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New early Eocene anaptomorphine primate (Omomyidae) from the Washakie Basin Wyoming with comments on the phylogeny and paleobiology of anaptomorphines.

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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) anaptomorphine, Anemorhysis sauagei, n. sp., is structurally intermediate between Teilhardina 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 Teilhardina 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 insectivory seen in extant prosimians. Incisor enlargement does not appear to be
associated with specialization in either fruits or insects but may have been a n
adaptation for specialized grooming or food manipulation.
0 1994 Wiley-Liss, Inc.
Fossil primates of North America first occur in the earliest Eocene, a time period referred 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 subfamily is the Anaptomorphinae (e.g., Szalay,
1976; Gingerich, 1981). Therefore, reconstruction of the phylogeny and behavior of
these early forms should help us to understand basal primate paleobiology.
Much of what we know about early anaptomorphines is based upon research in the
Bighorn Basin (including the Clarks Fork
0 1994 WILEY-LISS, INC
Basin) (e.g., Bown, 1974, 1976, 1979; Gingerich, 1981; Bown and Rose, 1984, 1987)
and the Wind River Basin (e.g., Stucky,
1982, 1984; Beard et al., 1992) of Wyoming.
I n the Bighorn Basin, latest Wasatchian
(Lostcabinian) fossils are sparsely represented (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.
B.A. WILLIAMS AND H.H. COVERT
324
TABLE 1 . Primates known from the Wasatch Formation of the Washakie Basin. Wvomine'
Omomyidae
Adapidae
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.
Cantius cf. uenticolus
Cantius frugiuorus
Copelemur australotutus
Copelemur tutus
Notharctus cf. robinsoni
Land mammal subage and biochron
Lostcabinian (Wa-7)
Lysitean (Wa-6)
Upper Graybullian (Wa-5)
Lower Graybullian (Wa-3-4)
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 a t 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, including approximately 300 jaws, isolated teeth,
and postcranial bones of primates. The new
material has been recovered from the Wasatch 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 described from near Bitter Creek was Tetonoides pearcei (Gazin, 1962), and Arapahovius 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
UCM
UM
USGS
usm
Carnegie Museum o f Natural History,
Pittsburgh, PA
University of Colorado Museum,
Boulder, CO
University o f Michigan Museum o f
Paleontology
United States Geological Survey,
Denver, CO;
United S t a t e s National Museum,
Washington, DC.
In this paper we 1) provide a taxonomic
list of primates occurring in Wasatchian deposits 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 Anemorhysis and Trogolemur; 3) discuss the
phylogenetic relationships among these
anaptomorphines; and 4)make suggestions
about their body size and dietary adaptations.
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 dimension) and W for width (maximum buccolingual dimension). Subscript numbers indicate 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 Colorado 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. Anemorhysis savagei, n. sp., is described from
325
NEW EARLY EOCENE PRIMATE
Lysitean strata near Bitter Creek Station.
In order to understand the phylogenetic relationship of the new Anemorhysis species to
other phenetically similar anaptomorphines, the following taxa were analyzed:
Teilhardina americana, Tetonoides pearcei,
Anemorhysis pattersoni, A . wortmani, A.
sublettensis, A. natronensis, and Trogolemur
myodes. The following University of Colorado dental specimens have been recently
recovered and were examined in this study:
Tetonoidespearcei, UCM 56408 P,-M3; UCM
56409 Mi-3, UCM 56894 P4-M,, UCM 56895
MI,, UCM 56898 P34, UCM 60947
UCM 65084 P4-M,, UCM 65309 P3-M,, UCM
65457 P,; Trogolemur myodes, UCM 59776
M,-partial M,, UCM 58957 MI. Specimens
ofAnemorhysis savagei, n. sp., that were examined are listed under type and hypodigm
below. Recently reported material of Trogolemur 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 variation exhibited among these species.
Data was entered in a MacClade 3.01
(Maddison and Maddison, 1992) file and analyzed 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 exhaustive search option in PAUP. A strict
consensus of all equally parsimonious alternatives was then computed.
Estimating body size and dietary
adaptation
Body size is a crucial aspect of a mammal’s adaption and can influence its dietary
regime. For example, a s 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-
TABLE 2. Characters a n d 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
separated; b = close together
9. P, paraconid size: a = absenthmall; b = large
10. P4 paraconid height: a = low (lower than protoconid);
b = high (nearly as high as protoconid
11. P, entoconid-hypoconid spacing: a = close together;
b = far apart; c = very far apart
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 a t anterior aspect of talonid (anterior
breadth (at conjunction of cristid obliqua and
posterior trigonid WallYposterior width
(entoconid-hypoconid);a = s . 7 2 ; 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,
(P,/P, S 5 5 )
dominantly frugivorous or insectivorous but
is too small to have been predominantly folivorous.
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 between 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 equations) or of particular subsets (e.g., tarsioid
equations), It is not clear which of the proposed estimates are most appropriate for
predicting body weight in omomyids. Gingerich (1981) notes that extant tarsiers and
other insectivorous and carnivorous mam-
present
absent
unknown (1/2)
Teilhardina
americana
Tetonoides
pearcei
1*>>12
present
P, paraconid presence
present
X
high
P, paraconid position
P, entoconid
absentltrace
absentltrace
two roots
P, root number
two roots
P4 root number
two roots
two roots
P, paraconid-metaconid widely separated close together
large
P, paraconid size
small
high
P, paraconid height
low
close
P, entoconid-hypoconid close
distance
narrow
P, talonid breadth
narrow
convergent
Molar cusp wall
convergent
orientation
narrow
M,: anterior breadth of narrow
talonid basin
Body size
135 g
135 g
M, nearly = or > M,
M,<cM,
M, length/M, length
M, shape
nearly square
nearly square
P3/p4 area
P, < P,
P,<P,
I,:I, size
P, presence
x indicatesnot applicable.
18
17
15
16
14
12
13
1
2
3
4
5
6
7
8
9
10
11
- Character
of
wide
vertical
wide
wide
<35 g
nearly square
nearlyequal
>35 g
unknown
nearly square
unknown
>35 g
Unknown
<35 g
unknown
nearly square nearly square
P3<P,
unknown
wide
wide
wide
vertical
wide
vertical
Unknown
unknown
unknown
Unknown
unknown
unknown
narrow
vertical
I, > > I,
present
absent
unknown
unknown
Unknown
Unknown
Unknown
Anemorhysis
pattersoni
two roots
two roots
widely separated
small
low
far
Anemorhysis
wortmani
I, 3> > I,
absent
present
low
X
small
absenvtrace
one root
two roots
two roots
two roots
two roots
close together close together close together
small
large
large
low
high
low
far
very far
far
Anemorhysis
sublettensis
dental characters in taxa discussed in text
Anemorhysis
sauagei
TABLE 3. Distribution
wide
vertical
<35 g
narrow
nearly equal
Unknown
P, < c P,
135 g
M, nearly = or > M,
nearly square
wide
wide
vertical
large
two roots
two roots
widely separated
small
low
far
wide
X
large
one root
one root
widely separated
small
low
far
X
absent
?absent
absent
I, > > > I,
Tmgolemur
myodes
unknown
I1 > 1,
Anemorhysis
natronensis
327
NEW EARLY EOCENE PRIMATE
TABLE 4. Shear quotients and body size estimates of extant small-hodied prosimians
and omomyid species discussed in text'
Species
Perodicticus potto
Galago crassicaudatus
Euoticus elegantulus
Anemorhysis pattersoni
Trogolemur myodes
Anemorhysis sublettensis
Galago alleni
Teilhardina amerieana
Anemorhysis natronensis
Anemorhysis wortmani
Loris tardigradus
Anemorhysis savagei
Tetonoides pearcei
Galagoides demidovii
Arctocebus ealabarensis
Galago senegalensis
N
LM2
Shear
ratio
8
6
6
1
3
1
7
5
1
2
6
2
5
3.41
3.76
2.36
1.75
1.58
1.59
2.81
1.70
1.59
1.69
2.88
1.69
1.65
1.94
3.56
2.17
1.54
1.69
1.86
1.89
1.89
1.92
1.95
1.97
1.98
2.00
2.00
2.09
2.10
2.13
2.18
2.46
_______.
8
6
7
Body size
(grams)
Diet
FIG
FIG
G
(F)
iF)
iF)
F
FA
(M)
(FA)
I
(1)
(1)
1
I
I
850-1,600
1,000-2,000
270-360
42 (144)
24 (78)
20 (70)
19G340
40 (136)
28 (93)
38 (130)
270-350
27 (87)
27 (91)
45-90
150-270
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 CharlesDominique (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.
mals have relatively large cheek teeth for
their body size. He groups omomyids with
modern tarsiers in the Tarsioidea and proposes 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 primarily 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 recovered 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 presented 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 distinguished 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 species-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; B o w
and Rose, 1987,1991).Washakie Basin omomyids and adapids were apparently equally
diverse. The presence of the anaptomor-
B.A. WILLIAMS AND H.H. COVERT
328
phine Trogolemur in the Lostcabinian (lateearly 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 having well-developed medial and lateral postprotocristids. None of these specimens preserve the antemolar dentition, so it is not
possible at present to determine the degree
of premolar reduction or incisor enlargement 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
a n extremely rare taxon (Bown and Rose,
1984). Four of the taxa represent new species.
Anemorhysis sauagei, n. sp., described below, is structurally transitional between
Teilhardina americana, the earliest known
and possibly most primitive North American omomyid (Bown, 1976,1979; Bown and
Rose, 1987), and more derived species of
Anemorhysis and Trogolemur. Two specimens 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.
1 4 ; Table 5; Appendices A-C).
Type species
Paratetonius? sublettensis Gazin, 1952.
Included species
A. sublettensis, A. pattersoni, A. wortmani, A. natronensis, A. savagei, n. sp.
Age and geographic distribution
Wasatchian to earliest Bridgerian LhL4
(early-middle Eocene) of Wyoming.
Fig. 1. Tetonoides and Anemorhysis, SEM photographs, occlusal views. Left: Tetonoides pearcei, holotype USNM 22426. Right dentary fragment preserving
P4-M3. R i g h t Anemorhysis pattersoni, USGS 476, holotype. 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 under molars. P, larger relative to P, than in
Trogolemur. Compared with Teilhardina
and Tetonoides, third and fourth premolar
talonid basins are broader and oblique cristids more buccally placed; entoconids farther from hypoconids relative to breadth of
talonid. P,, talonids longer than in Arapahovius, Tetonius, and Teilhardina crassidens. 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 unlike other anaptomorphines in having (especially on M,) broad trigonid basins, vertical
(non-convergent) cusp slopes, and talonid
basins that are buccolingually broad a t 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 preserving alveoli of I,-P,, and crowns of P,-M,. Right, top:
Anemorhysis natronensis holotype, CM 41137. Left dentary fragment preserving root of I,, partial crowns of I,
(partial), C,, P.,
Right, middle: Anemorhysis wortmanz holotype, USGS 6554. Right dentary fragment
preserving root of I,, P,-M,. Right, bottom: Anemorhysis sublettensis holotype, USNM 19205. Left dentary
fragment preserving crowns of P,-M,. Bar equals approximately 3.4 mm.
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 expanded than in Teilhardina or Tetonoides
but less so than in Trogolemur. Molar
enamel smooth, unlike Arapahouius, Strigorhysis, and some Absarokius.
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
UCM 56410, right mandibular body with
P,-M,, and alveoli for 11-P2.
The type and other UCM specimens are
from UCM Locality 88040, early Eocene,
Wasatch Formation, Washakie Basin, Wyoming. This locality is assigned to the Lysitean subage of the Wasatchian Land Mammal 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 Formation, Wind River Basin, Wyoming.
Hypodigm
Etymology
The type specimen and UCM 56413 right
isolated MI; UCM 56899, left MZp3;UCM
56900, left P,-M,; UCM 60914, right MI-,;
Named for Dr. Donald E. Savage in honor
of his contributions to understanding
Eocene faunas.
Anemorhysis savagei, sp. nov.
(Figs. 2 4 ,Appendices A-C).
Holotype
B.A. WILLIAMS AND H.H. COVERT
330
Fig. 3. Anernorhysis sauugei, n. sp., SEM stereophotograph,occlusal view. UCM 56410. Right P, = M,.
Bar equals approximately5.00 mm.
Diagnosis
Description
Differs from Anemorhysis wortmani, A.
natronensis, and probably A. pattersoni in
retaining P,. Molars and third and fourth
premolars absolutely smaller than corresponding teeth of A. pattersoni and A. wortmani. 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.
Among anaptomorphines, A. savagei is
most similar to other species of Anemorhysis, Trogolemur myodes, Tetonoides pearcei,
and Teilhardina americana. Several pertinent characters and their states are shown
in Table 2.
Incisor and canine roots or alveoli are
unknown for A. sublettensis and A. pattersoni. 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. Bottom: UCM 62682. Right dentary fragment preserving root of I,, alveoli of 12-P2,a n d crowns of P,-M,. B a r
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),a n d B e a r d e t a]. (1992). Oldest
1; youngest = Br.2. Abbreviations as in Table 1.
americana [Bown and Rose, 19871) but
less enlarged than in A. wortmani or Trogolemur myodes. P, is absent. The P, is singlerooted, and the alveolus is smaller than that
for P,. The relative sizes of Il-P2 (based
=
Wa-
on their alveoli) may be expressed in a
series from largest to smallest as:
I1 >> C, > P, > I,, where > means slightly
larger and >> means much larger. The
ratio is like that seen in Tetonoides pearcei.
332
B.A. WILLIAMS AND H.H. COVERT
The P, and P, trigonids are less molariform than in A. wortmani, A. sublettensis (P,
unknown), and T. pearcei. The third premolar has two roots and is a simple tooth dominated by a protoconid, without paraconid or
metaconid. A very faint cristid obliqua is
buccally oriented as in other species of Anemorhysis. 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 expression 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 hypoconid and small entoconid; these cusps are
far apart relative to the breadth of the talonid, as in other Anemorhysis; however, the
talonids of P, are relatively less buccolingually 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 absolutely smaller than in all species of Anemorhysis 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 specimens of A. sauagei. Its paraconid is closely
appressed to the metaconid and is strongly
lingual. Compared with Teilhardina americana, 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 characters in which these taxa differ. A Student’s t-test demonstrates that A, savagei
has an absolutely narrower M, ( t = -4.06,
P C .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 Anemorhysis 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 sample of the extremely rare notharctine adapid
Copelemur australotutus (Beard, 1988; Covert, 1990). Turtle Graveyard is situated approximately 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 pearcei. 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 difficult to assess their relative ages due to lack
of precision in interbasinal faunal correlation.
Phylogeny and character evolution
A parsimony analysis using a PAUP exhaustive search yields two trees of equivalent minimum parsimony. With uninformative 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 Anemorhysis 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 talonid basins but has apparently derived premolar trigonids featuring a distinct P, paraconid and a large, high P, paraconid.
2. Primitive Anemorhysis is specialized
from Teilhardina by having more complex
premolar talonids (broader basins) and derived molar morphology (non-convergent
cusp walls, anteriorly broad talonid basins).
Anemorhysis sauagei is the most primitive
NEW EARLY EOCENE PRIMATE
333
Teilhardina americana
Tetonoidespearcei
Anemorhysis savagei
Anemorhysis sublettensis
Anemorhysis wortmani
Anemorhysispattersoni
Anemorhysis natronenesis
Trogolemur myodes
Fig. 5. Strict consensus phylogeny.
member of the Anemorhysis clade, retaining
P, and premolariform P3+
3. More derived Anemorhysis show further 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 molarization of the premolars must be a parallelism
in Tetonoides and some Anemorhysis.
4. Anemorhysis sublettensis and A. wortmani 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 exceptionally long and wide talonid basin.
5. The source of Trogolemur comes from
within the Anemorhysis clade, but exact sister-group relationships are unclear. Trogolemur has the molar synapomorphies that
unite species ofAnemorhysis but is more derived than any species of that genus in having 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 demonstrates that there is an unresolved trichotomy 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
metaconid).
6. Anemorhysis natronensis is probably
the species least phenetically similar to
other members of the genus and is autapomorphic in several features, such as narrow
molars (especially M,), very large P, entoconid, 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 omomyids vary, such as molarization of the premolars and enlargement of the I, relative to the
I,, is difficult to assess. Selecting the outgroup in a cladistic analysis (thus determining polarity) can profoundly affect the interpretation of cladogenesis. By rooting the
network with Teilhardina americana we are
presuming that the dental morphology of
that taxon represents the morphology primitive for anaptomorphines. It has been accepted 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). Therefore, 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., Szalay, 1976; Gingerich, 1981; Bown, 1976,
1979; Bown and Rose, 1987).
It is possible that anaptomorphines are
not the most primitive members of Omomyidae. 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 a s Teilhardina (Gazin, 1958; Rose and
Bown, 1991).
The P, of Steinius vespertinus has a low
paraconid and only slightly developed metaconid, whereas the P, trigonid of Loveina
zephyri is well developed. Unfortunately,
the P, of the older and possibly more primitive species of Loueina, L. minuta, is unknown. The P, trigonids of the anaptomorphines under study here show considerable
variation (Table 3; Fig. 2). In T. belgica and
T. americana, the P, paraconid and metaconid are usually poorly developed, but in
later-occurring T. crassidens those cusps are
distinct and well developed.
If anaptomorphines are the most primitive omomyids, and Teilhardina is representative of the anaptomorphine morphotype,
then simple premolar trigonids are primitive for the family. I t is unclear what the
omomyine morphotype might be with regard
to premolar trigonid structure. Understanding the omomyid morphotype is further complicated 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 evidence that washakiins may be a primitive
outgroup to all other omomyids, and possibly all other omomyids, and possibly all
other primates (Williams and Kay, 1992).
Additionally, Steinius may be a n anaptomorphine rather than a n omomyine (Bown
and Rose, 1984, 1987; Williams and Kay,
1992). The development of complex premolars apparently occurred in more than one
group of anaptomorphines (e.g., Teilhardina, Tetonoides, Anemorhysis).
The polarity of central incisor enlargement in omomyids similarly has been debated. Small, equisized lower incisors were
probably primitive for primates (Cartmill
and Kay, 1987; Szalay et al., 1987; Gingerich e t al., 1991; Covert and Williams, 1991b)
and possibly for anaptomorphines as seen in
Teilhardina belgica (Gingerich, 1977). However, 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 a s
Tetonoides and Arapahovius (contra observations 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 indicate phylogenetic relatedness. Until the fossil 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 tarsioid estimate is used, it is apparent that
these anaptomorphines were well under
Kay’s threshold of 500 grams and were probably less than 50 grams (Table 4). Smith
(1993) noted that some body weight estimates 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 anaptomorphines 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 a s the smallest living
primates: the galagine Galagoides, the
cheirogaleid Microcebus, and the platyrrhine Cebuella. Anemorhysis savagei is one
of the smallest omomyids known. If A . sauagei evolved from the larger-bodied Teilhardina americana and represents the stem
member of its genus, it appears that Anemorhysis underwent some size reduction followed by size increase in some taxa (A.
pattersoni, A. wortmani).
NEW EARLY EOCENE PRIMATE
1
I
I
- I
I
1
1
’
more fruit and gum
I
1
I
I
I
335
I
I
more insects
I
W‘Trogolemur myodes (3)
I
Anemorhysis pattersoni (1)
I
Anemorhysis sublettensis (1)
I-W Teilhardina americana (5)
I@
Anemorhysis natronensis ( 1 )
Anemorhysis wortmani (2)
l-+l
W Anemorhysis savagei (2)
I
F0-I Tetonoides pearcei (5)
*Perodicticus potto (8)
I
t-0-I OtoEerrb crassicaudatus (6)
A ‘Euoticuselegantulus (6)
W Galagoides alleni (7)
K)-l Loris
I
I
tardigradus (6)
Galagoides demidovii (8)
)--0--1Arctocebus calabarensis (6)
1.2
I
I
1.4
I
I
1.6
I
I
1.8
I
I
2.0
senegalensis (7)
-Galago
I
I
I
2.2
I
I
2.4
I
I
2.6
Mean shearing ratio
Fig. 6. Diets of extant lorises and galagos and suggested 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-
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).
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 shearing ratios of Anemorhysis savagei and Tetonoides pearcei fall into the range of animals such as Galagoides demidovii and
Loris tardigradus that feed primarily on in-
sects. Anemorhysis pattersoni, A. sublettensis, and Trogolemur myodes have ratios
more similar to frugivores such as Euoticus
elegantulus and Galagoides alleni. The
other taxa do not show apparent specialization 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 generally comparable. As indicated by our results
and those of Strait, it is apparent that no
anaptomorphines (we excluded Loveina
from Anaptomorphinae) possessed the trenchant shearing blades of highly insectivorous 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 insufficient material). It appears that the earliest
anaptomorphines were opportunistic, mixed
feeders and that the dietary specializations observable in modern primates did not
occur until considerably later in their evolution.
It is interesting to note that there appears
to be litte association between trends in incisor 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 incisor enlargement while maintaining an insectivorous diet. Perhaps incisor enlargement 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 assemblage of fossil primates during the early
Eocene and documents the presence of several rare middle and late Wasatchian taxa.
This new material is especially noteworthy
because it samples the middle-late Wasatchian transition more completely than do Samples from either the Bighorn or Wind River
Basins. The Washakie Basin sample thus
increases our understanding of the taxonomic 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 premolar and molar features characteristic of
other species of Anemorhysis and is clearly
referable to this genus. A phylogenetic analysis of this and other small-bodied anaptomorphines indicates that this taxon is primitive relative to other members of the genus.
A. savagei is structurally intermediate between 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 renders further resolution of phylogenetic relationships 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. Recovery 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 associated with the extremes of dietary specialization known in extant primates. Incisor enlargement 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 manipulation.
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 specimens. We are also grateful t o Tom Bown,
Ken Rose, Peter Robinson, and Chris Beard
for many insightful conversations on anaptomorphine 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
Richard Kay. Leonard Krishtalka and Chris
Beard, Carnegie Museum, Tom Bown,
USGS, and Robert Emry, USNM, generously allowed us to examine specimens in
their care. Funding for this work has been
provided by the University of Colorado Museum W. Van Riper and W.H. Burt Fund
grants.
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Swofford D (1991) PAUP: Phylogenetic Analysis Using
Parsimony, Version 3.0s. Champaign, IL: Illinois Natural History Survey.
Szalay F (1976) Systematics of the Omomyidae (Tarsiiformes, Primates):Taxonomy, Phylogeny and adaptations. Bull. Am. Mus. Nat. Hist. 156:157450.
Szalay FS, Rosenberger AL, and Dagosto M (1987) Diagnosis and differentiation of the order Primates. Yrbk.
of Phys. Anthropol. 30t75-105.
Williams B, and Kay R (1992) Phylogenetic analysis of
Eocene primates suggests Omomyidae is not a monophyletic group. In: Proceedings of the XTVth Congress
of the International Primatological Society, p. 286.
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Anernorhysis from the Washakie Basin, Wyoming.
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APPENDIX A. Measurements (mmi o f teeth o f Anemorhvsis savapei. S D . now.
Specimen
UCM 56410
UCM 62682
UCM 56900
UCM 56899
UCM 56413
UCM 60915
UCM 60914
CM 39654
CM 28915
P,W
-~
0.90
0.95
-
P,L
P,W
P,L
M,W
MIL
M,W
M,L
M,W
M,L
1.15
1.10
1.05
1.10
1.25
1.20
1.30
1.30
1.35
1.35
1.50
1.75
1.70
1.65
1.50
1.35
1.65
1.70
1.25
-
-
1.90
-
-
-
-
-
1.50
1.55
1.15
1.95
1.15
1.50
1.45
1.45
1.35
1.40
1.35
1.70
1.75
1.75
1.75
1.75
1.25
2.00
-
-
-
-
1.30
1.40
-
-
-
1.50
1.45
1.50
-
-
1.75
1.80
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.
1.35
.27-1.43
.07
8
2.10
1.94-2.19
.08
8
11
1.10
1.05-1.15
.03
11
1.42
1.2Ck1.55
.ll
18
1.39
1.25-1.55
.09
18
1.60
1.451.75
.07
26
1.61
1.40-1.80
.10
26
1.96
30-2.10
.09
27
1.68
.40-1.85
.10
27
1.91
.8Ck2.10
.09
2.12
2.10-2.20
.05
4
.98
.95-1 .OO
.03
4
1.23
i.oai.30
.lo
5
1.20
i.oai.30
.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
-
-
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
.04
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
1.10, 1.15
-
2
.go, .95
-
2
Anemorhysis
sauagei
-
-
-
-
-
-
-
-
-
-
1.80
1.80, 1.90
1
-
2
-
1.30
-
1
1.55, 1.65
2
-
1.80
1.90, 1.95
-
-
-
1.90, 1.95
2
-
1.60, 1.85
-
2
-
1.90, 2.00
-
-
-
1.70
-
-
1
1.40
-
-
1
1.65
-
1
1
-
2
2
-
1.30
1.40
-
1
1.60
1
-
2
1.40
-
-
1
1.10
-
Anemorhysis
sublettensis
1.65, 1.75
2
-
-
1.50
-
1.40, 1.70
1
-
2
1.50, 1.60
2
-
1.30
-
1.35, 1.40
1.30, 1.40
-
-
-
-
1
2
-
-
1.40
-
1
1.40
1
-
-
1.15
-
1
Anemorhysis
natronensis
1.20
1
Anemorhysis
wortmani
-
2
-
Anemorhysis
pattersoni
APPENDIX B. Summarv statistics for anavtomorvhines discussed in text '
Tetonoides
pearcei
~~
2.04
2.00-2.10
.05
4
1.30
1.21-1.40
.08
4
1.67
1.50-1.75
.lo
5
1.52
1.3g1.60
.10
5
1.57
1.40-1.85
.19
5
.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
~
Trogolemur
.~ myodes
3
.84
.80-.90
.05
3
1.00-1.11
1.07
'Measurements are in millimeters. Specimen data from following sources: Teilhardina americana from Bown and Rose (1987:31; p 3 - 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.
OR
SD
n
mean
OR
SD
mean
OR
SD
n
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
Teilhardina
americana
M,
M,
M,
P,
1.22
1.14
.82
.86
1.15
(11)1.56
(18)2.24
(26) 3.16
(27) 3.21
( 8 ) 2.87
.70
.71
.98
1.12
1.29
(5) 1.57
(8)2.42
(6) 2.50
(3)2.38
.66
.65
.97
1.05
1.22
1.15
1.24
1.16
.77
.84
( 5 ) 1.74
(9) 2.50
(9) 2.67
(4)2.86
.70
.70
.94
.93
1.26
1.21
1.22
1.07
.83
.84
-
(2) 1.04
(4)1.21
1.15
1.15
1.12
.79
.79
-
-
.63
1.00
-
(2) 2.02
(213.32
(2) 3.32
Anemorhysis
puttersoni
Anemorhysis
sauugei
-
1.22
1.15
1.29
1.38
.83
.93
.86
.81
-
1.17
1.13
1.20
1.16
.83
.77
1.08
-
.79
.69
1.04
-
(11 2.52
(1)2.34
(1)1.61
(1)1.95
Anemorhysis
natronensis
(1)1.68
(2) 2.13
(2) 3.08
(2) 2.96
Anemorhysis
wortmani
APPENDIX C. Areas and ratios for anaptomorphines discussed in text
Tetonoides
pearcei
’Measurements are in millimeters. Areas are in square millimeters
M I N
M2 m
P, m
1L
P. wm. w
p4m
Area P,
Area P4
Area M,
Area M,
Area M,
Area P,/Area
Area P,/Area
Area MJArea
Area MJArea
p3m
Dimension
Teilhardina
americana
1.27
1.27
1.21
.85
.85
-
.90
.72
-
-
(1)2.15
(1) 2.38
(1)1.54
-
Anemorhysis
sublettensis
(I) 9 2
(1)1.77
(2) 2.27
(3) 2.60
(2) 2.56
.52
.77
.87
1.02
1.34
1.15
1.32
1.10
.83
.95
Trogolernur
myodes
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