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AMERICAN JOURNAL
OF
PHYSICAL ANTHROPOLOGY 93:323340 (1994)
New Early Eocene Anaptomorphine Primate (Omomyidae) From
the Washakie Basin, Wyoming, With Comments on the
Phylogeny and Paleobiology
of
Anaptomorphines
BLYTHE
A.
WILLIAMS
AND
HERBERT H.
COVERT
Department
of
Anthropology, University
of
Colorado, Boulder, Colorado
80309-0233
KEY
WORDS
Omomyidae, Anaptomorphine, Paleoprimatology,
Eocene
ABSTRACT
Recent paleontological collecting
in
the Washakie Basin,
southcentral Wyoming, has resulted in the recovery of over
100
specimens of
omomyid primates from the lower Eocene Wasatch Formation. Much of what
is known about anaptomorphine omomyids is based upon work in the Bighorn
and Wind River Basins of Wyoming. This new sample documents greater
taxonomic diversity of omomyids during
the
early Eocene and contributes to
our
understanding of the phylogeny and adaptations
of
some of these earliest
North American primates. A new middle Wasatchian (Lysitean) anaptomor-
phine,
Anemorhysis sauagei,
n. sp.,
is
structurally intermediate between
Teil-
hardina americana
and other species of
Anemorhysis
and may be
a
sister
group of other
Anemorhysis
and
Trogolemur.
Body size estimates
for
Anemorhysis, Tetonoides, Trogolemur,
and
Teilhar-
dina americana
indicate that these animals were extremely small, probably
less than
50
grams. Analysis of relative shearing potential of lower molars of
these taxa indicates that some were primarily insectivorous, some primarily
frugivorous, and some may have been more mixed feeders. Anaptomorphines
did not develop the extremes of molar specialization for frugivory
or
insec-
tivory seen in extant prosimians. Incisor enlargement does not appear to be
associated with specialization in either fruits
or
insects but may have been an
adaptation for specialized grooming
or
food manipulation.
0
1994
Wiley-Liss,
Inc.
Fossil primates of North America first oc-
cur in the earliest Eocene,
a
time period re-
ferred to
as
the Wasatchian Land Mammal
Age, approximately
56-51
million years ago.
These primates
are
usually placed in two
families, the Omomyidae and the Adapidae.
It
is commonly thought that the oldest and
apparently most primitive omomyid sub-
family is the Anaptomorphinae (e.g., Szalay,
1976;
Gingerich,
1981).
Therefore, recon-
struction
of
the phylogeny and behavior of
these early forms should help
us
to under-
stand basal primate paleobiology.
Much of what we know about early anap-
tomorphines
is
based upon research in the
Bighorn Basin (including the Clarks Fork
Basin) (e.g., Bown,
1974, 1976, 1979;
Gin-
gerich,
1981;
Bown and Rose,
1984, 1987)
and the Wind River Basin (e.g., Stucky,
1982, 1984;
Beard
et
al.,
1992)
of Wyoming.
In the Bighorn Basin,
latest
Wasatchian
(Lostcabinian) fossils
are
sparsely repre-
sented (Schankler,
1980;
Gingerich et al.,
1980;
Gingerich,
1991),
whereas in the Wind
River Basin
the
middle Wasatchian faunas
are poorly documented (Krishtalka et al.,
1987).
Received July 15,1992; accepted October
5,1993.
Address reprint requests
to
Blythe A. Williams, Department
of
Biological Anthropology and Anatomy, Campus
Box
3170, Duke
University Medical Center, Durham, NC 27710.
0
1994 WILEY-LISS,
INC
324
B.A.
WILLIAMS
AND
H.H.
COVERT
TABLE
1.
Primates known
from
the Wasatch Formation
of
the Washakie Basin. Wvomine'
Land mammal subage and biochron Omomyidae
Lostcabinian (Wa-7)
Lysitean (Wa-6)
Upper Graybullian (Wa-5)
Lower Graybullian (Wa-3-4)
Absarokius
cf.
abbotti
Trogolemur
myodes
Loueina minuta
Chlororhysis knightensis
c.f.
Chlororhysis
anaptomorphine sp. indet.
Anemorhysis sauagei,
n. sp.
Arapahouius gazini
Tetonoides pearcei
Steinius
sp.
Tetonius matthewi
anautomomhine n.
SD.
Adapidae
Cantius
cf.
uenticolus
Cantius frugiuorus
Copelemur australotutus
Copelemur tutus
Notharctus
cf.
robinsoni
Cantius
cf.
abditus
Copelemur australotutus
Cantius trigonodus
Copelemur praetutus
Cantius
cf.
mckennai
'
Faunal zones listed from youngest (Lostcabinian) to oldest (Lower Graybullian). Sources utilized for compilation
of
table include Gazin
(19621,
Savage and Waters
(19781,
Savage and Russell
(19831,
and collections at the University
of
Colorado Museum and University of California
Museum of Paleontology. Faunal zonations here and in Table
4
follow those summarized in Krishtalka
et
al.
(1987)
Sandcouleean-Blacksforkian).
Alternative zonations (Wa-l to Wa-7)
follow
Gingerich
(1989)
and
(Br-1
to Br-2)
Gunnel1
(1989).
Since 1987, the University
of
Colorado
Museum has been collecting early Eocene
fossils from near Bitter Creek Station and
Table Rock in the northwestern part
of
the
Washakie Basin, southcentral Wyoming.
These efforts have resulted in the recovery
of
over
3,000
mammalian specimens, includ-
ing approximately
300
jaws, isolated teeth,
and postcranial bones of primates. The new
material has been recovered from the Wa-
satch Formation and samples most of the
Wasatchian (Table
1).
To date, the primate
fauna from this basin is largely undescribed.
The first early Eocene anaptomorphine de-
scribed from near Bitter Creek was
Te-
tonoides pearcei
(Gazin, 1962), and
Arapa-
hovius gazini
was described 20 years later
by Savage and Walters (1978). Many new
specimens
of
these taxa (Covert and
Williams, 1991a; Williams et al., 1991;
Williams and Covert, 1992a,b) document
and clarify some aspects
of
their anatomy.
Abbreviations
CM Carnegie Museum
of
Natural History,
Pittsburgh,
PA
UCM University of Colorado Museum,
Boulder, CO
UM
University
of
Michigan Museum
of
Paleontology
USGS United States Geological Survey,
Denver,
CO;
usm
United States National Museum,
Washington, DC.
In this paper we
1)
provide a taxonomic
list of primates occurring in Wasatchian de-
posits
of
the Washakie Basin; 2) present an
emended generic diagonsis
for
Anemorhysis
and describe a new species of this taxon that
may be structurally transitional between
Teilhardina americana
and species of
Ane-
morhysis
and
Trogolemur; 3)
discuss the
phylogenetic relationships among these
anaptomorphines; and
4)
make suggestions
about their body size and dietary adapta-
tions.
MATERIALS AND
METHODS
Measurements
All
dental measurements taken with an
optical retical on a Wild M5 microscope at
x25.
Tooth measurements are denoted as
L
for length (maximum mesiodistal dimen-
sion) and W for width (maximum buccolin-
gual dimension). Subscript numbers indi-
cate lower teeth; superscript numbers
indicate upper teeth. Teeth are denoted as I
for
incisor, C
for
canine,
P
for premolar, and
M
for
molar.
Washakie Basin primates
Primate taxa present in University of Col-
orado Museum collections from early Eocene
deposits in the Washakie Basin are listed in
Table
1.
Approximately 100
of
the primate
specimens in this collection are omomyids,
and the remaining 200 are adapids.
Anemo-
rhysis savagei,
n. sp., is described from
NEW
EARLY EOCENE PRIMATE
325
Lysitean strata near Bitter Creek Station.
In order to understand the phylogenetic re-
lationship of the new
Anemorhysis
species to
other phenetically similar anaptomor-
phines, the following
taxa
were analyzed:
Teilhardina americana, Tetonoides pearcei,
Anemorhysis pattersoni, A. wortmani,
A.
sublettensis,
A.
natronensis,
and
Trogolemur
myodes.
The following University of Colo-
rado dental specimens have been recently
recovered and were examined in this study:
Tetonoidespearcei,
UCM 56408 P,-M3; UCM
MI,, UCM 56898
P34,
UCM 60947
65457 P,;
Trogolemur myodes,
UCM 59776
M,-partial M,, UCM 58957 MI. Specimens
ofAnemorhysis savagei,
n. sp., that were ex-
amined are listed under type and hypodigm
below. Recently reported material of
Trogo-
lemur myodes
from Nevada (Emry, 1990)
was also studied.
Phylogeny reconstruction
In this study, 16 dental characters of the
lower teeth (no upper teeth of
Anemorhysis
have yet been described) of
Anemorhysis,
Trogolemur,
and outgroups
Teilhardina
and
Tetonoides
were analyzed. These characters
are listed in Table 2, and the character
states exhibited by each taxon are provided
in Table
3.
Two- and three-state characters
were used to document the range of varia-
tion exhibited among these species.
Data was entered in
a
MacClade 3.01
(Maddison and Maddison, 1992) file and an-
alyzed with the PAUP 3.0s (Swofford, 1991)
program. The most parsimonious tree with
Teilhardina
and
Tetonoides
as
outgroups
was determined through the use of the ex-
haustive search option in PAUP. A strict
consensus of all equally parsimonious alter-
natives was then computed.
Estimating body size and dietary
adaptation
Body size is a crucial aspect of a mam-
mal’s adaption and can influence
its
dietary
regime. For example, as discussed in Kay
(19751, Kay and Simons (1980), and Kay and
Covert (1984),
a
primate less than
500
grams (a limit known
as
“Kay’s threshold
[Gingerich, 19811) is likely to have been pre-
56409 Mi-3, UCM 56894 P4-M,, UCM 56895
UCM 65084 P4-M,, UCM 65309 P3-M,, UCM
TABLE
2.
Characters and character states
used in this analysis
1.
11:1,
size: a
=
I,
=
or
slightly
>
12,
b
=
I,
>
I,,
c
=
I,
>
>
I,
(with
I,
root extending below cheek tooth row)
2.
P,
presence: a
=
absent; b
=
present
3.
P,
paraconid presence:
a
=
absent;
b
=
present
4.
P,
paraconid position:
a
=
low;
b
=
high
5.
P, entoconid:
a
=
absent
or
trace; b
=
small; c
=
large
6.
P,
root number:
a
=
two roots; b
=
one root
7.
P4
root number: a
=
two roots; b
=
one root
8.
P,
paraconid-metaconid spacing: a
=
widely
9.
P,
paraconid size: a
=
absenthmall; b
=
large
separated; b
=
close together
10.
P4
paraconid height: a
=
low (lower than protoconid);
11.
P,
entoconid-hypoconid spacing:
a
=
close together;
12.
P,
talonid width: a
=
narrow; b
=
wide
13.
Molar cusp wall orientation (M, entoconid-hypoconid
distance/total talonid width: a
=
S.72, b
=
>.73):
a
=
convergent;
b
=
non-convergent (vertical)
14. M, breadth at anterior aspect
of
talonid (anterior
breadth (at conjunction
of
cristid obliqua and
posterior trigonid WallYposterior width
(entoconid-hypoconid);
a
=
s.72;
b
=
>.73):
a
=
narrow; b
=
wide
15. Body size (based on M, size using tarsioid model
[Gingerich,
19821):
a
=
>35 g; b
=
135
g
16. Length
of
M,flength M,: a
=
M, nearly equal to or
longer than M, (M,/M,
s
.80);
b
=
M,, slightly shorter
than M, (M,/M,
=
.81-.89);
c
=
M, shorter than M,
(M,/M,
3
.90)
17.
M,
shape:
a
=
nearly square
(LW
<
1.25);
b
=
narrow
(L/W
2
1.26)
18.
P,
area relative to
P,
area: a
=
P,
nearly equal to
or
slightly smaller than
P,
(P,/P,
3.75);
b
=
P,
<
P4
(P,/P,
=
.56-.74; P,
1
1
smaller than
P,
b
=
high (nearly as high as protoconid
b
=
far apart; c
=
very far apart
(P,/P,
S
55)
dominantly frugivorous or insectivorous but
is
too small to have been predominantly foli-
vorous.
Among primates, body size is known to be
closely correlated with molar size (Kay,
1975; Gingerich et al., 1982). Body weights
for fossil primate taxa can be estimated
through the use of regression equations be-
tween body weight and MI area in samples
of living primate species (e.g., Gingerich,
1981; Gingerich et al., 1982; Conroy, 1987).
Several of these equations have been used to
estimate body size in omomyids. These
equations
are
based on data from various
groups composed either of a wide array of
primates (e.g., generalized primate equa-
tions) or of particular subsets (e.g., tarsioid
equations), It
is
not clear which of the pro-
posed estimates
are
most appropriate for
predicting body weight in omomyids. Gin-
gerich (1981) notes that extant tarsiers and
other insectivorous and carnivorous mam-
TABLE
3.
Distribution
of
dental characters in taxa discussed in text
-
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
Character
I,:I,
size
P,
presence
P,
paraconid presence
P,
paraconid position
P,
entoconid
P,
root number
P4
root number
P,
paraconid-metaconid
P,
paraconid size
P,
paraconid height
P,
entoconid-hypoconid
P,
talonid breadth
Molar cusp wall
orientation
M,: anterior breadth of
talonid basin
Body size
M, length/M, length
distance
Teilhardina
americana
unknown
(1/2)
present
absent
absentltrace
two roots
two roots
widely separated
small
low
close
X
narrow
convergent
narrow
135
g
M,<cM,
Tetonoides
pearcei
1*>>12
present
present
high
absentltrace
two roots
two roots
close together
large
high
close
narrow
convergent
narrow
135
g
M, nearly
=
or
>
M,
Anemorhysis
sauagei
I,
> >
I,
present
absent
absenvtrace
two roots
two roots
close together
small
low
far
X
narrow
vertical
wide
<35
g
Anemorhysis
sublettensis Anemorhysis
wortmani
unknown
Unknown
unknown
unknown
Unknown
unknown
two roots
close together
large
high
very far
wide
vertical
wide
<35
g
unknown
I,
3>
>
I,
absent
present
low
small
one root
two roots
close together
large
low
far
wide
vertical
wide
>35
g
Unknown
Anemorhysis
pattersoni
unknown
unknown
Unknown
Unknown
Unknown
two roots
two roots
widely separated
small
low
far
wide
vertical
wide
>35
g
unknown
Anemorhysis
natronensis
I1
>
1,
unknown
absent
large
two roots
two roots
widely separated
small
low
far
wide
vertical
wide
<35
g
Unknown
X
Tmgolemur
myodes
I,
>
>
>
I,
?absent
absent
large
one root
one root
widely separated
small
low
far
wide
vertical
wide
135
g
M, nearly
=
or
>
M,
X
M, shape nearly square nearly square nearly square nearly square nearly square nearly square narrow nearly square
18
P3/p4
area
P,
<
P,
P,<P,
P3<P,
unknown
nearlyequal
unknown
nearly equal
P,
<
c
P,
x
indicates
not
applicable.
NEW EARLY EOCENE PRIMATE
327
TABLE
4.
Shear quotients and body size estimates
of
extant small-hodied prosimians
and omomyid species discussed in text'
Shear Body size
Species
_______.
N LM2
ratio
Diet (grams)
Perodicticus potto
8
3.41 1.54 FIG 850-1,600
Galago crassicaudatus
6 3.76 1.69 FIG 1,000-2,000
Euoticus elegantulus
6 2.36 1.86 G 270-360
Anemorhysis pattersoni
1 1.75 1.89 (F) 42 (144)
Trogolemur myodes
3 1.58 1.89 iF) 24 (78)
Anemorhysis sublettensis
1 1.59 1.92 iF) 20 (70)
Galago alleni
7 2.81 1.95 F 19G340
Teilhardina amerieana
5 1.70 1.97 FA 40 (136)
Anemorhysis natronensis
1 1.59 1.98
(M)
28 (93)
Anemorhysis wortmani
2 1.69 2.00 (FA) 38 (130)
Loris tardigradus
6 2.88
2.00
I
270-350
Anemorhysis savagei
2 1.69 2.09
(1)
27 (87)
Galagoides demidovii 8
1.94 2.13 1 45-90
Arctocebus ealabarensis
6 3.56 2.18
I
150-270
Galago senegalensis
7 2.17 2.46
I
230-300
'Body size ranges
for
extant lorisids are from Charles-Dominique (1977). Body size estimates for fossil taxa based on Gingerich's Equation
1
for
generalized primates and, in parentheses, Equation
3
for
tarsioids (1981,
p.
355). Dietary information
for
extant taxa are from Charles-
Dominique (19771, Bearder and Martin (19801, and Hladik (1979). Inferred diets of fossil species given
in
parentheses.
N
=
sample size available
for estimation of sheat quotient;
LM:,
=
lower second molar occlusal length;
I
=
insects;
G
=
gum;
F
=
fruit.
Tetonoides pearcei
5 1.65 2.10
(1)
27 (91)
mals have relatively large cheek teeth for
their body size. He groups omomyids with
modern tarsiers in the Tarsioidea and pro-
poses the use of the tarsier model. Strait
(1991, in press) has suggested that use of the
tarsioid regression may underestimate body
size because not all omomyids were primar-
ily insectivorous, and because omomyids
may not be fossil tarsiers. However, Dagosto
and Terranova (1992) have used postcranial
measurements to estimate body weight and
suggested that some measurements of the
known omomyid postcrania indicate that
generalized primate estimates may be too
large. Among the anaptomorphines under
study here, postcranial remains are known
only for
Tetonoides
pearcei,
having been re-
covered from deposits associated with that
taxon. These elements are extremely small
and indicate that the tarsioid model may be
most accurate for these anaptomorphines.
It
is
our intention
to
obtain a very general
idea
of
the size of the omomyids discussed
here in order to better evaluate their dietary
adaptations (i.e., to determine if they were
under
500
grams). Therefore, we have pre-
sented two body weight estimates in Table 4,
one using Gingerich's (1981) generalized
primate equation and the other his tarsioid
equation.
A
frugivorous primate can be distin-
guished from an insectivorous one by the
relative molar shear crest development
(Kay, 1975; Kay and Simons, 1980; Kay and
Covert, 1984; Covert, 1985; Strait, 1991,
1993). On unworn teeth, shear development
can be expressed in terms of a ratio (Strait,
1991, 1993) by calculating the summed
length of shearing crests 1-6 (as defined by
Kay and Hiiemae U9741 and illustrated by
Strait [19931) on
M2
divided by the occlusal
length of that tooth. The mean shear ratio
for an extinct sample can be compared with
a
series of extant primates
of
similar size.
Among small extant primates, frugivorous
species have smaller shear ratios than do
insectivorous species (Kay, 1975; Kay and
Covert, 1984; Strait, 1991, 1993, in press).
In order
to
suggest dietary adaptations
among the anaptomorphines discussed
here, shear ratios were calculated.
RESULTS
Washakie
Basin
primates
The primates of the Washakie Basin were
taxonomically diverse (Table
1).
The spe-
cies-level diversity during the early Eocene
was about as great in this basin as in either
the Wind River (Stucky, 1984; Beard et al.,
1992)
or
Bighorn Basins (Bown, 1979; Bow
and Rose, 1987,1991). Washakie Basin omo-
myids and adapids were apparently equally
diverse. The presence
of
the anaptomor-
B.A.
WILLIAMS
AND
H.H.
COVERT
328
phine
Trogolemur
in the Lostcabinian (late-
early Eocene)
is
the earliest occurrence for
this taxon, known previously only from
Bridgerian-aged (middle Eocene) deposits
(Szalay, 1976; Emry, 1990; Beard
et
al.,
1992). These molar specimens resemble
both
Trogolemur
and
Anemorhysis
in having
broad trigonid basins on
M2-,.
They more
closely resemble
Trogolemur myodes
in
hav-
ing well-developed medial and lateral post-
protocristids. None of these specimens pre-
serve the antemolar dentition,
so
it
is
not
possible
at
present to determine the degree
of premolar reduction
or
incisor enlarge-
ment in this sample.
Arapahovius gazini
and
Copelemur praetutus
are
unique to the
Washakie Basin,
as
is
a
new, undescribed
species of anaptomorphine that may be the
sister taxon to
Tetonoides.
The presence of
Loveina minuta
is also of note because
it
is
an extremely rare taxon (Bown and Rose,
1984). Four of the taxa represent new spe-
cies.
Anemorhysis sauagei,
n. sp., described be-
low, is structurally transitional between
Teilhardina americana,
the earliest known
and possibly most primitive North Ameri-
can omomyid (Bown, 1976,1979; Bown and
Rose, 1987), and more derived species of
Anemorhysis
and
Trogolemur.
Two speci-
mens from early Eocene deposits in the
Wind River Basin, Wyoming, may also be
referrable to this new species.
Systematic
paleontology
Order Primates Linnaeus, 1758; family
Omomyidae Trouessart, 1879; subfamily
Anaptomorphinae Cope, 1883.
Genus
Anemorhysis
Gazin, 1958 (Figs.
14; Table 5; Appendices A-C).
Type species
Paratetonius? sublettensis
Gazin, 1952.
Included species
mani,
A.
natronensis,
A.
savagei,
n.
sp.
Age and geographic distribution
(early-middle Eocene) of Wyoming.
A.
sublettensis,
A.
pattersoni,
A.
wort-
Wasatchian to earliest Bridgerian
LhL4
Fig.
1.
Tetonoides
and
Anemorhysis,
SEM
photo-
graphs, occlusal
views.
Left:
Tetonoides pearcei,
holo-
type
USNM
22426.
Right dentary fragment preserving
P4-M3.
Right
Anemorhysis
pattersoni,
USGS
476,
holo-
type.
Left dentary fragment preserving
P,-M,.
Emended diagnosis
Anaptomorphines with
I,
root enlarged
relative to
I,
as
in
Tetonius, Tetonoides,
and
Arapahovius
but
less
hypertrophied than
Trogolemur,
in which
the
I,
root extends un-
der molars.
P,
larger relative to
P,
than in
Trogolemur.
Compared with
Teilhardina
and
Tetonoides,
third and fourth premolar
talonid basins are broader and oblique cris-
tids more buccally placed; entoconids far-
ther from hypoconids relative to breadth of
talonid.
P,,
talonids longer than in
Arapa-
hovius, Tetonius,
and
Teilhardina crassi-
dens.
Molars less basally inflated than
in
Tetonius.
P4
is
two-rooted, unlike that of
Trogolemur.
First and second lower molar
morphology similar to
Trogolemur
but un-
like other anaptomorphines in having (espe-
cially on
M,)
broad trigonid basins, vertical
(non-convergent) cusp slopes, and talonid
basins that are buccolingually broad at the
conjunction of the cristid obliqua with the
posterior trigonid wall. Medial and lateral
postprotocristids on lower molars less we11
NEW EARLY EOCENE PRIMATE
329
Fig. 2.
Anemorhysis
species and
Tetonoides pearcei,
SEM photographs,
lingual
view, premolars and molars
only. Left, top:
Anemorhysis pattersoni
holotype,
USGS
476. Left dentary fragment preserving crowns of P,-M,.
Left,
middle:
Anemorhysis savagei,
n. sp., holotype
UCM 56410. Right dentary fragment preserving crowns
of P,-M, and alveoli of Il-P2. Left, bottom:
Tetonoides
pearcei,
UCM 56408. Right dentary fragment preserv-
ing alveoli of
I,-P,,
and crowns of P,-M,. Right, top:
developed than in
Trogolemur.
Third molars
larger relative
to
Ma than in
Teilhardina
(similar to
Trogolemur)
but smaller than in
Tetonoides.
M, talonids more broadly ex-
panded than in
Teilhardina
or
Tetonoides
but less
so
than in
Trogolemur.
Molar
enamel smooth, unlike
Arapahouius, Strigo-
rhysis,
and some
Absarokius.
Anemorhysis savagei,
sp.
nov.
(Figs.
24,
Appendices A-C).
Holotype
P,-M,, and alveoli for 11-P2.
UCM
56410,
right mandibular body with
Anemorhysis natronensis
holotype, CM 41137. Left den-
tary fragment preserving root
of
I,,
partial crowns of
I,
(partial), C,, P,. Right, middle:
Anemorhysis wort-
manz
holotype, USGS 6554. Right dentary fragment
preserving root
of
I,,
P,-M,. Right, bottom:
Anemorhy-
sis sublettensis
holotype, USNM 19205. Left dentary
fragment preserving crowns
of
P,-M,. Bar equals ap-
proximately 3.4 mm.
UCM
60915,
left P,-MI; UCM
62682,
right
P,-M, and root for I,, alveoli 12-P,; CM
39654,
left P,-M,; CM
28915,
right
Age and geographic distribution
The type and other UCM specimens are
from UCM Locality
88040,
early Eocene,
Wasatch Formation, Washakie Basin, Wyo-
ming. This locality
is
assigned to the Ly-
sitean subage of the Wasatchian Land Mam-
mal Age (Wa,). CM
39654
comes from Lysite
Flats Locality
7
and CM
28915
from Lysite
Flats, Davis Draw Locality, both from
Lysitean age deposits in the Wind River For-
mation, Wind River Basin, Wyoming.
Hypodigm Etymology
The type specimen and UCM
56413
right Named for Dr. Donald E. Savage in honor
isolated MI; UCM
56899,
left
MZp3;
UCM of his contributions to understanding
56900,
left P,-M,; UCM
60914,
right
MI-,;
Eocene faunas.
330
B.A.
WILLIAMS
AND
H.H.
COVERT
Fig.
3.
Anernorhysis sauugei,
n.
sp.,
SEM
stereophotograph, occlusal view.
UCM
56410. Right
P,
=
M,.
Bar equals approximately
5.00
mm.
Diagnosis
Differs from
Anemorhysis wortmani,
A.
natronensis,
and probably
A.
pattersoni
in
retaining
P,.
Molars and third and fourth
premolars absolutely smaller than corre-
sponding teeth
of
A. pattersoni
and
A.
wort-
mani.
Differs from
A.
wortmani
in lacking a
paraconid on
P3,
and differs from
A.
wortmani
and
A.
sublettensis
in having a smaller, lower,
and more crestiform
P,
paraconid. Differs
from all other
Anemorhysis
in having a less
buccolingually broad
P,
talonid.
Description
Among anaptomorphines,
A.
savagei
is
most similar to other species
of
Anemorhy-
sis, Trogolemur myodes, Tetonoides pearcei,
and
Teilhardina americana.
Several perti-
nent characters and their states are shown
in Table
2.
Incisor and canine roots
or
alveoli are
unknown for
A. sublettensis
and
A.
patter-
soni.
In
A.
savagei
the I, alveolus is enlarged
relative
to
the I, alveolus, similar to the
condition in
T.
pearcei
(and apparently
T,
NEW EARLY EOCENE PRIMATE
331
Fig.
4.
Anemorhysis sauugei,
n.
sp.,
SEM
photograph, buccal view.
Top:
UCM
56410,
holotype.
Bot-
tom:
UCM
62682.
Right dentary fragment preserving
root
of
I,,
alveoli of
12-P2,
and crowns
of
P,-M,.
Bar
equals approximately
3.4
mm.
TABLE
5.
Stratimaohic occurrences
of
anaDtomorDhines discussed in text
Species
Anemorhysis natronensis
Trogolemur amplior
Trogolemur myodes
Anemorhysis sublettensis
Anemorhysis
wort mani
Anemorhysis savagei
Tetonoides pearcei
Anemorhysis pattersoni
Teilhardina amerieana
Land mammal subage
and biochron
~-
Gardnerbuttian (Br-1)
Gardnerbuttian (Br-1)
Lostcabinian-Blacksforkian
(Wa-7
to
Br-2)
Lost Cabin (Wa-7)
Lysite (Wa-6)
Lysite (Wa-6)
Upper Graybull (Wa-5)
Middle-Upper Graybull
(Wa-4 to Wa-5)
Middle Sandcouleean
(Wa-1)
Location
Wind River Basin
Wind River Basin
Washakie and Bridger Basins
Green River Basin
Bighorn Basin
Washakie Basin
Wasbakie Basin
Bighorn Basin
Bighorn Basin (including Clarks
Fork
Rasini
~
'Taxondistributions based upon datacompiledfromGazin(1952,1962), Bown and
Rose
(1984,1987,1991), andBeardet a]. (1992). Oldest
=
Wa-
1; youngest
=
Br.2. Abbreviations as in Table
1.
americana
[Bown and Rose,
19871)
but on their alveoli) may be expressed
in
a
less enlarged than in
A.
wortmani or Trogo-
series from largest
to
smallest as:
lemur myodes.
P,
is absent. The
P,
is single-
I1
>>
C,
>
P,
>
I,,
where
>
means slightly
rooted, and the alveolus
is
smaller than that larger and
>>
means much larger. The
for
P,.
The relative sizes of
Il-P2
(based ratio
is
like that seen in
Tetonoides pearcei.
332
B.A.
WILLIAMS
AND H.H.
COVERT
The P, and
P,
trigonids are less molari-
form than in
A.
wortmani,
A.
sublettensis
(P,
unknown), and
T.
pearcei.
The third premo-
lar has two roots and is a simple tooth domi-
nated by a protoconid, without paraconid
or
metaconid.
A
very faint cristid obliqua
is
buccally oriented as in other species of
Ane-
morhysis.
This tooth has a small hypoconid
but lacks an entoconid. The
P4
has
a
low,
lingually positioned paraconid and a low
metaconid. There is variation in the expres-
sion of the paraconid; in some specimens
(UCM 56410, CM 39654) it is more crestlike,
whereas in others (UCM 62682, UCM
60915) it is a very small distinct cusp. The
paraconid in all specimens is lower and
smaller than in
A.
wortmani,
A.
sublettensis,
and
T. pearcei.
The metaconid is low and
small. There is a small but distinct hypo-
conid and small entoconid; these cusps are
far apart relative
to
the breadth of the tal-
onid, as in other
Anemorhysis;
however, the
talonids of
P,,
are relatively less buccolin-
gually expanded than in other
Anemorhysis
(especially less
so
than in
A. sublettensis).
The premolar and molar buccal cingula are
weakly developed.
The first and second lower molars are ab-
solutely smaller than in all species of
Ane-
morhysis
except
A. sublettensis. A. savagei
has a relatively smaller M, paraconid and
more closely appressed
MI-,
paraconids and
metaconids.
The
M,
is unknown for other species
of
Anemorhysis
but is preserved in three speci-
mens
of
A.
sauagei.
Its paraconid is closely
appressed
to
the metaconid and
is
strongly
lingual. Compared with
Teilhardina ameri-
cana,
the
M,
is larger relative to the
M,,
but
it is relatively smaller than in
Tetonoides
pearcei
or
Trogolemur myodes.
A.
savagei
and
T.
pearcei
overlap in size;
however, there are several additional char-
acters in which these taxa differ.
A
Stu-
dent’s t-test demonstrates that
A,
savagei
has an absolutely narrower M,
(t
=
-4.06,
PC
.001) and a shorter M,
(t
=
-4.38,
P
<
.007) (AppendixB).
Stratigraphic occurrence and relative
ages
Table
5
depicts the mammalian subages
and corresponding faunal zones in which
Teilhardina americana, Tetonoides pearcei,
Trogolemur myodes,
T.
amplior,
and species
of
Anemorhysis
are known to occur. The
UCM Washakie Basin sample of
Anemorhy-
sis savagei
comes from a single locality
(UCM locality
880401,
referred to as “Turtle
Graveyard by Savage and Waters (1978).
This locality has also produced hypodigm
specimens of
Arapahouius gazini
(Savage
and Waters, 1978) and the best known sam-
ple of the extremely rare notharctine adapid
Copelemur australotutus
(Beard, 1988; Co-
vert, 1990). Turtle Graveyard
is
situated ap-
proximately 70 meters higher in the main
body
of
the Wasatch Formation than
is
the
Bitter Creek Promontory Hill (UCM locality
88039), the type locality of
Tetonoides pear-
cei.
Whereas there is some overlap in the
mammalian faunas of these localities, the
primates are distinctly different (Table 1).
Anemorhysis savagei
may be slightly
younger than the oldest known species of
the genus,
A.
pattersoni;
however,
it
is diffi-
cult to assess their relative ages due
to
lack
of precision in interbasinal faunal correla-
tion.
Phylogeny and character evolution
A
parsimony analysis using a PAUP ex-
haustive search yields two trees of equiva-
lent minimum parsimony. With uninforma-
tive characters excluded, these networks are
23 steps long and have a consistency index
of
0.69 (rescaled
=
0.451,
a retention index
of
0.65, and a homoplasy index of
0.33.
A
strict
consensus of these trees is shown in Figure
5.
Each of the most parsimonious trees has
the following features:
1.
Tetonoides pearcei
is outside the
Ane-
morhysis
clade. It resembles
Teilhardina
in
primitive characters such as the retention
of
P, and in having convergent molar cusp
walls and narrow premolar and molar tal-
onid basins but has apparently derived pre-
molar trigonids featuring a distinct P, para-
conid and a large, high
P,
paraconid.
2.
Primitive
Anemorhysis
is
specialized
from
Teilhardina
by having more complex
premolar talonids (broader basins) and de-
rived molar morphology (non-convergent
cusp walls, anteriorly broad talonid basins).
Anemorhysis sauagei
is
the most primitive
NEW EARLY EOCENE PRIMATE
Teilhardina americana
Tetonoides pearcei
Anemorhysis savagei
Anemorhysis sublettensis
Anemorhysis wortmani
Anemorhysis pattersoni
Anemorhysis natronenesis
Trogolemur myodes
Fig.
5.
Strict consensus phylogeny.
333
member of the
Anemorhysis
clade, retaining
P,
and premolariform
P3+
3.
More derived
Anemorhysis
show fur-
ther trends toward premolar specialization
including
P,
loss and, in some taxa, greater
premolar trigonid andlor talonid complexity.
The primitive premolar trigonid morphology
of
A.
savagei
demonstrates that molariza-
tion of the premolars must be a parallelism
in
Tetonoides
and some
Anemorhysis.
4.
Anemorhysis sublettensis
and
A.
wort-
mani
are sister taxa, sharing
P,
trigonid
character states (large paraconid that is
close to the metaconid).
A.
sublettensis
is
further derived in having a
P,
with an ex-
ceptionally long and wide talonid basin.
5.
The source of
Trogolemur
comes from
within the
Anemorhysis
clade, but exact sis-
ter-group relationships are unclear.
Trogo-
lemur
has the molar synapomorphies that
unite species
ofAnemorhysis
but is more de-
rived than any species of that genus in hav-
ing an
I,
root that is dramatically enlarged
and runs underneath the premolars and in
having a
P,
that is single-rooted and just
half the size
of
P,.
A
strict consensus of the
three most parsimonious trees demon-
strates that there
is
an unresolved trichot-
omy linking
A.
pattersoni,
A.
natronensis,
and
Trogolemur myodes,
which share the
apparently primitive
Ps4
trigonid structure
seen in
Teilhardina americana
(small, low
paraconid that
is
widely separated from the
metaconi d)
.
6.
Anemorhysis natronensis
is probably
the species least phenetically similar
to
other members of the genus and is autapo-
morphic in several features, such as narrow
molars (especially
M,),
very large
P,,
ento-
conid, and a central incisor that is only
slightly larger than the lateral incisor
(Beard et al., 1992).
Comments on premolar and incisor
evolution
The polarity of several of the characters in
which anaptomorphines and other omomy-
ids vary, such as molarization
of
the premo-
lars and enlargement
of
the
I,
relative to the
I,,
is difficult to assess. Selecting the out-
group in a cladistic analysis (thus determin-
ing polarity) can profoundly affect the inter-
pretation of cladogenesis. By rooting the
network with
Teilhardina americana
we are
presuming that the dental morphology
of
that taxon represents the morphology prim-
itive for anaptomorphines.
It
has been ac-
cepted by many researchers that Anapto-
334
B.A.
WILLIAMS
AND
H.H.
COVERT
morphinae is the most primitive subfamily
of the Omomyidae and gave rise to the
later-occurring Omomyinae (e.g., Simpson,
1940; Szalay, 1976; Gingerich, 1981). There-
fore, the dental morphology of the oldest
known anaptomorphines
(Teilhardina
americana
and the European
Teilhardina
belgica)
has been thought to represent the
primitive condition for omomyids (e.g., Sza-
lay, 1976; Gingerich, 1981; Bown, 1976,
1979; Bown and Rose, 1987).
It is possible that anaptomorphines
are
not the most primitive members of Omomy-
idae. Taxa attributed by many authors to
the subfamily Omomyinae, such
as
Loveina
and
Steinius
(Gazin, 1958; Szalay, 1976;
Gingerich, 1981; Bown and Rose, 1987;
Honey, 1990), are possibly more primitive in
some features than
are
anaptomorphines
such as
Teilhardina
(Gazin, 1958; Rose and
Bown, 1991).
The
P,
of
Steinius vespertinus
has
a
low
paraconid and only slightly developed meta-
conid, whereas the
P,
trigonid
of
Loveina
zephyri
is well developed. Unfortunately,
the
P,
of the older and possibly more primi-
tive species of
Loueina,
L.
minuta,
is
un-
known. The
P,,
trigonids of the anaptomor-
phines under study here show considerable
variation (Table
3;
Fig. 2). In
T.
belgica
and
T.
americana,
the
P,
paraconid and meta-
conid are usually poorly developed, but in
later-occurring
T.
crassidens
those cusps
are
distinct and well developed.
If anaptomorphines are the most primi-
tive omomyids, and
Teilhardina
is represen-
tative of the anaptomorphine morphotype,
then simple premolar trigonids
are
primi-
tive for the family. It
is
unclear what the
omomyine morphotype might be with regard
to premolar trigonid structure. Understand-
ing the omomyid morphotype
is
further com-
plicated in that Omomyinae may not be a
monophyletic group. The oldest and possibly
most primitive subtribe of Omomyinae is
Washakiini (Honey, 1990). There
is
some ev-
idence that washakiins may be
a
primitive
outgroup to all other omomyids, and possi-
bly all other omomyids, and possibly all
other primates (Williams and Kay, 1992).
Additionally,
Steinius
may be an anapto-
morphine rather than an omomyine (Bown
and Rose, 1984, 1987; Williams and Kay,
1992). The development of complex premo-
lars
apparently occurred in more than one
group of anaptomorphines (e.g.,
Teilhar-
dina, Tetonoides, Anemorhysis).
The polarity of central incisor enlarge-
ment in omomyids similarly has been de-
bated. Small, equisized lower incisors were
probably primitive
for
primates (Cartmill
and Kay, 1987; Szalay et al., 1987; Ginger-
ich et al., 1991; Covert and Williams, 1991b)
and possibly for anaptomorphines
as
seen in
Teilhardina belgica
(Gingerich, 1977). How-
ever, the central incisor of
T.
americana
may
have been moderately enlarged (Bown and
Rose, 1987). Central incisor enlargement is
seen in many anaptomorphine taxa such as
Tetonoides
and
Arapahovius (contra
obser-
vations by Beard
et
al., 1992),
Anemorhysis,
Tetonius, Pseudotetonius,
and
Tatmanius,
as
well
as
in
some other omomyids. This
is
apparently a homoplastic convergence in
several omomyid groups and may not indi-
cate phylogenetic relatedness. Until the fos-
sil record for primitive omomyids
is
better
known, it
is
impossible to determine the
dental morphotype for these primates.
Body size and diet
Whether the generalized primate
or
tar-
sioid estimate
is
used,
it
is apparent that
these anaptomorphines were well under
Kay’s threshold of 500 grams and were prob-
ably less than
50
grams (Table
4).
Smith
(1993) noted that some body weight esti-
mates are biased by log transformation and
require corrections of up to 19%. Even if the
maximum corrections indicated by Smith
are applied, predictions for all of the anapto-
morphines examined here are less than 200
grams. These estimates, in addition to the
overall size of their jaws and of the known
postcrania of
Tetonoides,
suggest that they
were about
as
small as the smallest living
primates: the galagine
Galagoides,
the
cheirogaleid
Microcebus,
and the platyr-
rhine
Cebuella. Anemorhysis savagei
is
one
of the smallest omomyids known. If
A. sau-
agei
evolved from the larger-bodied
Teilhar-
dina americana
and represents the stem
member of
its
genus,
it
appears that
Anemo-
rhysis
underwent some size reduction fol-
lowed by size increase in some taxa
(A.
pattersoni,
A.
wortmani).
NEW EARLY EOCENE
PRIMATE
335
1
I
II
I
11’1
IIIII
I
-
-
more
fruit
and
gum
I
more insects
W‘
Trogolemur myodes
(3)
I
Anemorhysis pattersoni
(1)
I
Anemorhysis sublettensis
(1)
I-W
Teilhardina americana
(5)
I@
Anemorhysis natronensis
(1)
l-+l
Anemorhysis wortmani
(2)
I
W
Anemorhysis savagei
(2)
F0-I
Tetonoides pearcei
(5)
I
*
Perodicticus potto
(8)
t-0-I
OtoEerrb crassicaudatus
(6)
A
‘Euoticus elegantulus
(6)
W
Galagoides alleni
(7)
K)-l
Loris tardigradus
(6)
I
Galagoides demidovii
(8)
I
)--0--1
Arctocebus calabarensis
(6)
-Galago senegalensis
(7)
I
II II
II
I
I
IIIII
I
1.2
1.4
1.6
1.8
2.0 2.2 2.4 2.6
Mean shearing ratio
Fig.
6.
Diets of extant lorises and galagos and sug-
gested diets of fossil anaptomorphines discussed in text.
Mean shearing ratio
=
sum of shear crests
1-6
on
MJM, occlusal length. Black circles indicate fossil taxa
and white circles indicate extant taxa. Vertical lines on
either side of taxon indicate standard deviation. Num-
Shear ratios for the available samples of
the anaptomorphines discussed in the text
as well as several lorises and galagos are
provided in Table
4
and Figure
6.
The shear-
ing ratios of
Anemorhysis savagei
and
Te-
tonoides pearcei
fall into the range of ani-
mals such as
Galagoides demidovii
and
Loris tardigradus
that feed primarily on in-
bers in parentheses indicate sample size. Vertical line
dividing animals that are predominantly fruit and gum
eaters from those that are predominantly insect eaters
is based on information from Charles-Dominique
(1977),
Bearder and Martin
(19801,
and Hladik
(1979).
sects.
Anemorhysis pattersoni,
A.
subletten-
sis,
and
Trogolemur myodes
have ratios
more similar
to
frugivores such as
Euoticus
elegantulus
and
Galagoides alleni.
The
other taxa do not show apparent specializa-
tion in either direction and may have been
mixed feeders, eating both insects and
fruits. Strait
(1991)
analyzed shearing ra-
336
B.A.
WILLIAMS
AND
H.H. COVERT
tios
of
some
of
these taxa using a different
method of measurement. Her results differ
from ours for individual taxa but are gener-
ally comparable. As indicated by
our
results
and those
of
Strait,
it
is apparent that no
anaptomorphines (we excluded
Loveina
from Anaptomorphinae) possessed the tren-
chant shearing blades
of
highly insectivo-
rous forms such as
Galago senegalensis
nor
the extremely bunodont molars seen in
primates that are specialized frugivorel
gummivores such as
Perodicticus potto
(with
the possible exception
of
Gazinius,
a middle
Eocene North American anaptomorphine
not evaluated here
or
by Strait due to insuf-
ficient material). It appears that the earliest
anaptomorphines were opportunistic, mixed
feeders and that the dietary specializa-
tions observable in modern primates did not
occur until considerably later in their evolu-
tion.
It is interesting
to
note that there appears
to
be litte association between trends in inci-
sor
enlargement and a dietary shift from an
insectivorous
to
a frugivorous diet. Although
Trogolemur
exhibits the largest incisors and
was frugivorous,
Anemorhysis sauagei
had
already commenced this trend toward inci-
sor
enlargement while maintaining an in-
sectivorous diet. Perhaps incisor enlarge-
ment in these taxa was a compromise for a
range
of
adaptive roles, such as improved
oral manipulation of all food items (not just
fruit), grooming, and defense.
CONCLUSIONS
The Washakie Basin had a diverse assem-
blage of fossil primates during the early
Eocene and documents the presence of sev-
eral rare middle and late Wasatchian taxa.
This new material is especially noteworthy
because it samples the middle-late Wasatch-
ian transition more completely than do Sam-
ples from either the Bighorn
or
Wind River
Basins. The Washakie Basin sample thus
increases our understanding
of
the taxo-
nomic diversity, phylogenetic relationships,
and paleobiology of these minute primates.
The Lysitean-age fauna near Bitter Creek
Station contains a previously unknown
primitive anaptomorphine,
Anemorhysis
savagei.
This taxon shares the derived pre-
molar and molar features characteristic
of
other species
of
Anemorhysis
and is clearly
referable
to
this genus.
A
phylogenetic anal-
ysis of this and other small-bodied anapto-
morphines indicates that this taxon is prim-
itive relative to other members of the genus.
A.
savagei
is structurally intermediate be-
tween
Teilhardina americana
and the group
that includes
Anemorhysis
and
Trogolemur
and may be the stem member of the latter
group. The paucity
of
data on the anterior
and upper dentitions of these omomyids ren-
ders further resolution
of
phylogenetic rela-
tionships impossible at present.
It
appears that
Trogolemur
may be seated
within the
Anemorhysis
radiation, making
Anemorhysis
paraphyletic with respect to
Trogolemur.
The occurrence of
Trogolemur
in Lostcabinian deposits in the Washakie
Basin is noteworthy because
it
is
older than
previously known samples of that genus. Re-
covery
of
the antemolar dentition
of
this
early-occurring
Trogolemur
might help to
understand the apparent transition from
Anemorhysis
to
Trogolemur.
Teilhardina, Tetonoides, Trogolemur,
and
Anemorhysis
were in the size range
of
the
smallest extant primates. They probably ate
a variety
of
fruits and insects. Neither these
nor other anaptomorphines show the range
of shearing potential on their molars associ-
ated with the extremes of dietary specializa-
tion known in extant primates. Incisor en-
largement in anaptomorphines does not
appear
to
be associated with specialization
in either frugivory
or
insectivory but may be
related
to
various adaptive roles, including
specialized grooming, defense,
or
food ma-
nipulation.
ACKNOWLEDGMENTS
We
thank Diana Ayers-Darling, Mark
Hamrick, Carol Harrisville-Wolff, David
Hobbs, Mark Anthony, Robert Anemone,
John Wible, and others who have helped
to
collect the Washakie Basin primate speci-
mens. We are also grateful
to
Tom Bown,
Ken Rose, Peter Robinson, and Chris Beard
for many insightful conversations on anap-
tomorphine evolution and
to
Richard Kay,
Ken Rose, Mary Maas, Suzanne Strait, Tom
Bown, and two anonymous reviewers for
helpful comments on the manuscript. Shear
crest data for extant taxa are courtesy
of
NEW EARLY EOCENE PRIMATE
337
Richard Kay. Leonard Krishtalka and Chris
Beard, Carnegie Museum, Tom Bown,
USGS, and Robert Emry, USNM, gener-
ously allowed us to examine specimens in
their care. Funding for this work has been
provided by the University
of
Colorado
Mu-
seum
W.
Van Riper and
W.H.
Burt
Fund
grants.
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APPENDIX
A.
Measurements (mmi
of
teeth
of
Anemorhvsis savapei.
SD.
now.
P,W P,L P,W P,L M,W MIL M,W M,L M,W
M,L
Specimen
UCM
56410
0.90
1.15 1.05 1.20 1.35 1.75 1.50 1.65 1.25 1.90
UCM
62682 0.95 1.10 1.10 1.30 1.35 1.70 1.35 1.70
- -
UCM
56900
- -
1.25 1.30 1.50 1.65
UCM
56899
- -
- -
-
1.45 1.70
UCM
56413
-
UCM
60915
-
-
1.15 1.50 1.45 1.75
UCM
60914
-
- - -
1.35 1.75 1.50 1.75 1.25 2.00
CM
39654
- -
1.30 1.40 1.40 1.75 1.45 1.80
CM
28915
-
- - -
1.35 1.75 1.50 1.70
'Fmm
the Washakie Basin, Wyoming
iUCM
specimens) and Wind
River
Basin, Wyoming
iCM
specimens) and other taxa referred
to
in this
paper. Dashes indicate no teeth.
-~
-
-
-
-
-
-
- -
1.50 1.55 1.15 1.95
- - - -
-
-
-
-
-
-
-
-
APPENDIX
B.
Summarv statistics for anavtomorvhines discussed in text
'
Teilhardina
americana
n
mean
OR
SD
n
mean
OR
SD
n
mean
OR
SD
n
mean
OR
SD
n
mean
OR
SD
n
mean
OR
SD
n
mean
OR
SD
n
mean
OR
SD
n
mean
OR
SD
n
mean
OR
SD
11
1.10
1.05-1.15
.03
1.42
1.2Ck1.55
.ll
1.39
1.25-1.55
.09
1.60
1.451.75
.07
1.61
1.40-1.80
.10
26
1.96
30-2.10
.09
27
1.68
.40-1.85
.10
1.91
.8Ck2.10
.09
8
1.35
.27-1.43
.07
8
2.10
1.94-2.19
.08
11
18
18
26
27
Tetonoides
pearcei
4
.98
.95-1
.OO
.03
4
1.23
.lo
5
1.20
.09
5
1.45
1.33-1.52
.07
9
1.43
1.33-1.50
.08
9
1.75
1.65-1.91
.09
9
1.58
1.52-1.65
.05
9
1.69
1.59
.08
4
1.35
1.27-1.40
.06
4
2.12
2.10-2.20
.05
i.oai.30
i.oai.30
Anemorhysis
sauagei
2
-
.go, .95
-
2
-
1.10, 1.15
-
5
1.17
1.05-1.30
.10
5
1.34
1.20-1.50
.ll
8
1.40
1.35-1.50
.06
8
1.73
1.65-1.75
.04
6
1.47
1.35-1.50
.06
6
1.70
1.55-1
.SO
.09
3
1.22
1.15-1.25
.06
3
1.95
1.90-2.00
.05
Anemorhysis
pattersoni
-
2
1.30, 1.40
2
1.50, 1.60
2
1.65, 1.75
2
1.90, 2.00
2
1.60, 1.85
2
1.90, 1.95
-
-
-
-
-
-
-
-
-
-
-
-
Anemorhysis
wortmani
1
1.20
1
1.40
2
-
-
-
-
-
1.35, 1.40
-
2
-
1.40, 1.70
-
2
1.60
2
-
-
-
1.90, 1.95
-
2
-
1.55, 1.65
-
2
-
1.80, 1.90
-
-
-
-
-
-
-
-
-
Anemorhysis
natronensis
1
1.15
1
1.40
1
1.30
1
1.50
1
1.40
1
1.80
1
1.30
1
1.80
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Anemorhysis
sublettensis
.~
-
-
1.10
1
1.40
1
1.30
1
1.65
1
1.40
1
1.70
-
-
-
-
-
-
-
-
-
-
-
Trogolemur
myodes
~~ ~
3
.84
.80-.90
.05
1.00-1.11 1.07
.06
3
1.23
1.20-1.25
.03
3
1.24
1.10-1.43
.17
5
1.42
1.27-1.50
.11
5
1.67
1.50-1.75
.lo
5
1.52
1.3g1.60
.10
5
1.57
1.40-1.85
.19
4
1.30
1.21-1.40
.08
4
2.04
2.00-2.10
.05
3
'Measurements are in millimeters. Specimen data from following sources:
Teilhardina americana
from
Bown and Rose
(1987:31;
p3-M~ only; individual specimens listed in appendix) and authors' measurements
(M,
only, specimens
UM
67424, 71091, 75610,76600;
UW
6896,7098;
USGS
7179,12194)
for
A.
pattersonr
and
A.
wortrnanz
from Bown and Rose
(1987;
specimens and measurements listed in appendix), for
Trogolemur
myodes from Emry
(1990:194;
USNM specimens only) and authors' measurements (AMNH
12599,
YPM
13523,
UCM
58776,
UCM
58957),
A.
natronensis
from Beard et al.
(1992;
CM
41137),
T. pearceL
from Gazin
(1962;
specimens YPM
14084,
USNM
22382
and
223831,
and author's measurements (specimens USNM
22426,
and UCM
56408, 56409, 56894, 56895, 56898, 60947, 65084, 65309, 65457).
Abbreviations: n
=
sample size;
OR
=
observed range;
SD
=
standard deviation.
APPENDIX
C.
Areas and ratios
for
anaptomorphines discussed in text
Teilhardina Tetonoides Anemorhysis Anemorhysis Anemorhysis Anemorhysis Anemorhysis Trogolernur
Dimension
americana pearcei
sauugei
puttersoni wortmani natronensis sublettensis myodes
Area
P,
(11)
1.56
(4)
1.21
(2)
1.04
-
(1)
1.68
(1)
1.61
-
(I)
92
Area P4
(18)
2.24
(5)
1.74
(5)
1.57 (2) 2.02
(2)
2.13
(1)
1.95
(1)
1.54
(1)
1.77
Area
M,
(26) 3.16 (9)
2.50
(8)
2.42
(21
3.32
(2)
3.08
(11
2.52
(1)
2.15
(2)
2.27
Area M,
(27) 3.21 (9) 2.67 (6) 2.50
(2)
3.32 (2) 2.96
(1)
2.34 (1)
2.38
(3) 2.60
- -
-
(2)
2.56
Area P,/Area
P,
.70
.70
.66
-
.79
.83
-
.52
Area M,
(8)
2.87 (4) 2.86
(3)
2.38
-
Area P,/Area M,
.71 .70 .65 .63 .69 .77 .72 .77
Area MJArea M,
.98 .94 .97
1.00
1.04 1.08
.90
.87
p3m
p4m
MIN
M2
m
P,
m1
L
-
-
1.02
1.29 1.26 1.22
-
1.17 1.22
-
1.34
1.15
1.21
1.15
1.15
1.13
1.15 1.27
1.15
1.22
1.22 1.24 1.15 1.20 1.29 1.27 1.32
1.14 1.07 1.16 1.12
1.16
1.38 1.21 1.10
.82 .83 .77 .79
.81
.83
.85
.83
P.
wm.
w
.86
.84 .84 .79 .86 .93
.85
.95
-
-
Area MJArea M,
1.12 .93 1.05
’
Measurements are in millimeters. Areas are in square millimeters
