations, Bulletin, Department of Geology 5(29):
other fragments, LACM 24957; P2, LACM 17687;
436, 13 July 1910.
skeletal fragments, LACM 24578, 24580; distal tibia,
Hypolagus vetus, Dice, University of California Pub-
LACM 39462; maxillary fragment with P2, LACM
lications, Bulletin, Department of Geology 10(12):
20428. Layer Cake-cranium with complete dentition,
181, 23 March 1917.
LACM 24969; cranium lacking basicranium, LACM
Holotype -Left mandible with P3-M2, UCMP 23444; mandible with P3-M3, LACM 24972; distal
12565, Hemphillian Miocene, UCMP locality 1100, femur, LACM 39459; pelvis and skeletal fragments,
Thousand Creek local fauna, Humbolt County, Ne- LACM 24665; rostrum with incisors and P3_p4, LACM
vada. 24966; palate with complete dentition, LACM 24967;
Emended Diagnosis-H. vetus differs from Hypo- mandible with P3, LACM 24968; astragalus, calca-
lagus parviplicatus in having a more deeply incised neum, proximal end of femur, humerus, scapula, distal
PER on P3, and usually an uncrenulated TH. It differs ends of femora with articulated tibiae, LACM 23425;
from H. oregonensis in lacking an AR on P3; from H. distal end of pes with metatarsals, LACM 3946 1; left
ringoldensis in its significantly smaller size, lack of an mandible with P3-P4, LACM 17667; mandible with
AR, and sigmoid TH; and from H. gidleyi in being P3, LACM 24750; three articulated pedes with meta-
significantly smaller in size, and having a sigmoid TH tarsals, tarsi, and other fragments, LACM 24719; cra-
in 90 per cent of P3's compared to only 20 per cent in nium, SDMNH 20445, IVCM, 1*. Truth or Conse-
H. gidleyi. From H. regalis, H. voorhiesi, and H. ari- quences-P3, USGS-M, 1*.
zonensis, H. vetus differs in having a sigmoid TH, and Geologic Age and Distribution - Hemphillian of Ar-
a PER that is not strongly deflected anteriorly. A few izona, Idaho, Nevada, Oregon; Blancan of California
specimens of H. vetus are small in size and may rep- and Baja California.
resent immature individuals. These smaller-sized Description--The proportions of the skull in Hy-
specimens can be distinguished from species that polagus
arevetus closely resemble those in Sylvilagusflor-
significantly smaller in size than H. vetus by idanus (Fig. 11). Comparing the crania ofH. vetus with
the fol-
lowing characters: from H. furlongi by a sigmoid the cranium
TH; of H. gidleyi reveals that the choana in
from H. edensis by a markedly shallower AER; and the latter is narrower, the maxillary tooth row is short-
from H. fontinalis and H. tedfordi by a more deeply er, and the front upper incisor is wider than in the
inflected PER on P3. former. If the known skulls of H. vetus and H. gidleyi
Referred Specimens--Asterisk (*) refers to number lacked dentitions, they could reasonably be referred to
of uncataloged specimens at a locality. Hemphillian. Sylvilagus (see White, 1984, figs. 1-3). Of the 25 mea-
Redington-P3, AMNH(FAM) 108736, 110551. surements taken on the skull of H. gidleyi, only one,
Rome-P3, LACM(CIT) 6513, USGS-M, 1*. Star Val- the depth of the anterior end of the zygomatic arch,
ley-mandibles, IMNH 27888, 29115, 29129; P3, fits into the observed range of that in Lepus capensis,
IMNH 27889, 27891, 27893, 27896, 27921-27923, while all others fit into comparable ranges in Sylvila-
27925, 27926, 29113, 29114. Thousand Creek-frag- gus. This does not confirm the analysis of Campbell
mentary skull, AMNH(FAM), 1*, LACM(CIT), 2*; (1969) of the postcranial skeletal elements of Hypo-
mandibles, AMNH(FAM), 2", LACM(CIT), 2*,lagus near vetus (=H. gidleyi) in which he concluded
that the latter species was functionally nearest L. eu-
UCMP, 1"; P3, LACM(CIT), 53*.
Blancan. Arroyo Seco--skull with complete upperropaeus (=L. capensis), the feral hare that occurs in
and lower dentitions, distal one third of humerus, northestern United States (Hall, 1981).
Hypologus vetus(55) _ _ __
H. par viplica tus( 1)
H. gidley/(4 0) f -
H. rego/is(40)
H. foninao/is ( 17) r(1
H. ledfordi(42)
H. furlongi/(20)
H. voorhiesi(23)
H. edensis(35)
H. oregonensis(108)
H. ringo/densis(59)
H. orizonensis(2)
Lepoides /epoides(21)
Pewe/ogus dowsonoe(1 6)
. Rmexicanus (2)
PH. opochensis(ll) i L
40 . 45 50 55 60 65 70%
PER/WP X100
FIGURE 6. Dice-Leraas diagrams of the degree to which the posteroexternal reentrant (PER) exten
the occlusal surface of P3, expressed in percentages.
The P2 in H. vetus is oval in cross section, and has regalis are here referred to H. vetus bcause they closely
a deep and smooth-sided MAR and shallow EAR, both resemble specimens of H. vetus from the Anza-Borrego
of which are filled with cement. The post-P2 teeth have Desert section and from Thousand Creek, and PER on
slight crenulations on the anterior walls of the internalP3 is not deflected anteriorly as in H. regalis.
reentrants, which become less crenulated progressively Specimens from Las Tunas, Baja California, which
posteriorly to M2. The M3 is ovoid in cross section Miller (1980) tentatively referred to H. vetus, are here
with thick enamel on the anterior and medial edges of referred to H. vetus because the enamel patterns on P3
the occlusal surface. closely resemble those from Thousand Creek and other
The P3 is triangular in cross section with the labial localities, and the size of P3 fits into the range of H.
side directed from lateral to slightly anterior. AER vetus.
is
usually smooth-sided and extends from 9 to 28 per
cent across the occlusal surface, averaging 18 per cent HYPOLAGUS PAR VIPLICATUS Dawson
(Fig. 5). PER extends from 40 to 62 per cent across (Fig. 7G)
the occlusal surface, averaging 52 per cent (Fig. 6). TH
Hypolagusparviplicatus Dawson, University of Kansas
is sigmoid in 90 per cent and straight in 10 per cent
Paleontological Contributions, Vertebrata 6:45, 1
of the specimens, is deflected from 9 degrees anteriorly
May 1958.
to 8 degrees posteriorly, averaging 1.5 degrees ante-
riorly (Fig. 2). TN is slightly crenulated in 15 of the Holotype--Right mandible with P3-M3, UCMP
60 specimens examined. The enamel patterns on P4- 11571, Barstovian (middle Miocene), Virgin Valley,
M3 are essentially as in all other species of Hypolagus. Humbolt County, Nevada, UCMP locality 1065.
Remarks -Specimens from Radec and Pechanga that Diagnosis - Hypolagus parviplicatus differs from H.
Golz, Jefferson, and Kennedy (1977) referred to H. fontinalis and H. tedfordi in its markedly larger size
Al A2 A3 A4 BI B2 B3
B4 Cl C2 C3 C4 DI D2
D3 D4 El E2 E3 E4 F F2
F3 F4 G H II 12 13 14 JI
J2 J3 J4 K LI L2 L3 L4 MI
M2 M3 M4 NI N2 N3 N4 01 02
03 04 P QI Q2 RI R2 R3 R4
SI S2
0I5mm
FIGURE 7. Enamel patterns on th
and sp. AMNH(FAM) 108730, UNSM 91155, 1090-74, AMNH(FAM) 108721; B, Hypolagus ringoldensis UWBM 35237,
MUBD 9389C, UMMP V56573A, UNSM 91154; C, H. gidleyi USNM 336218, 23573 (holotype), WTSU 4468B, UWBM
42496; D, H. oregonensis SDSM 11515, LACM(CIT) 57682, OU F2063, F4037; E, H. regalis UMMP V29678, V41362,
MUBD 12045, UNSM 91184; F, H. vetus USGS-M 1578, AMNH(FAM) 108736, LACM 24971, LACM(CIT) locality 63;
G, H. parviplicatus UCMP 11571 (holotype); H, H. fontinalis UCMP 28525; I, ?Hypolagus apachensis LACM(CIT) 5183A,
5183 (holotype), 123647:36, 38; J, H. tedfordi n. sp. LACM(CIT) 123645, 123647:29, 123644, UCMP locality V6742; K,
Hypolagus cf. H. tedfordi UF 23992; L, H. fontinalis UCMP 33985B, 29540, 33425, 29630; M, H. edensis USGS-M 1431,
LACM(CIT) 1972, IMNH 33639, LACM 24930; N, H. furlongi IMHN 29573, 30596, WTSU 4468G, IMNH 29564; 0, H.
voorhiesi UNSM 51844, 91131, 85366, 92004; P, Hypolagus cf. H. voorhiesi UCMP 78497; Q, H. arizonensis UALP 1013A
(holotype), 1013B; R, Pewelagus dawsonae LACM 13742, LACM locality 6331, IVCM 2486, 308; S, ?Pewelagus mexicanus
IGCU 4158 (holotype), 4075.
and less well incised AER; and from all other species Referred Specimens - Asterisk (*) refers to the num-
of Hypolagus in a markedly less well incised PER on ber of uncataloged specimens at a locality. Blancan.
P3. Cita Canyon-P3, WTSU 1235, 3193. Coffee Ranch-
Description - Hypolagus parviplicatus is known only P3, MUBD 11612A-11612F; P2, MUBD 11612G; P3,
from the holotype. P3 is longer than wide, trigonid UCMP 30726, 30726A. Flat Iron Butte-mandible,
markedly large, AER and PER are shallow and cement- IMNH 29584; P3, IMNH 5389. Hagerman-mandi-
filled, and TN is strongly crenulated. The patterns of bles, IMNH 29534-29536; P3, IMNH 4978, 30616,
the occlusal surfaces of P4-M3 are essentially as in other 34849, 34858; UMMP, V48946, V49711, V50222; P3,
species of Hypolagus. The swelling marking the base V53473; USGS-M 1*; USNM 12641, 336221, 336222;
of P3 is present below the anterior portion of P3 at the P3, USNM 336210, 336212, 336213. Ninefoot Rap-
occlusal surface. There is a slight pitting on the man- ids-P3, IMNH 29570. Oreana--mandible, IMNH
dible below P3-P4, and the masseteric fossa extends 29533. Oshkosh-P3, UNSM 52584. Railroad Can-
anteriad to just below the anterior portion of M3. yon-mandible, AMNH(FAM) 108732. Red Corral-
P3, WTSU 4468A, B, C, D. Sand Point-P3, IMNH
HYPOLAGUS cf. H. PAR VIPLICATUS
30409. Taunton- mandible, UWBM 42593; P3, 42494,
No additional specimens from the late Barstovian, 42496, 46225-46227. Trench Canyon-P3, UCMP
Quarry C, LACM(CIT) locality 172, approximately 9 73332. Chamita- mandible, AMNH(FAM) 116845.
miles north of Tonopah, Nevada, which Dawson (1958) Geologic Age and Distribution - Hemphillian of
referred to Hypolagus near H. parviplicatus have be- Texas and New Mexico, Blancan of Idaho, Nebraska,
come available. Dawson's (1958) analysis can there- Texas, and Washington (Fig. 12).
fore not be confirmed or modified. The specimens are Description--The cranium in Hypolagus gidleyi
(USNM 23573, the holotype) is essentially as in Syl-
of immature individuals and cannot be used in deter-
mining the size-range of H. parviplicatus. vilagus floridanus except that the choana is propor-
Specimen UWMG is a right mandible with P3-M3 tionally narrower, length of maxillary tooth row at al-
from Trail Creek Quarry in the early Valentinian veoli is shorter, and width of anterior upper incisor is
Ash
wider. The narrow choana may be related to the im-
Hollow formation (Cassiliano, 1980). It differs from
Hypolagus parviplicatus in P3 having a triangularmaturity
cross of the specimen. The mandible is propor-
tionally
section, relatively smaller trigonid, only a trace of ce- as in S. palustris except that it is deeper below
MI. The proportions of the skull in H. gidleyi strongly
ment in AER, labial surface of P3 facing anterolaterad,
suggests this species to be referable to a cottontail eco-
a relatively large posterior lobe on M3, and in being
morph.
slightly smaller in size. It resembles H. parviplicatus
in P3 having a crenulated TN, PER being incised 48.6 The P2 is ovoid in cross section with a deep and
smooth-sided MAR and shallow EAR, both of which
per cent of occlusal surface, and similar enamel pat-
terns on P4-M2. The parameters of variation areare filled with cement. P3_M2 have internal reentrants
un-
known for H. parviplicatus, and this specimen is only extending approximately one half way across occlusal
tentatively referred to the latter species. surface, cement-filled, and with crenulations on the
enamel of the anterior walls of the reentrants, which
HYPOLAGUS GIDLEYI new species become less crenulated progressively posteriorly as in
(Fig. 7C) other species of Hypolagus. M3 is ovoid in cross section
with enamel on the anterior and lateral margins of the
Holotype - Complete skull of an immature individ- occlusal surfaces.
ual, USNM 23573, Blancan Pliocene, from the em- The P3 is triangular in cross section and faces lat-
bayment south of the horse quarry, Hagerman local erally and slightly anteriorly. TH is straight in 82 pe
fauna, in the Glenns Ferry formation (Hibbard, 1969,cent and sigmoid in 18 per cent of observed specimens
figs. lA, B, 3F, G). PER is deflected from 10 degrees anteriorly to 11 de-
Diagnosis--Hypolagus gidleyi differs from H. vetus grees posteriorly, averaging one degree anteriorly (Fig
in its significantly larger size, and having TH straight 2). TN on PER is crenulated on 38 per cent of speci-
in 90 per cent of specimens. It differs from H.fontinalis mens. The post-P3 teeth are essentially as in other
and H. tedfordi in its markedly larger size, more deeplyspecies of Hypolagus.
incised PER, and straight TH. It differs from H. edensis Remarks - Specimens from the Chamita formation
in a markedly less well incised AER and markedly and those from Coffee Ranch that were referred to H.
larger size. From H. furlongi it differs in its markedly vetus by MacFadden, Johnson, and Opdyke (1979) are
larger size. It differs from H. voorhiesi, H. regalis, and here referred to Hypolagus gidleyi because TH on P3
H. arizonensis in PER not being deflected anteriorly. is straight and the P3's are larger than in H. vetus.
From H. parviplicatus it differs in a more deeply incised
AER and PER on P3. From H. oregonensis it differs HYPOLAGUS cf. H. GIDLEYI
in lacking an AR and a straight TH. It differs from H.
ringoldensis in lacking an AR and being significantly Specimens from the Barstovian of Wood Mountain,
smaller in size. Saskatchewan that Storer (1975) tentatively referred to
PER on P3 extends from 36 to 51 per cent of the
Hypolagus vetus are here tentatively referred to H. gid-
leyi because TH on P3 in the specimens as describedocclusal width, averaging 43 per cent, and TH is sig-
moid. PER, furthermore, is deflected 1.4 degrees pos-
and figured by Storer is straight and not sigmoid, and
the upper cheek teeth seem to have more crenulations
teriorly, ranging from 3 degrees anteriorly to 7 degrees
than in H. vetus. AIR and PIR are absent from all posteriorly (Fig. 2), and TN is crenulated in 29 per cent
specimens of H. vetus and H. gidleyi which were ex-
of specimens observed. AER on P3 varies from 14 to
amined, but present, although shallow, in the Sas- 27 per cent of the occlusal width, averaging 21 per
kachewan specimens. cent.
As noted by Dawson (1958), the mental foramen i

present below the posterior half of the lower diastem
HYPOLAGUS FONTINALIS Dawson
with variable pitting below P3 and P4. The anterior
(Fig. 7H, L) border of the masseteric fossa is situated below the
talonid of M2. There is a swelling on the mandible a
Hypolagus fontinalis Dawson, University of Kansas,
Paleontological Contributions, Vertebrata 6:48,the
1 base of the lower incisor, and the latter extends
May 1958. posteriorly under P3. A postalveolar foramen is pres-
ent.
Holotype - Incomplete left mandible with P3-M2,
Remarks - When Lindsay (1972) referred specimens
UCMP 33425, from UCMP locality V3322, Big Spring from Barstow to Hypolagus parviplicatus he included
Canyon, Bennet County, South Dakota. in his analysis specimens from Tonopah along with
Emended Diagnosis--Hypolagus fontinalis differs the holotype. The specimens from Tonopah, as Daw-
from H. tedfordi in the posterior deflection of PER, son (1958) noted, have internal reentrants that extend
and its significantly larger anteroposterior length of P3.
only a short distance down the side of P3, thus indi-
From H. parviplicatus it differs in its markedly smallercating the specimens to be immature and of doubtful
size and in an uncrenulated TN on PER. From all other
species assignment. The Barstow specimens differ from
species of Hypolagus, H. fontinalis differs in a signif-the holotype of H. parviplicatus in not having a strongly
icant shallower inflection of PER on P3. expanded trigonid and by markedly smaller size.
Referred Specimens -Clarendonian. From Dawson
(1958): 29540,
UCMP Big Spring
FMNHCanyon--incomplete mandibles,
P15812, P15813, P15814. Todd HYPOLAGUS cf. H. FONTINALIS
County, South Dakota--incomplete mandible, SDSM
5549. Quinn Canyon-incomplete mandibles, FMNH LACM(CIT) 107301 is a left P3 from the Claren-
P26938, P26939, P26940. Fish Lake Valley-incom- donian of Santa Margarita, San Luis Obispo, Califor-
plete right and left mandibles, anterior upper incisor, nia. This specimen is tentatively referred to Hypolagus
UCMP 29630. fontinalis because since AER on P3 is more deeply
From Clark et al. (1964): Fish Lake Valley-left incised than in H. vetus. It is larger in size than spec-
mandible with P4-M2, MCZ 7640; left premaxilla withimens of H. fontinalis.
upper incisors, MCZ 7641; isolated teeth, MCZ 7642; UCMP45031 is a right P3 from the Hemphillian of
postcranial fragments including proximal and distal Old Stiener Ranch, Lander County, Nevada. It is
ends of humerus, astragali, calcanea, and navicular, smaller in size than any specimen of Hypolagus vetus.
MCZ 7643. Although the specimen is morphologically similar to
H. fontinalis, its occurrence in the Hemphillian and
Barstovian.
with Barstow--incomplete
P3-M1, UCMP 28525; incomplete right mandible
its morphological
mandible with resemblance (except in size) to H.
vetus makes this specimen only tentatively referrable
P3-P4, UCMP 78514; incomplete mandible with M1-
to H. fontinalis.
M3, UCMP 25826; P3, UCMP 78497, 78507, 78513,
131051; 2 P3's, P2, 4 lower cheek teeth, 10 upper cheek
teeth, and bone fragments, UCMP 35397; upper cheek
HYPOLAGUS TEDFORDI new species
tooth, UCMP 131050; DP3, UCMP 78495, 78504,
(Fig. 7J)
78509; deciduous upper premolars, UCMP 78496,
78505, 78506, 78508, 78510, 78511, 78512, 78515, Holotype -Incomplete left mandible with P3,
78519; upper cheek teeth, UCMP 78498, 78520, 79144; LACM(CIT) 132 from locality LACM(CIT) 64, Apache
tooth fragments, UCMP 28528. Canyon, Ventura County, California.
Geologic Age and Distribution - Barstovian of Cal- Diagnosis - H. tedfordi differs from H. fontinalis in
ifornia, Clarendonian ofNebraska, Nevada, and South its significantly smaller size, and anterior deflection of
Dakota (Fig. 12). PER on P3. From all other species of Hypolagus it
Description - P2 has a shallow EAR and deep MAR. differs in a markedly less well incised PER. It is sig-
The internal reentrants on p3-M2 extend laterad more nificantly smaller in size than H. vetus, H. gidleyi, H.
than one half of the width of the occlusal surface, and regalis, H. oregonensis, and H. ringoldensis.
the walls of these reentrants are strongly crenulated on Referred Specimens--Asterisk (*) refers to numbers
P3 and less so progressively posteriorly on p4-M2. of uncataloged specimens at a locality. Clarendonian.
Apache Canyon - left mandible with P3-M1, HYPOLAGUS FURLONGI Gazin
LACM(CIT) 123647, 123647-11; left mandible with (Fig. 7N)
P3-P4, LACM(CIT) 123645, 123647; right mandible
Hypolagus furlongi Gazin, Proceedings of the United
with P3-P4: LACM(CIT) 123647-13, 123647-35; P3,
States National Museum 83(2976):118, 1934.
LACM(CIT) 63, 123647-32, 123647-38, 123644. Ri-
cardo-right mandible with P3-M,, UCMP, 1*; right Holotype - Right mandible with I-M2, LACM(CIT)
P3, UCMP, 14*; left P3, UCMP, 16*. Quatal Canyon- 1321, Grand View local fauna, Glenns Ferry forma-
right P3, USGS-M, 1*. tion, Idaho.
Hemphillian. Mulholland--left P3, UCMP, 1*. Wik- Emended Diagnosis - Hypolagusfurlongi differs from
ieup-left mandible with P3-P4, UALP 24; right man- H. voorhiesi, H. regalis, and H. arizonensis in not hav-
dible with P3-M3, AMNH(FAM) 116844. Reding- ing a strong anterior deflection of PER. Its deeply in-
ton-right mandible with P3-M3, AMNH(FAM) flected PER sets it apart from H. fontinalis, H. tedfordi,
108734. and H. parviplicatus. From H. edensis it differs in hav-
Geologic Age and Distribution - Clarendonian of ing a significantly shallower AER. From H. vetus it
California, Hemphillian of Arizona and California (Fig. differs in a straight TH and more deeply incised PER.
12). From H. gidleyi it differs in its significantly smaller
size with no overlap of observed ranges; and from H.
Etymology--Named
Description for Richard
- The holotype H. Tedford.
and topotypes of H. ted- oregonensis and H. ringoldensis it differs in lacking an
fordi were recovered from the same locality as ?Hy- AR on P3 and by its markedly smaller size.
polagus apachensis, and the upper and lower dentitions Referred Specimens--Asterisk (*) refers to number
have not been found in association; it is therefore not of uncataloged specimens at a locality. Blancan. Red
possible to refer the upper dentitions to either species Corral-left P3, WTSU 4093A, 4464, 4467A, 4477A,
with certainty. Wood (1937, figs. 8 and 10) noted dif- B, C; right P3, WTSU 4463, 4477D, E. Del Valle-
ferences between two P2's that he attributed to different right P3, USGS-M, 1*. Ninefoot Rapids-right man-
stages of wear. The post-P2 teeth are characteristically dible with P3-P4, IMNH 29575; left P3, IMNH 29573,
leporid in structure. The mandibular dentition in H. 30381. Grand View--left mandible with P3-M2, IMNH
tedfordi is characteristically archaeolagine in structure. 29564; right mandible with P3-P4, IMNH 29825; right
In P3 the AER is uncrenulated and extends mediad mandible with P3, IMNH 31394; left P3, IMNH 29913,
from 12 to 29 per cent, averaging 20 per cent across 29915, 30395; right P3, IMNH 29826, 29828, 29830,
the occlusal surface, and PER from 34 to 55 per cent, 29916, 29925, 30396. Big Springs-right P3, UNSM
averaging 45 per cent, respectively. TH is sigmoid to 51638, 85363, 91121.
straight, and TN is crenulated in 14 per cent of spec- Geologic Age and Distribution - Blancan of Califor-
imens. PER is deflected from a line perpendicular to nia, Idaho, Nebraska, and Texas (Fig. 12).
a line of orientation (Fig. 2) from 9 degrees anteriorly Description --Hypolagus furlongi is comparable in
to 4 degrees posteriorly, averaging 3 degrees anteriorly. size to H. edensis, H. tedfordi, and H. voorhiesi. AER
Remarks--The holotype and topotypes of H. ted- on P3 extends from 15 to 29 per cent, averaging 21 per
fordi were previously referred to Hypolagus apachensis cent, and PER extends from 50 to 61 per cent, aver-
Gazin (Gazin, 1930; Wood, 1937; Dawson, 1958), but aging 55 per cent of the distance across occusal surface
differ from the latter species in being slightly larger, of P3. There is a very shallow AR in 20 per cent of the
having PER and AER more deeply incised, lacking specimens of H. furlongi. PER is deflected from a line
internal reentrants that extend to the base of the crown perpendicular to a line of orientation (Fig. 2) 19 degrees
of the tooth in adults, and having TH deflected anteriad anteriorly to 10 degrees posteriorly, averaging 2 de-
and not posteriad as in ?Hypolagus apachensis (Fig. 2).grees posteriorly. The upper dentition and post-P3 den-
The retainment of AIR and PIR in adults of ?Hypo- tition is essentially as in H. edensis. TN on P3 is cren-
lagus apachensis suggests that this species is referableulated in 71 per cent and AER in 17 per cent of
to the Leporinae, and that it may represent a new genus specimens.
(Figs. 2, 5, 6, and 71).
HYPOLAGUS EDENSIS Frick
HYPOLAGUS cf. H. TEDFORDI
(Fig. 7M)
(Fig. 7K)
Hypolagus edensis Frick, University of California Pub-
UF 23992, a right P3 from the Mobil Mine, Lower
lications, Bulletin, Department of Geology 12(5):
Bone Valley, Hillsborough County, Florida, has a rel-
348, 1921.
atively shallow inflection of PER on P3 and general
Hypolagus limnetus Gazin, United States National
proportions that closely resemble comparable struc-
Museum, Proceedings 83(2976):114, 1934.
tures in specimens ofHypolagus tedfordi. Although this
specimen is tentatively referred to H. tedfordi, the ir-Holotype - Fragmentary right mandible with P3-M,
regularities on the enamel of the anteromedial edge ofofan immature individual, UCMP 23376 from UCMP
P3 suggests that this may be an immature individual. locality 3269, Riverside County, California, Mount
The sides of this tooth are parallel. Eden local fauna.
Emended Diagnosis--H. edensis is distinguishable Arlington--P3, LACM(CIT) 57682, 57684. Christm
from all other species of Hypolagus by the presence of Valley--P3, USGS-M, 1*. McKay Reservoir--m
a deep and smooth-sided AER on P3. dibles, UO F2063, F2097, F3663, F4036, F4067; pal-
Referred Specimens--Asterisk (*) refers to number atal fragments, SDSM 11506, 1*; mandibles, SDSM
of uncataloged specimens at a locality. Blancan. Hag- 11500-11504, 11521, 11522, 11524, 11525.
erman-cranium, right mandible and atlas (Gazin, Geologic Age and Distribution--Late Hemphillian
1934, fig. 2), USNM 12619; fragmentary mandible, of Oregon.
UMMP V54782, USNM 336217; P3, IMNH 4985, Description--The proportions of the mandibles in
4986, 4988, 30905, 33639, 33641, UMMP V49714, Hypolagus oregonensis resemble those in H. vetus (Fig.
V53372A, V53372B, V53372C. Panaca- fragmentary 9). P3 resembles that of H. vetus but unlike the latter,
mandible, AMNH(FAM) 108705, 108711. Red Cor- there is a well-developed AR. AER on P3 is incised
ral-fragmentary mandible, WTSU 4094, 4465, 4468E, across the occlusal surface from 13 to 21 per cent,
4478F. Arroyo Seco-P3, LACM 23174, 1*. Del Valle- averaging 49 per cent. AER is crenulated in 30 per cent
P3, USGS-M, 1*. Elk Hills-P3, USGS-M, 1*. Arroyo of specimens observed. In all other structures, the man-
Pequeno, Kings County, California-P3, UCMP, 1*. dibles resemble those in H. vetus.
Mission Viejo-P3, LACM 119109. Ninefoot Rap- Remarks--AR is present on P3 in all specimens o
ids-P3, IMNH 29569, 30383. Taunton-P3, UWBM H. oregonensis, although in some specimens it is barel
42529, 46224. discernible (Fig. 7).
Hemphillian. Kern River-P3, LACM(CIT) 1972, The tiny "enamel lakes" on P3 of H. oregonensis
1973, 1976, 1977. Pinole Tuff-P3, UCMP 57805, described by Shotwell (1956) are also present in all
64790. Redington-P3, AMNH(FAM) 110553. known leporids. They seem to be dentinal tubules.
Geologic Age and Distribution - Hemphillian of Ar-
izona and California, Blancan of California, Idaho, Ne- HYPOLAGUS cf. H. OREGONENSIS
vada, Texas, and Washington (Fig. 12).
UO F2907 and SDSM 11521 (P3's) from McKay
Description--The known skull (USNM 12619) of Reservoir and LACM(CIT) 57684 from Arlington, are
Hypolagus edensis is proportionally similar to the skull
tentatively referred to H. oregonensis since they occur
in Sylvilagus bachmani and S. nuttallii. The width of
with specimens of that species at the two localities. On
the choana is smaller than in the above species, but its
the other hand, they are also indistinguishable from
small diameter may reflect an immature individual
the P3's of H. vetus which occur at Truth or Conse-
(Fig. 10).
quences and Rome. McKay Reservoir and Arlington
The P3 in Hypolagus edensis varies little from lo-
are biostratigraphically older than Truth or Conse-
cality to locality. Specimens from Kern River are larger
quences, and the possibility of H. vetus occurring at
than those from other localities but are clearly referable the two localities thus cannot be ruled out.
to H. edensis. The holotype is the smallest specimen
of H. edensis and is of an immature individual. AER
HYPOLAGUS RINGOLDENSIS Gustafson
on P3 is deeply incised, ranging from 24 to 46 per cent
(Fig. 7B)
of the occlusal width, averaging 34 per cent. PER is
incised across the occlusal surface of P3, ranging Hypolagus
from ringoldensis Gustafson, Museum of Natural
45 to 59 per cent, averaging 51 per cent of occlusal History, University of Oregon, Bulletin 23:15,
width. TH is straight in 71 per cent and sigmoid in 29 March 1978.
per cent, and TN is crenulated in 49 per cent and
Holotype - Mandible with P3-M2 and broken M3,
uncrenulated in 58 per cent of observed specimens.
AER is uncrenulated.
UWBM 45015 from UWBM locality A8814, White
Bluffs south of Ringold Coulee, Franklin County,
Washington, from a conglomerate about 45 feet above
HYPOLAGUS OREGONENSIS Shotwell the White Bluffs Tuff.
(Fig. 7D) Emended Diagnosis - Hypolagus ringoldensis differs
from H. oregonensis in its significantly larger size, hav-
Hypolagus oregonensis Shotwell, Geological Society of
ing TH on P3 30 per cent sigmoid as opposed to 64
America Bulletin 67:727, June 1956.
per cent in H. oregonensis, and AER and PER being
Holotype--Left mandible with P3-M3, UO F4094, more deeply incised across the occlusal surface of P3.
late Hemphillian, Miocene, McKay Reservoir, 5 miles It differs from all other species of Hypolagus by the
south of Pendleton, Umatilla County, Oregon, in the presence in 40 per cent of observed specimens of a
Dalles Formation. strongly incised AR on P3. From H. regalis it differs
Emended Diagnosis -Hypolagus oregonensis differs in the absence of a strongly anteriorly deflected and
from H. ringoldensis in having a sigmoid TH on shallower
P3, PER. From all other species of Hypolagus it
and being significantly smaller in size. From all differs
other in its significantly larger size and more deeply
species of Hypolagus it differs in having an AR.incised AER, except in H. edensis.
Referred Specimens --Asterisk (*) denotes number Referred Specimens - Blancan. Beck Ranch- man-
of uncataloged specimens at a locality. Hemphillian.dible, MUBD 9393, 9394; P3, MUBD 9388A-9388C,
9389A-9389D, 12040, 12043-12045, 12061-12064, AR and straight TH, which closely resembles speci-
12089-12092. Benson-P3, UALP 1505. Fox Can- mens of H. ringoldensis. The Rancho El Ocote speci-
yon-mandible, UMMP V24786, 24787; P3, UMMP men is tentatively referred to H. ringoldensis because
V24784A-V24784D. Rexroad, locality 3- P3, UMMP it resembles the latter in all characters except an ex-
V46572, V46573A-V46573E, V46574A-V46574F. tremely incised AR.
White Bluffs--incomplete palate, UWBM 46206;
mandible, UWBM 35237, 40259, 40286, 40293,
HYPOLAGUS REGALIS Hibbard
46843; P3, UWBM 4613, 34977, 35118A, 35118B,
35128, 35145A, 35145B, 40313A, 40313B, 40393, (Fig. 7E)
40396, 40428A, 40428B, 41548, 41975, 42148, 42220, Hypolagus regalis Hibbard, American Midland Nat-
45020, 46169, 50720. uralist 21(2):510, 10 May 1939.
Hemphillian. Santee--mandible, UNSM 2020-73,
2583-78; P3, UNSM 1084-70, 2573-78, 2582-78, 2584- Holotype-Left mandible with I-M2, KU 4572, lo-
78, 2591-78, 2592-78, 2595-78, 2596-78, 91151, cality 3, Rexroad local fauna, Meade County, Kansas,
in the Rexroad formation.
91157; P2, UNSM 85378, 91164, 91166, 91171, 91173,
Emended Diagnosis - Hypolagus regalis differs from
91177, 91191-91195. White Cone- P2, UALP WC8 1;
H. voorhiesi in a more deeply incised AER and more
P3, UALP WC111, WC 1 WC 112, WC 113. Upper Bone Val-
ley-P3, UF 95700. strongly anteriorly deflected PER. From H. arizonensis
Geologic Age and Distribution - Hemphillian of Ne- it differs in its markedly larger size with no overlap of
braska, Florida, and Arizona, Blancan of Kansas, Tex- observed ranges. From all other species of Hypolagus
as, and Washington (Fig. 12). it differs in a strong anterior deflection of PER on P3.
Description - Hypolagus ringoldensis is significantly Referred Specimens--Asterisk (*) denotes the num-
larger in size than all other species of Hypolagus. The ber of uncataloged specimens at a locality. Blancan.
mandible has proportions similar to those in other Beck Ranch-P3, MUBD 12039, 12041, 12046, 9388A,
species of the genus and of Sylvilagus aquaticus (Fig. 9388C, 9389D, 12042, 12044, 12045. Rexroad, lo-
9). P3 resembles that of other species of the genus but cality 3--cranial fragment, UMMP V4236; mandibles,
is significantly larger, has AER incised from 14 to 33 UMMP V26375, V28684, V29678; P3, UMMP
per cent, averaging 21 per cent of occlusal width, and V39777, 14*, V56572, V56574B, V56574C, V56574H,
PER similarly incised from 42 to 65 per cent, averaging V56575A-V56575K. Wendell Fox--skull fragment,
55 per cent. PER is deflected from a line perpendicular UMMP V35075; P3, UMMP V41362, V56673A,
to a line of orientation (Fig. 2) from 12 degrees ante- V56673B, V56673D.
riorly to 10 degrees posteriorly, averaging 1 degree Hemphillian. Santee-P3, UNSM 1080-75, 91184,
91179.
posteriorly. AER is crenulated in 22 per cent, and PER
in 40 per cent of observed specimens. The post-P3 Geologic Age and Distribution--Late Hemphilli
dentition is as in other species of the genus. P2 has a of Nebraska, Blancan of Kansas and Texas (Fig. 12).
deeply incised MAR and a shallow EAR, thus resem- Description --P2 in H. regalis has a deep MAR and
bling the comparable structure in other species of the shallow EAR, both of which are filled with cement. P3
genus.
is quite distinctive in the sharp deflection or forward
Remarks - Specimens from Beck Ranch that were turning of the innermost part of PER. AER and TH
referred to Hypolagus regalis by Dalquest (1978) are are crenulated in 42 per cent of observed specimens;
here referred to H. ringoldensis because there is an AR P4-M2 are as in other species of the genus; and the
on P3 in some specimens and, as Dalquest (ibid.) point- posterior lobe of M3 is the same in size as the anterior
ed out, some specimens resemble those from Fox Can- lobe (Hibbard, 1969, fig. 2B).
yon that are now referable to H. ringoldensis. Remarks - Hibbard (1939) referred specimens from
Specimens from White Cone that Baskin (1979) ten- Rexroad locality 3, Wendell Fox, and Fox Canyon to
tatively referred to Hypolagus vetus are here referred H. regalis, specimens that in this study are referred to
to H. ringoldensis because they are significantly larger H. ringoldensis, and which Hibbard (1969) excluded
in size than specimens of H. vetus and H. gidleyi, and from H. regalis.
TH on P3 is straight and not sigmoid as it is in H.
vetus.
HYPOLAGUS VOORHIESI new species
(Figs. 8B, E, 70)
HYPOLAGUS cf. H. RINGOLDENSIS
Holotype - Left mandible with P3-M2, UNSM
BUNM A7960, a mandible from Albuquerque, pos- 51653, from UNSM locality AP103, Big Springs, Ne-
sibly from the Chamita formation, Hemphillian, is ten- braska, Big Springs local fauna.
tatively referred to H. ringoldensis. It is close, mor- Diagnosis --Hypolagus voorhiesi differs from all
phologically, to the latter species but lacks an AR onspecies of Hypolagus, except H. arizonensis and H.
P3. regalis, in having a strong anterior deflection of PER,
IGCU 5006 from the Hemphillian of Rancho El and an anterior abrupt deflection of innermost PER.
Ocote, Guanajuato, Mexico (Miller and Carranza-Cas- It differs from H. arizonensis in a shallower AER and
tafieda, 1982, fig. 2D) is a P3 with a strongly incised less well-developed anterior deflection of PER. From
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FIGURE 8. Diagrams of the occlusal surfaces of cheek teeth in Hypolagus voorhiesi n. gen. and sp., Lepoides lepoides n. gen.
and sp., and Hypolagus ringoldensis. A, C right P2's and D right P3-M2 (holotype) of L. lepoides. F. right P2 of H. ringoldensis.
H. regalis it differs in its significantly smaller size and Blanco--right mandible with P3-M2, Joe Taylor Col-
less well-developed anterior deflection of PER. lection (WTSU). Broadwater-left P3, UNSM 83423.
Referred Specimens --Blancan. Big Springs--max- White Rock--left P3, UMMP V60633A. Deer Park-
illary fragment with P3-M', UNSM 47348; left P2, left P3, UMMP V31955. Ninefoot Rapids-left P3,
UNSM 85352, 85353, 85377, 91130, 91133, 91135; IMNH 30382, 36957; right P3, IMNH 29572.
right P2, UNSM 85355-85359; left mandible with P3- Geologic Age and Distribution--Late Blancan (late
M2, UNSM 92004; left P3, UNSM 51635, 51843, Pliocene) of Idaho, Kansas, and Nebraska (Fig. 12).
51844, 85366, 85372, 85375, 85386, 85388, 85393, Etymology-Named for Michael R. Voorhies.
85394, 91113, 91126, 91127, 91129, 98269; right P3, Description--The upper dentition and post-P3 den-
UNSM 51638, 85362, 85363, 85371, 85373, 85374, tition in Hypolagus voorhiesi resemble those of other
85376, 85387, 85389, 85390, 85395, 91121, 91137. species of Hypolagus. In size, it is significantly smaller
than in all species of the genus, except H. tedfordi, H. Lepus, 7 species, N=170
furlongi, and H. edensis, which are of comparable size.
PER on P3 is crenulated in 48 per cent of observed m5- Sylv//ogus, 9species,N=251
mm.
specimens and inflected from 43 to 66 per cent across
the occlusal surface, averaging 57 per cent. AER is
crenulated in 40 per cent of observed specimens and
extends across the occlusal surface from 17 to 27 per
cent, averaging 20 per cent. PER is deflected from a 4- 0
line perpendicular to a line of orientation (Fig. 2) 15
degrees anteriorly to 2 degrees posteriorly, averaging - 0
6 degrees anteriorly. The innermost part of PER turns

abruptly anteriad. 0 e 0 Te
4.Q
Remarks--UMMP V31955 is a left P3 from Deer
Park, which Hibbard (1956) referred to Hypolagus sp.;
and UMMP V60633A is a left P3 from White Rock, 3 -0 Lepoides lepoides
which Eshelman (1975) referred to Hypolagusfurlongi. S0 0 Hypologus ringoldensis
Both specimens are here identified as H. voorhiesi be- Se H oregonensis
cause in both, the innermost PER is abruptly deflected
0 H gidleyi
anteriorly.
Q H. vetus
2-
?HYPOLAGUS cf. . VOORHIESI
1 5 20 25 mm
UCMP 78497 is a right P3 from the Barstow local diastema: 3 I - P
fauna, Barstovian (middle Miocene). Morphologically
FIGURE 9. Scatter diagram with the anteroposterior length
this specimen (Fig. 70, P) resembles the P3's ofofH.
P3(APL P3) plotted against I-P3 diastema. The two ovoid
voorhiesi, but differs in having a thickened TH. circles
Al- inclose dots of extant species of Sylvilagus and Lepus
though it occurs with a series of specimens of H. fon-
(N = number of specimens), which were erased. The dots
tinalis, its morphology is quite different from the latter,
representing species of archaeolagines are plotted for com-
parison with the two extant genera.
a difference in magnitude that suggests it is not refer-
able to H. fontinalis. If it is H. voorhiesi, it would
represent an enormous extension of range for this
species.
HYPOLAGUS ARIZONENSIS Downey LEPOIDES new genus

Hypolagus arizonensis Downey, Journal of Paleontol- Type Species -Lepoides lepoides new species.
ogy 36(5):1112, September 1962. Diagnosis - Significantly larger in size than all other
species of archaeolagines and near in size to Lepus
Holotype - Incomplete right mandible with I-P3,
arcticus, P2 with three anterior reentrants, AR on P3,
UALP 1013A from UALP locality 15-10, Tusker local
diastema I-P3 longer than in other archaeolagines, pro-
fauna. Cochese County, Arizona.
portions of mandible resemble comparable structures
Emended Diagnosis--Hypolagus arizonensis differs in jack rabbits of the genus Lepus. PER is strongly
from H. regalis by its significantly smaller size with no
deflected posteriorly.
overlap of observed ranges. From H. voorhiesi it differs
Etymology - Lepoides, Latin, Lepus, hare, and Greek,
in a more strongly developed anterior deflection of
adjectival suffix -oides, as in.
PER on P3, and more deeply inflected AER. From all
other species of Hypolagus it differs in a strong anterior
LEPOIDES LEPOIDES new species
deflection of PER and abrupt anterior deflection of
innermost PER.
(Figs. 8A, C, D, 7A)
Referred Specimen - This species is known from Holotype the - UNSM 91196, incomplete mandible with
holotype and UALP 1013B, a mandible with I-P3. P3-M2, from late Hemphillian sediments in an un-
Geologic Age and Distribution - Late Blancan of Ar- named formation, Santee local fauna, in channel fill
izona (Fig. 12). near Santee, Knox County, Nebraska.
Description -The P3 in UALP 1013A, the holotype, Diagnosis--Same as for genus.
has an AER with crenulations and TN that is slightly Referred Specimens--Asterisk (*) denotes the num-
crenulated. PER is strongly deflected anteriorly and ber of uncataloged specimens at a locality. Blancan.
the innermost PER is abruptly deflected anteriorly. Panaca--incomplete palate, AMNH(FAM) 108718,
The P3 in UALP 1013B is as in the holotype but AER 108720, 108727, 108728; mandible, AMNH(FAM)
has only tiny crenulations. The P4 is characteristic of 108721, 108730; P3, AMNH(FAM) 108716, 108717,
that in other archaeolagines but has slight crenulations 108719, 108722, 108725, 108731.
on the thin enamel. Hemphillian. Santee -mandibles: UNSM, 1*, 91197,
.3 .2 .1 o .1 .2 .3
Lepus. This suggests that perhaps L. lepoides is a jack
Length of cranium p; rabbit ecomorph.
Lindsay (personal communication) suggested that
Bony palate to basioccipital perhaps Lepoides evolved into Lepus or Sylvilagus. In
Lepoides there is no trace of an internal reentrant on
Width of choana P3, yet PER extends 51 to 65, averaging 55 per cent
of occlusal width, while in Nekrolagus and Pratilepus,
which have PIR's, PER extends across the occlusal
Zygomatic breadth surface 55 (N = 7) and 54 (N = 19) per cent respec-
tively, percentages that are not statistically different
Diastema 12- P2 from those in Lepoides. There is no evidence that the
F
A B C Lepus-pattern of Hibbard (1963) ever develops by the
Maxillary tooth row(alveoli) D G KH simple extension of PER across the occlusal surface of
P3, but there is evidence that the nekrolagus-pattern
evolved into the Lepus-pattern.
Width of I
Genus PEWELAGUS
Anteroposterior length of R
Pewelagus White, Carnegie Museum of Natural His-
Diastema I - P G) H tory, Special Publication 9:47, 25 September 1984.
Type species Pewelagus dawsonae.
Mandibular tooth row (alveoli)
Emended Diagnosis--Size small, approximately as
in Hypolagus voorhiesi and Sylvilagus bachmani; tym-
A. Pewe/lgus dowsonae E. H vetus
panic bullae proportionally and actually larger than in
B. Sylvilogus bochmoni F S.foramen
any known leporid; floridanus
lacerum laterally dis-
C.Hypologus edensis G. Lepoides lepoides placed; basicranial-basifacial axis as in Sylvilagus;
D. H. gidleyi H. Lepus arcticus choana narrow as in Ochotona princeps; P2 ovoid in
cross section with MAR inflected at medial half and
FIGURE 10. Ratio diagram comparing measurements extendingofposteriad; EAR shallow or absent; PER on
the skull in species of archaeolagines with Sylvilagus flo-
P3 extends 36 to 55 per cent across occlusal surface,
ridanus (standard), S. bachmani, and Lepus arcticus.
and AER The
extends 20 to 36 per cent across occlusal
stippling was included to facilitate comparisons. A, C, D, G
surface, with two or more crenulations.
are known from single skulls. B, E, F, and H are samples
with means. Geologic Age and Distribution - Blancan of Arizona,
California, Nevada, Texas, and Guanajuato, Mexico.
PEWELAGUS DAWSONAE White

(Fig. 7R)
91199; P3, UNSM 91140, 91146, 91148, 91150, 91153,
Pewelagus dawsonae White, Carnegie Museum of Nat-
91156, 91158, 91161, 1080-75, 1090-74, 2020-73; P2,
ural History Special Publication 9:47, 25 Septem-
UNSM 85362-85377, 91152, 91160, 91162, 91163, ber 1984.
91167-91170, 91172, 91174-91176, 91178.
Geologic Age and Distribution--Late Hemphillian Holotype--A cranium lacking left tympanic bulla,
of Nebraska and mid Blancan of Nevada (Fig. 12). zygomatic arches, and anterior ends of nasals; LACM
Description - Specimens ofLepoides lepoides are the 24664, from LACM locality 6583, Upper Tapiado
largest known for the Archaeolaginae. The depth of the Wash, Anza-Borrego Desert State Park, San Diego
mandible below MI, length of the I-P3 diastema, and County, California, and 1,526 m (5,020 ft.) below top
the length of the mandibular tooth row at alveoli, all of section, Arroyo Seco local fauna, in the Palm Spring
fall within the ranges of comparable measurements of formation.
jack rabbits of the genus Lepus (Fig. 10). The enamel Emended Diagnosis--Pewelagus dawsonae is distin-
patterns of post-P2 and post-P3 dentitions are essen- guishable from ?Pewelagus mexicanus by its smaller
tially as in Hypolagus. The enamel pattern on P3 has size, smaller P2 with an uncrenulated MAR, and a TN
a strong posterior deflection of PER (Fig. 2) and strong- on P3 that is usually uncrenulated.
ly incised AR. AER is crenulated in 13 per cent, and Referred Specimens --Blancan. Vallecito Creek-
PER in 52 per cent of observed specimens. There are right mandible with P3-P4 and calcaneum, LACM
three reentrants on P2 with MAR the deepest, IAR 40146; right mandible with P3, IVCM 308. Arroyo
next deepest, and EAR the shallowest. P2 closely re- Seco-left mandible with DP3, M1, LACM 19266; left
sembles the P2 in Lepus. P3 and fragments, LACM 13742; skull with left man-
Remarks--The proportions of the mandible in L. dible (immature), LACM 16833; right mandible with
lepoides are similar to those in jack rabbits of the genus P4-M3, LACM 13745; rostral fragment with I'-I2 and
Hemingfordian Barstovian Clarendonian Hemphillian Blancan
PHypologus
Sopochensis Pewelagus
dowsonoe
?Pewelogus mexiconus
v oorhesi
Hypologus _ Hypologus
f. voorhiesi H reglis
H. tedfordi
H. orionensis
) I~ HH edensis
rchoeHore Hfoninsiis
H p or vip/cH H furlongi
H ringoldensis
FIGURE 11. Diagram of a suggested phylogeny of North A

leading to species refer to structures characterizing each sp
AER devoid of cement. 3. AER with cement. 4. AER usually uncrenulated. 5. AER crenulated. 6. AER crenulated in 40 per
cent of specimens. 7. AER deeply incised. 8. AER smooth-sided. 9. P3 and P4 incompletely molariform. 10. PER strongly
deflected anteriorly. 11. PER proportionally shallow. 12. PER deeply incised. 13. PER deflected posteriorly. 14. Innermost
maxillary fragment with P2_M3, LACM 23228; left doming of parietals, is nearest Sylvilagus auduboni,
mandible with P3-M1, LACM 13744; left mandible and far greater than in Romerolagus, Paleolagus hyp-
with P3-M2, LACM 23050; left mandible with P3-M,, sodus, and Ochotona."
LACM 23432; skeletal fragments, left maxilla with P3- The P2 is essentially as in Hypolagus and Brachy-
M2, mandible with P3-M1, proximal epiphysis of tibia, lagus with an ovoid cross section, a deep and smooth-
and fragment of right innominate, LACM 39458; left sided MAR, and a shallow or absent EAR. The internal
mandible with P3, LACM 23439; right mandible with reentrants on P3_M2 have crenulations on the enamel
P3-M2, LACM 13728; fragmentary cranium, LACM of the anterior walls which become progressively re-
17650; left P3, LACM 19562; left mandible with P3- duced toward the back of the tooth row. Crenulations
M2, LACM 24970; right P3, LACM 23174. Layer on the posterior walls of the reentrants are almost a
Cake-skull with right mandible (immature), LACM sent. M3 is roughly oblong in cross section with th
17671. Beck Ranch-left P3, MUBD 12036, 12037; enamel on the anterior and anterolabial sides of the
right P3, MUBD 12038. Coso Mountains-right P3, tooth.
LACM(CIT) 1978. Panaca-left P3, AMNH(FAM) AER on P3 extends from 26 to 41 per cent of occlusal
width, averaging 33 per cent, and PER extends from
108709. Red Corral-left P3, WTSU 4468A. Blanco--
right P3, TTU 6151. 42 to 54 per cent, averaging 50 per cent, respectively.
Geologic Age and Distribution - Blancan of Califor- AER is crenulated and TN is crenulated in 63 per cent
nia, Nevada, and Texas. of observed specimens.
Description--From White (1984:48-49): "Skull. Remarks - The P3 (TTU P615 1) and P2 (TTU P6150)
Tympanic bullae relatively large and laterally com- from the Blanco local fauna which Dalquest (1975)
pressed; their posterior ends extend slightly posteriad referred to Hypolagus sp. are referable to Pewelagus
of occipital condyles separating paroccipital process dawsonae because of the morphological similarity to
from squamosal process of squamosal; anterior ends the latter species.
extend almost into pterygoid fossae. Wall of bulla sim-
ple, without sponge-like structures present in Scalopus, ?PEWELAGUS MEXICANUS
Ochotona, Perognathus, Thomomys, Geomys, and (Miller and Carranza-Castafieda)
Mustela, and without mastoid portion delimited by a (Fig. 7S)
septum as in Dipodomys. Basisphenoids and basioc-
cipitals constricted between bullae, suture between them Hypolagus mexicanus Miller and Carranza-Castafieda,
Journal of Vertebrate Paleontology 2:97, May
posteriad of anterior ends of bullae. Most of left bulla 1982.
missing in holotype but ectotympanic present and su-
tured to basioccipital and posterior portion of basi- Holotype - From Miller and Carranza-Castafieda,
sphenoid. Immediately posterior to level of cavernous 1982. Separate teeth belonging to a single individual;
foramen, entotympanic, following contour of bulla, di- outer and inner upper incisors, P2, three upper molar-
verges laterally to basisphenoid, leaving a slit-like iform teeth, lower incisor, P3 and a lower molariform
opening between the two bones, which is here inter- tooth, IGCU 4158, from unnamed deposits of early
preted as representing the pharyngeal opening of the Blancan age, 2.5 km east of Rancho Viejo and 2 km
Eustachian tube. Foramen lacerum situated laterad of south of Tierra Blanca, Guanajuato, Mexico.
anterior end of bullae and medioposteriad of lateral Emended Diagnosis. - ?Pewelagus mexicanus differs
lamina of pterygoid fossa. Orbital fissure and optic from P. dawsonae in its larger size, more deeply incised
foramen as in other leporids, as is anterior half of AER on P3, and crenulations in MAR on P2.
cranium. Fenestrations on side of rostrum as in other Geologic Age and Distribution - Early Blancan of
leporids, but those in basicranial region almost absent, Guanajuato, Mexico.
probably reflecting the forward extension ofbullae, and Description - From Miller and Carranza-Castaneda
consequent lateral displacement of foramen lacerum (1982:97). "The upper first incisor possesses a narrow,
from its normal median position. Interparietal visible; shallow groove which lies slightly internal to the mid-
supraoccipital small as in Brachylagus. External au- line of the anterior surface. A short bevel with a distinct
ditory meatus enlarged and directed dorsoposteriad. bench marks the occluding surface of this tooth. P2 has
Palatal bridge extends from middle of M' to middle a well-developed reentrant which reaches a little more
of P2. than one-half the anteroposterior surface and is di-
The 'pistol grip'-like flexion of the posterior end of rected posteroexternally. A shallow groove exists be-
skull as expressed by the basicranial-basifacial axis and tween the reentrant and the labial border of the tooth.
4---
PER abruptly turned anteriorly. 15. TH usually sigmoid. 16. TH usually straight. 17. Trigonid relatively enlarged. 18. A
present. 19. Huge tympanic bullae. 20. I-P3 diastema elongated. 21. AIR and/or present in adults. 22. AIR and PIR absent
in adults. 23. P2 with MAR only. 24. P2 with MAR and often EAR. 25. P2 with MAR, EAR, and IAR. 26. Choana narrow as
in Ochotona. 27. PER less anteriorly deflected than in H. regalis and H. arizonensis. 28. Size-range with no overlap with that
in H. regalis. 29. Shallow AR in 12 per cent and TN crenulated in 70 per cent of specimens.
The three upper molariform teeth have internal reen- H. arizonensis, known from two adult individuals, is
trants that extend labially from a little more than half morphologically similar to H. regalis but differs in its
to nearly three-fourths the tooth width. All three teeth very small size. The observed ranges of the two species
possess strong crenulations on the posterior border. P3 do not overlap. H. voorhiesi differs from the other two
is heart-shaped in occlusal view. Reentrants exist only species in a less well-developed anterior deflection of
on the external surface of tooth. The anteroexternal PER, and a less well-incised AER on P3. The first ap-
reentrant is well-developed, narrow anteroposteriorly pearance of a specimen with a strongly anteriorly de-
and extends more than one-third of the anterior grind- flected PER is in the Barstow local fauna in a specimen
ing surface. Distinct crenulations mark the posterior tentatively referred to H. voorhiesi (Fig. 7P).
enamel border. The posteroexternal reentrant is di-Hypolagus oregonensis and H. ringoldensis make up
rected lingually and reaches more than half way across the H. oregonensis group, which is characterized by
the posterior occlusal surface. Its posterior enamel the bor-presence of an AR on P3. H. oregonensis has a
der also shows distinct crenulations. In the lower mo- sigmoid, and H. ringoldensis a straight TH on P3. The
lariform tooth the reentrant traverses the entire occlu- latter species seems derivable from the former as sug-
sal surface. The posterior border of the reentrant is gested by Gustafson (1978).
moderately crenulated. In the monotypic genus Lepoides there is an AR on
IGCU 4075, a P3, closely resembles the P3 of the P3, PER is strongly deflected posteriorly, and P2 has
holotype. The only difference observed is in the pos-three reentrants. The presence of an AR in this genus
terior borders of both the anteroexternal and postero- may relate it to the H. oregonensis group, assuming
external reentrants. In this specimen the anteroexternalthat an AR appeared only once in the archaeolagines.
reentrant has but a single crenulation on its posteriorHibbard (1963) argued that an AR appeared only once
border (about the same magnitude as those in the P3 in the Leporinae.
holotype) and the posterior border of the posteroex- The enormous tympanic bullae and related changes
ternal reentrant is wavy, not crenulated." of the cranium of Pewelagus makes this a most dis-
AER extends from 40 to 41 per cent and PER ex- tinctive genus of the Archaeolaginae.
tends from 60 to 61 per cent of the occlusal width of Archaeolagus (sensu lato) is here viewed as the base
P3. from which the various lines of archaeolagines are de-
Remarks-The skull of Pewelagus dawsonae is so rived (see Fig. 11). As shown by Dawson (1958), the
markedly different from the skulls of other leporids Hypolagus vetus and H. oregonensis groups are deriv-
that until the skull of?P. mexicanus is known, the latter able from known species of Archaeolagus. The H. re-
is tentatively referred to Pewelagus. galis group, Pewelagus, and Lepoides are not derivable
from any known species of Archaeolagus. Light might
be shed on this problem with a more detailed revision
DISCUSSION of Archaeolagus.
The peak of archaeolagine diversity was reached in
The Archaeolaginae treated in this paper midare pre-
Blancan times (Hagerman, Taunton, Panaca, Beck
sumed to be derived from Archaeolagus Ranch), as proposed
and the entire subfamily became extinct at the
by Dawson (1958), and are segregated into endthree gen-
of Blancan times. At that time there were 11 species
era: Hypolagus, Lepoides, and Pewelagus. of archaeolagines and 14 species of leporines, and of
Within Hypolagus there are three species groups:
the 14 leporines, only Lepus sp. (from Big Springs and
The H. vetus group, H. regalis group, and H. orego-
Borchers), Aztlanolagus, Aluralagus, and Sylvilagus
nensis group. hibbardi survived into the Irvingtonian. Thus 21 species
The Hypolagus vetus group is made up of ofthespecies
Leporidaeinbecame extinct by the end of Blancan
which the PER on P3 is not strongly deflected anteriorly
times (11 species of archaeolagines and 10 species of
or posteriorly, and lacks an AR. The species in this leporines). If the extinction of the archaeolagines was
group are: H. vetus, H. gidleyi, H. parviplicatus, H. caused by competition with the leporines, why did an
edensis, H. tedfordi, H. furlongi, and H. fontinalis. This almost equal number of archaeolagines and leporines
group is subdivided into primitive species in which become extinct in the Blancan? It may be that the
PER is less well-developed: H. tedfordi, H. fontinalis, leporine species that became extinct were, as for the
and H. parviplicatus, and two derived groups: a group contemporary archaeolagines, presumably not so "ad-
of small-sized individuals that includes H. edensis, in
vanced" as the species of Sylvilagus and Lepus. In the
which AER is deeply incised and smooth-sided, and
Anza-Borrego Desert section, Hypolagus vetus died out
H. furlongi, in which AER is weakly incised and TH
about 1400 m below the top of the section, and Syl-
is straight; and a group of large-sized individuals, which
include H. vetus in which TH is sigmoid, and H. gidleyi, vilagus hibbardi makes its first appearance 1100 m
in which TH is straight. below the top of the section. This was interpreted by
The Hypolagus regalis group is characterized by a White (1984) as indicating an ecological replacement
strong anterior deflection of PER, and the innermost ofH. vetus by S. hibbardi. The skull proportions in the
part of PER is abruptly deflected anteriorly. This group two species are remarkably similar.
includes H. regalis, H. arizonensis, and H. voorhiesi. Clearly identifiable specimens ofarchaeolagines are,
So
B, r (nHLOCAL
insN
S0 Ninefoot
SBig Rapids, ID
Sproadwaterings, NE
0 0 ' Grand View, ID
< OW Elk Hills, CA Z- t
E Hogermanfoot Rapids, E
White Rock, KS
Blanco, TX
N Tusker,
FlatAZ iron Butte, ID
Flat Cit
Iron Butte, ID Canyon,T
Cita Canyon,TX
Broadwater, NE
Coso Mountains,CA
z A Sand Point, I D
1 BRedC or ra,TX
CoBenson,NAZ
DPanaca, NVE
E Hagerman, ID D- - ET T
MoTaunton, W C
Deer iPark, KS
Fox Canyonle Tuff, KSKS
Rexroad loc.3,KS
Del Valle, AZ
Ranchoffee Ranch, GT
Truth or
Consequences,NM
White Bluffs, WA
McKay Reservoir, OR
Mount Eden,
G Christmas Volley,CA
OR .
Ainpachle TufCf, C A
Wikieup, AZ K J K K
I White Cone, AZ
Coffee Ranch, TX
SBig SpRedington, AZ SD-
a1
W
Rome, OR P
a Thousand Creek, NV-P--
Ricardo, CA P
b Apache Canyon, CA-P
Fish Lake Valley, NV-P
wr Big Spring Canyon, SD-P
4t
Of-J I
FIGURE 12. Diagram plotting the stratigraphic occurrence of species of Hypolagus, Lepoides, and Pewelagus. The vertical
lines represent stratigraphic ranges. The dots mark the stratigraphic position of each local fauna per species, and the horizontal
small lines represent the occurrence of specimens in the Anza-Borrego Desert section. The letters next to the vertical lines
correspond to the letters to the left of the names of local faunas sharing stratigraphic positions.
up to now, unknown from the mid-Hemphillian, and REFERENCES
only two species, H. tedfordi and H. vetus, are known
Alvarez, T. 1963. Nueva especie de Archaeolagus (Lepor-
from below and above the mid-Hemphillian. The bio-
idae) basada en restos procedentes de Sonora, Mexico.
stratigraphic use of the archaeolagines is primarily to Acta Zoologica Mexicana 6(5): 1-4.
be found in the late Hemphillian and Blancan species. Barnosky, A. D. (in press). Arikareean, Hemingfordian, and
Barstovian mammals from the Coulter Formation, Jack-
Acknowledgements -The writer gratefully acknowl-
son Hole, Wyoming. Carnegie Museum of Natural His-
edges the benefits of discussions with a number of col-
tory, Bulletin 26.
leagues, especially with Everett H. Lindsay, Charles A. Baskin, J. A. 1979. Small mammals of the Hemphillian
Repenning, Richard H. Tedford, Richard J. Zakrzew- age White Cone local fauna, northeastern Arizona. Jour-
ski, Michael R. Voorhies, and Gerald E. Schultz. Rich- nal of Paleontology 53(3):695-708.
ard Estes, Mary R. Dawson, Anthony D. Barnosky, Brown, B. 1908. The Conard Fissure, a Pleistocene bone
and S. David Webb constructively reviewed the manu- deposit in northern Arkansas; with descriptions of two
script and contributed greatly to its improvement. For genera and twenty new species of mammals. American
the privilege of studying specimens in their care, the Museum of Natural History, Memoir 9:155-208.
writer thanks William A. Akersten, Idaho Museum of Campbell, K. E., Jr. 1969. Comparing postcranial skeletons
Natural History; Sankar Chatterjee, The Museum, of Pliocene rabbits. Michigan Academician, Papers,
Michigan Academy of Science, Arts, and Letters 1:99-
Texas Tech University; William A. Clemens and J. H. 115.
Hutchison, Museum of Paleontology, University of Cassiliano, M. 1980. Stratigraphy and vertebrate paleon-
California, Berkeley; Walter W. Dalquest, Biology De- tology of the Horse Creek-Trail Creek area, Laramie
partment, Midwestern University; Philip D. Ginger- County, Wyoming. Contributions to Geology, Univer-
ich, Museum of Paleontology, University of Michigan; sity of Wyoming 19(1):25-68.
Everett H. Lindsay, Department of Geosciences, Uni- Clark, J. B., Dawson, M. R. and Wood, A. E. 1964. Fossil
versity of Arizona; Thomas Logan, Museum of South- mammals from the lower Pliocene of Fish Lake Valley,
west Biology, University of New Mexico; Philip Bjork Nevada. Bulletin, Museum of Comparative Zoology 131:
29-63.
and James E. Martin, Museum of Geology, South Da-
kota School of Mines and Technology; Jason A. Lil- Dalquest, W. W. 1975. Vertebrate fossils from the Blanc
local fauna of Texas. Occasional Papers, The Museum,
legraven, Museum of Geology, University of Wyo-
Texas Tech University 30:1-52.
ming; Larry D. Martin, Museum of Natural History, 1978. Early Blancan mammals of the Beck Ranch
University of Kansas; George J. Miller, Imperial Val- local fauna of Texas. Journal of Mammalogy 59(2):269-
ley College Museum; William Orr, Oregon State Mu- 298.
seum of Anthropology; Clayton E. Ray, National Mu- Dawson, M. R. 1958. Later Tertiary Leporidae of North
seum of Natural History; John M. Rensberger, Burke America. University of Kansas Paleontological Contri-
Museum, University of Washington; Charles A. Re- butions: Vertebrata 6:1-75.
penning, United States Geological Survey, Denver, Dice, L. R. 1929. The phylogeny of the Leporidae, with
Colorado; Gerald E. Schultz, Geology Department, description of a new genus. Journal of Mammalogy 10(4):
340-344.
West Texas State University; Richard H. Tedford,
American Museum of Natural History; Michael R. Downey, J. S. 1962. Leporidae of the Tusker local f
from southeastern Arizona. Journal of Paleontology
Voorhies, University of Nebraska State Museum; S.
36(5):1112-1115.
David Webb, Florida State Museum, University of
Eshelman, R. E. 1975. Geology and paleontology of the
Florida; David P. Whistler and Theodore Downs, Los
early Pleistocene (Late Blancan) White Rock fauna from
Angeles County Museum of Natural History; and north-central Kansas. Claude W. Hibbard Memorial
Richard J. Zakrzewski, Sternberg Memorial Museum, Volume 4, University of Michigan, Papers on Paleon-
Fort Hays Kansas State University. tology 13:1-60.
Wade E. Miller, Geology Department, Brigham Galusha, T. and Blick, J. C. 1971. Stratigraphy of the San
Young University, kindly made available to the writer F6 Group, New Mexico. Bulletin, American Museum o
camera lucida drawings of the teeth of ?Pewelagus Natural History 144(1):1-127.
mexicanus. Gazin, C. L. 1930. A Tertiary vertebrate fauna from the
upper Cuyama drainage basin, California. Carnegie In-
The stratigraphic data in Figure 12 were obtained
by the writer and in consultation with a number stitution
of of Washington, Publication 404:55-76.
colleagues, especially E. H. Lindsay, C. A. Repenning, 1934. Pliocene hares from Idaho. Proceedings,
G. E. Schultz, and M. R. Voorhies, and by reference United States National Museum 83(2976):111-121.
to Repenning (in press) and Lindsay, Johnson,Gustafson,
and E. P. 1978. The vertebrate faunas of the Plio-
cene Ringold formation, south-central Washington. Bul-
Opdyke (1975).
letin, Museum of Natural History, University of Oregon
The writer gratefully acknowledges the use of facil-
23:1-62.
ities and services of Idaho Museum of Natural History,
Hall, E. R. 1981. The mammals of North America. Seco
Department of Geosciences, University of Arizona,
edition. John Wiley and Sons, 1181 pp.
and the Florida State Museum, University of Florida. Hibbard, C. W. 1939. Four new rabbits from the upper
Special thanks are due to Mr. and Mrs. E. Hadley Pliocene of Kansas. American Midland Naturalist 21:
Stuart, Jr. for their financial support, without which 501-513.
this research would have been impossible. 1956. Vertebrate fossils from the Meade formation
of southwestern Kansas. Papers, Michigan Academy of Cenozoic mammals: their temporal record, biostratig-
Science, Arts, and Letters 41:145-203. raphy, and biochronology. University of California Press,
1963. The origin of the P3 pattern of Sylvilagus, Berkeley.
Caprolagus, Oryctolagus, and Lepus. Journal of Mam- Shotwell, J. A. Hemphillian mammal assemblage from
malogy 44(1):1-15. northeastern Oregon. Bulletin, Geological Society of
1969. The rabbits (Hypolagus and Pratilepus) from America 67:717-738.
the Upper Pliocene, Hagerman local fauna of Idaho. Stevens, M. S., Stevens, J. B. and Dawson, M. R. 1969.
Michigan Academician, Papers of the Michigan Acad- New early Miocene formation and vertebrate local fau-
emy of Science, Arts, and Letters 1:81-97. na, Big Bend National Park, Brewster County, Texas.
Lindsay, E. H. 1972. Small mammal fossils from the Bar- Pearce-Sellards Series, Texas Memorial Museum 15:1-
stow formation, California. University of California 53.
Publications in Geological Sciences 93:1-104. Storer, J. E. 1975. Tertiary mammals of Saskatchewan, Part
- , Johnson, N. M. and Opdyke, N. D. 1975. Prelim- III: the Miocene fauna. Life Sciences Contributions,
inary correlation of North American land mammal ages Royal Ontario Museum 103:1-134.
and geomagnetic chronology. Claude W. Hibbard Me- White, J. A. 1984. Late Cenozoic Leporidae (Mammalia,
morial Volume 3, University of Michigan, Papers on Lagomorpha) from the Anza-Borrego Desert, southern
Paleontology 12:111-119. California. Special Publication, Carnegie Museum of
MacFadden, B. J., Johnson, N. M. and Opdyke, N. D. 1979. Natural History 9:41-57.
Magnetic polarity stratigraphy of the Mio-Pliocene Wilson, R. W. 1937. A new genus of lagomorph from the
mammal-bearing Big Sandy formation of western Ari- Pliocene of Mexico. Southern California Academy of
zona. Earth and Planetary Science Letters 44(1979):349- Science, Bulletin 36:98-104.
364. 1939. Rodents and lagomorphs of the late Tertiary
Miller, W. E. 1980. The late Pliocene Las Tunas local fauna Avawatz fauna, California. Carnegie Institution of
from southernmost Baja California, Mexico. Journal of Washington Publication 514:31-38.
Paleontology 54(4):762-805. Wood, A. E. 1937. Additional material from the Tertiary
and Carranza-Castafieda, 0. 1982. New lago- of the Cuyama Basin of California. American Journal
morphs from the Pliocene of central Mexico. Journal of of Science 33(193):29-43.
Vertebrate Paleontology 2(1):95-107.
Repenning, C. A. (in press). Biochronology of the microtine
rodents of the United States. In Woodburne, M. O. (ed.), Received 29 November 1985; accepted 3 January 1987.
TABLE 1. Cranial measurements in mm of Hypolagus, Lepoides, and Pewelagus.
H. H. H. rin-
Hypolagus H. fur- eden- orego- golden- H. Lepoides Pewelagus
vetus H. gidleyi longi sis nensis sis regalis lepoides dawsonae
Greatest length of crani- X 71.8 76.0 - 67.0 - 57.7

um s - - -- - - -
OR - 62.9-78.4 -
N 1 3 - 1 - - 1
Condylopremaxillary X - 68.2 - - - - - - -
length of cranium s - -
OR - - - - - - - - -
N - 1 - - - -
Length of anterior pala- X 19.2 18.0 - 15.5 - - - - 12.9

tine foramen s - - - -
OR 18.3-19.8 - - - - - - - -
N 3 1 - 1 - - - 1
Width of anterior pala- X 6.6 5.8 - 6.3 - 8.4 - - 4.3

tine foramen s 0.884 - - - -
OR 5.3-8.0 - - - - - - - -
N 6 1 - 1 - - - 1
Palatal length X 5.3 7.6 4.9 5.5 7.0 7.3 6.4 9.0 4.4
s 0.639 - - - - - - - -
OR 4.5-6.8 - - - - - 5.1-7.6 - -
N 16 1 1 1 2 1 2 1 1
Palate to basioccipital X 22.9 21.4 - 20.4 - - - - 17.8

5 - - - - - - - - -
OR 22.7-23.2 -
N 2 1 - 1 - - - 1
TABLE 1. (Continued)
H. H. H. rin-
Hypolagus H. fur- eden- orego- golden- H. Lepoides Pewelagus
vetus H. gidleyi longi sis nensis sis regalis lepoides dawsonae
Width of choana X 6.5 4.8 - 4.5 - - 7.2 - 3.5

s 1.03 - - - - - - - -
OR 5.1-7.6 - - - - - 6.5-7.8 - -
N 6 1 - 1 - - 2 - 1
Length of basioccipital X 10.3 10.9 - 10.1 - - - - 10.4
OR 9.3-12.0 -
N 3 1 - 1 - - - 1
Anteroposterior diame- X 12.2 10.5 - 13.3 - - - 18.8
ter of bulla s 1.09 - - -
OR 10.7-13.2 -
N 6 1 1 1
Width of bulla X 9.2 8.5 - 10.5 - - 10.1
s 1.50 - - - - - - - -
OR 7.6-10.7 - - - - - - - -
N 6 1 - 1 - - - 1
Distance between bullae X 6.8 7.3 - 7.0 - 4.0
S - - - - - - - - -
OR 6.1-9.6 -
N 3 1 - 1 - - 1
Width across occipital X 12.1 14.9 - 12.5 - 10.0

condyles s - - - -
OR 10.7-13.0 - - - - - - - -
N 4 1 - 1 - - - 1
Least postorbital con- X 12.1 14.4 - 10.8 - 10.5

striction s - - - -
OR - - - - - - - - -
N 1 1 - 1 - - - 1
Depth anterior end zy- X 5.0 5.9 4.2 5.8 6.1 6.5 7.1 7.0 4.2
gomatic arch s 0.639 - - - - - - - -
OR 4.1-5.7 - - - - - - - -
N 9 1 1 1 2 1 1 1 2
Skull width anterior end X 31.7 35.0 - 30.7 - - 37.8 - 24.5
zygomatic arch s 2.98 - - - - - - - -
OR 26.9-35.8 -
N 6 1 - 1 - - 1 - 1
Width across supraorbi- X 21.4 18.5 -

tal ridges s - -
OR 20.0-22.7 - - - - - - - -
N 2 1 - - - -
Diastema 12_p2 at alveo- X 18.8 19.5 - 17.1 - - - - 14.6

li s - - - -
OR 15.7-22.7 19.5 - - - - - - 13.7-15.5
N 3 1 - 1 - - - 2
Alveolar length I2_p2 X 14.0 15.9 12.9 12.1 15.6 - 15.6 17.4 10.4
s 1.06 - - - - - - - -
OR 12.4-15.5 - - - - - - 16.9-17.8 9.2-11.9
N 13 1 1 1 1 - 1 3 3
Anteroposterior length X 1.2 2.0 - 1.7 - - - 1.2
ofI' s 0.285 - - - - - - - -
OR 1.4-2.3 - - - - - - - 1.0-1.4
N 10 1 - 1 - - - - 2
Width of I' X 2.6 2.9 - 2.3 - - - - 1.9

s 0.326 - - - - - - - -
OR 2.0-3.0 - - - - - - - 1.7-2.1
N 12 1 - 1 - - - - 2
TABLE 2. Mandibular measurements in mm of species of Hypolagus.
H. par- H.
Hypolagus vipli- H. H. H. H. H. orego-
vetus catus gidleyi fontinalis tedfordi furlongi edensis nensis
Anteroposterior X 3.1 3.2 3.3 2.8 2.6 2.6 2.6 3.2
length of P3 s 0.25 - 0.177 0.09 0.14 0.12 0.18 0.16
OR 2.3-3.6 - 3.0-3.6 2.6-2.9 2.2-2.8 2.4-2.8 2.0-2.9 3.0-4.0
N 94 1 38 17 39 18 3 14
Width of P3 X 2.8 2.5 3.0 2.4 2.3 2.3 2.3 3.0

s 0.18 - 0.20 0.11 0.18 0.16 0.19 0.16
OR 2.3-3.3 - 2.6-3.6 2.2-2.5 1.9-2.5 2.1-2.5 1.9-2.6 2.6-3.1
N 91 1 38 17 39 18 39 14
Length of I-P3 X 16.1 - 16.7 - - - 12.9 17.5

s 1.12 - 1.34 - - - - 1.11
OR13.9-18.0 - 14.1-18.6 - - - 12.5-13.1 15.7-18.8
N 12 - 8 - - - 3 10
Mandibular depth X 12.6 12.0 13.1 11.5 8.6 10.5 12.1 13.4
below M, s 1.11 - 0.50 - 0.24 - - 0.68
OR 11.4-14.4 - 12.4-13.9 11.1-12.1 8.3-9.0 - 10.8-13.4 12.0-14.8
N 6 1 9 3 7 1 2 15
Alveolar length X 15.1 13.1 16.6 - - 12.9 12.6 16.6

P3-M3 s 1.16 - 0.72 - - - - 0.80
OR 13.0-17.1 - 15.1-17.5 - - - 11.9-12.9 15.2-17.4
N 11 1 8 - - 1 3 6
TABLE 3. Mandibul
Hypolagus H. H. H. arizo- Lepoides Pewelagus ?P. mexi-

ringoldensis regalis voorhiesi nensis lepoides dawsonae canus
Anteroposterior length X 3.5 3.3 2.6 2.1 3.9 2.4 2.9
of P3 s 0.20 0.22 0.14 - 0.22 0.26 -
OR 3.0-4.0 2.7-3.6 2.2-2.9 - 3.5-4.3 2.0-2.8 2.8-3.0
N 60 43 24 2 21 17 2
Width of P3 X 3.2 2.9 2.3 2.0 3.4 2.1 2.7

s 0.23 0.28 0.16 - 0.18 0.26 -
OR 2.7-3.7 2.4-3.7 2.0-2.6 1.9-2.0 3.0-3.7 1.7-2.5 2.6-2.8
N 60 43 24 2 21 17 2
Length of I-P3 diastema X 17.9 16.3 12.3 - 21.7 11.8 -

s 1.11 - - - 1.24 - -
OR 16.6-19.5 16.1-16.7 11.2-13.4 - 20.5-23.3 11.4-12.1 -
N 7 3 2 - 5 2 -
Mandibular depth below X 13.8 13.7 11.2 - 15.2 - -

M, s - - - 1.19 - -
OR 13.7-14.0 - 10.7-11.6 - 14.2-17.0 - -
N 2 1 2 - 5 - -
Alveolar length P3-M3 X 16.9 16.4 13.0 - 19.3 11.2 -

S - - - - - - -
OR 16.6-17.2 - 12.5-13.5 - 18.7-19.9 - -

N 2 1 2 - 2 1 -
TABLE 4. List of local faunas discussed, given by state.
Arizona: Benson, Redington, Tusker, White Cone, Wikieup.

Baja California: Las Tunas.
California: Apache Canyon, Arroyo Pequefio, Arroyo Seco, Barstow, Coso Mountains, Del Valle, Elk Hills, Kern River, Layer
Cake, Mission Viejo, Mount Eden, Mulholland, Pechanga, Pinole Tuff, Radec, Ricardo, Quatal Canyon, Santa Margarita,
Trench Canyon, Vallecito Creek.
Florida: Lower Bone Valley, Upper Bone Valley.
Guanajuato: Rancho El Ocote, Rancho Viejo.
Idaho: Flat Iron Butte, Grand View, Hagerman, Ninefoot Rapids, Oreana, Railroad Canyon, Sand Point, Star Valley.
Kansas: Deer Park, Fox Canyon, Rexroad locality 3, Wendell Fox, White Rock.
Nebraska: Big Springs, Broadwater, Oshkosh, Quinn Canyon, Santee.
New Mexico: Albuquerque, Chamita, Truth or Consequences.
Nevada: Fish Lake Valley, Panaca, Stiener Ranch, Tonopah, Thousand Creek, Virgin Valley.
Oregon: Arlington, Christmas Valley, McKay Reservoir, Rome.
South Dakota: Big Spring Canyon, Todd County.
Saskatchewan: Wood Mountain.
Texas: Beck Ranch, Blanco, Cita Canyon, Coffee Ranch, Red Corral.
Washington: Taunton, White Bluffs.
Wyoming: Trail Creek Quarry.
ALFRED SHERWOOD ROMER PRIZE
The recipients of the 1987 Alfred Sherwood Romer Prize for the best student papers
of the Society of Vertebrate Paleontology are Stephen M. Gatesy (Museum of Comparative Zoology, Harvard
University) for his paper entitled "Dinosaur Limb Kinematics and Theropod Evolution" and Mark A. Norell
(Department of Biology, Yale University) for his paper entitled "The Phylogenetic Determination of Taxonomic
Diversity: Implications for Terrestrial Vertebrates at the K-T Boundary."

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The Archaeolaginae (Mammalia, Lagomorpha) of North America, Excluding Archaeolagus

THE ARCHAEOLAGINAE (MAMMALIA, LAGOMORPHA) OF NORTH AMERICA,

ABSTRACT-The Archaeolaginae of North America, excluding Archaeolagus and Panolax, include

© 1987 by the Society of Vertebrate Paleontology 425

were slightly different on comparable drawings of the

426 JVP 7(4), December 1987

JVP 7(4), December 1987 427

anterior AER PER

SYSTEMATIC PALEONTOLOGY zygomatic root more anterior to cheek teeth in the

428 JVP 7(4), December 1987

~·/''' ·~·' -L· '''.··

JVP 7(4), December 1987 429

430 JVP 7(4), December 1987

JVP 7(4), December 1987 431

432 JVP 7(4), December 1987

JVP 7(4), December 1987 433

434 JVP 7(4), December 1987

As noted by Dawson (1958), the mental foramen i

JVP 7(4), December 1987 435

436 JVP 7(4), December 1987

JVP 7(4), December 1987 437

JVP 7(4), December 1987 439

6 degrees anteriorly. The innermost part of PER turns

HYPOLAGUS ARIZONENSIS Downey LEPOIDES new genus

440 JVP 7(4), December 1987

PEWELAGUS DAWSONAE White

JVP 7(4), December 1987 441

FIGURE 11. Diagram of a suggested phylogeny of North A

442 JVP 7(4), December 1987

JVP 7(4), December 1987 443

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a Thousand Creek, NV-P--

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446 JVP 7(4), December 1987

TABLE 1. Cranial measurements in mm of Hypolagus, Lepoides, and Pewelagus.

Greatest length of crani- X 71.8 76.0 - 67.0 - 57.7

Length of anterior pala- X 19.2 18.0 - 15.5 - - - - 12.9

Width of anterior pala- X 6.6 5.8 - 6.3 - 8.4 - - 4.3

Palate to basioccipital X 22.9 21.4 - 20.4 - - - - 17.8

JVP 7(4), December 1987 447

Width of choana X 6.5 4.8 - 4.5 - - 7.2 - 3.5

Length of basioccipital X 10.3 10.9 - 10.1 - - - - 10.4

Width across occipital X 12.1 14.9 - 12.5 - 10.0

Least postorbital con- X 12.1 14.4 - 10.8 - 10.5

Width across supraorbi- X 21.4 18.5 -

Diastema 12_p2 at alveo- X 18.8 19.5 - 17.1 - - - - 14.6

Width of I' X 2.6 2.9 - 2.3 - - - - 1.9

448 JVP 7(4), December 1987

Width of P3 X 2.8 2.5 3.0 2.4 2.3 2.3 2.3 3.0

Length of I-P3 X 16.1 - 16.7 - - - 12.9 17.5

Alveolar length X 15.1 13.1 16.6 - - 12.9 12.6 16.6

Hypolagus H. H. H. arizo- Lepoides Pewelagus ?P. mexi-

Width of P3 X 3.2 2.9 2.3 2.0 3.4 2.1 2.7

Length of I-P3 diastema X 17.9 16.3 12.3 - 21.7 11.8 -

Mandibular depth below X 13.8 13.7 11.2 - 15.2 - -

Alveolar length P3-M3 X 16.9 16.4 13.0 - 19.3 11.2 -

OR 16.6-17.2 - 12.5-13.5 - 18.7-19.9 - -

JVP 7(4), December 1987 449

Arizona: Benson, Redington, Tusker, White Cone, Wikieup.

ALFRED SHERWOOD ROMER PRIZE

450 JVP 7(4), December 1987

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