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New postcranial elements for the earliest Eocene fossil primate Teilhardina belgica

Authors:
New postcranial elements for the earliest Eocene fossil primate Teilhardina belgica
Daniel L. Gebo
a
,
*
, Thierry Smith
b
, Marian Dagosto
c
a
Department of Anthropology, Northern Illinois University, DeKalb, IL 60115, United States
b
Direction Earth and History of Life, Royal Belgian Institute of Natural Sciences, B-1000, Brussels, Belgium
c
Department of Cell and Molecular Biology, Feinberg School of Medicine, Northwestern University, Chicago, IL 60611, United States
article info
Article history:
Received 1 July 2011
Accepted 14 March 2012
Available online 15 June 2012
Keywords:
Omomyidae
Primate origins
Positional behavior
Ankle
Knee
Dormaal
abstract
Teilhardina belgica is one of the most primitive fossil primates known to date and the earliest haplorhine
with associated postcranials, making it relevant to a reconstruction of the ancestral primate morphotype.
Here we describe newly discovered postcranial elements of T. belgica. It is a small primate with an
estimated body mass between 30 and 60 g, similar to the size of a mouse lemur. Its hindlimb anatomy
suggests frequent and forceful leaping with excellent foot mobility and grasping capabilities. It can now
be established that this taxon exhibits critical primate postcranial synapomorphies such as a grasping
hallux, a tall knee, and nailed digits. This anatomical pattern and behavioral prole is similar to what has
been inferred before for other omomyids and adapiforms. The most unusual feature of T. belgica is its
elongated middle phalanges (most likely manual phalanges), suggesting that this early primate had very
long ngers similar to those of living tarsiers.
Ó2012 Elsevier Ltd. All rights reserved.
Introduction
The fossil primate Teilhardina is regarded as one of the most
primitive of all early Eocene fossil primate genera recovered to date
(Szalay, 1976;Szalay and Delson, 1979;Bown and Rose, 1987;
Gunnell and Rose, 2002;Ni et al., 2004;Rose, 2006;Smith et al.,
2006;Beard, 2008). The European, North American and Asian
Teilhardina species are known primarily from dental elements,
leaving its body anatomy largely unknown. Until recently (Rose
et al., 2011), the lone exception was the type species Teilhardina
belgica to which Teilhard de Chardin (1927) referred three post-
cranial elements: a calcaneus, a talus, and a distal metacarpal.
Simpson (1940) commented on the calcaneus pictured by Teilhard
de Chardin and compared it with one attributed to Hemiacodon,
indicating that both were very similar in overall morphology.
Szalay (1976) would later show that the talus and metacarpal
gured by Teilhard de Chardin do not belong to a primate but
actually are a rodent-like astragalus and a frog humerus, respec-
tively. Szalay (1976) discovered and commented on additional
tarsal elements of T. belgica from Dormaal, identifying these tarsals
from collections housed at the Royal Belgian Institute of Natural
Sciences (Brussels, Belgium). From these same collections, and
some new ones, we have been able to recover additional postcranial
elements of T. belgica from the locality of Dormaal including two
tali, two calcanei, a rst metatarsal, a distal femur, and four
phalanges. These new elements are described below.
The earliest Eocene locality of Dormaal is located in western
Belgium and is part of the southern North Sea Basin. The Dormaal
fossil mammal assemblage is famous for its richness and diversity
(Smith and Smith, 1996) and for having yielded the earliest Euro-
pean members of several modern mammal orders including
artiodactyls, creodonts and carnivorans (Smith et al., 1996;Smith
and Smith, 2001,2010). Fossil mammals from the Dormaal
locality (Brabant, Belgium) come from continental sand deposits of
the uvio-lagoonal Tienen Formation, near the PaleoceneeEocene
boundary in Belgium (Steurbaut et al., 1999;Smith, 1999,2000;
Smith and Smith, 2003;Smith et al., 2006). The Dormaal mammal
fauna was chosen as reference level MP7 of the mammalian bio-
chronological scale for the European Palaeogene (Schmidt-Kittler,
1987).
In his original description of the primate from Dormaal, Teilhard
de Chardin (1927) named the species Omomys belgica. Subsequent
to the original description, Simpson (1940) argued for a generic
Abbreviations: AMNH, American Museum of Natural History, New York, USA;
BFI, University of Montpellier, France; CM, Carnegie Museum, Pittsburgh, USA;
DMNH, Denver Museum of Natural History, Denver, USA; GMH, Geiseltal Museum
Halle, Halle, Germany; IVPP, Institute of Vertebrate Paleontology and Paleoan-
thropology, Beijing, China; RBINS, Royal Belgian Institute of Natural Sciences,
Brussels, Belgium; UCM, University of Colorado Museum, Boulder, Colorado; UM,
University of Michigan, Ann Arbor, Michigan; USGS, United States Geological
Survey, Baltimore, Maryland; Zurich NR, A/V, University of Zurich-Irchel, Zurich,
Switzerland; UW, University of Washington, Burke Museum, Seattle, Washington;
YPM, Yale Peabody Museum, New Haven, Connecticut.
*Corresponding author.
E-mail addresses: dgebo@niu.edu (D.L. Gebo), thierry.smith@naturalsciences.be
(T. Smith), m-dagosto@northwestern.edu (M. Dagosto).
Contents lists available at SciVerse ScienceDirect
Journal of Human Evolution
journal homepage: www.elsevier.com/locate/jhevol
0047-2484/$ esee front matter Ó2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jhevol.2012.03.010
Journal of Human Evolution 63 (2012) 205e218
distinction from Omomys, naming Teilhardina after its discoverer.
T. belgica is generally considered to be more primitive than species
of Teilhardina from western North America (Szalay, 1976;Bown and
Rose, 1987;Smith et al., 2006;Beard, 2008) but more derived than
the recently discovered T. asiatica from China (Ni et al., 2004). We
view Teilhardina taxonomically as a basal member of the Anapto-
morphinae within the Omomyidae, a fossil primate family that also
includes the Omomyinae. The Omomyidae, Microchoeridae, and
Tarsiidae represent separate haplorhine families within Tarsii-
formes. We follow the taxonomic scheme of Szalay and Delson
(1979) and Gunnell and Rose (2002) with the exception of
elevating the Microchoeridae to the level of family. In this paper, we
restrict our use of the taxon Primates to the extant and fossil
Strepsirhini and Haplorhini, excluding potential primate sister taxa
such as Plesiadapiformes, Dermoptera, or Scandentia.
Materials and methods
We examined specimens from the site of Dormaal housed at the
RBINS (Brussels, Belgium), discovering several new postcranial
elements that can be allocated to T. belgica (listed in Table 1), the
only recognized primate species from this locality (Teilhard de
Chardin, 1927;Smith and Smith, 1996). The new specimens come
from miscellaneous bone boxes of undetermined specimens
collected by screen washing during four eld campaigns organized
by the RBINS at Dormaal between 1924 and 1949. These excava-
tions have all been done in the same stratigraphic horizon in the
western wall of a short sunken road, site of the classical Dormaal
locality that exposed the uviatile deposits of the base of the Tie-
nen Formation (Rutot and Van den Broeck, 1884;Casier, 1967;
Smith and Smith, 1996; map sheet 33/5-6; GPS coordinates: N
50
47
0
50
00
E05
04
0
48
00
).
Figs. 1e3illustrate the linear measurements and angles that
were measured using calipers for the larger specimens and
a microscopic reticle for the smallest elements. The measurements
follow Dagosto (1986),Gebo et al. (1991,2001), and Dagosto et al.
(1999). Comparative data for other extinct and extant primates
are also drawn from these publications, where further information
regarding these specimens is contained. The Appendix lists the
comparative fossil sample.
Results: descriptions and comparative anatomy
Body size
The absolute dimensions of all of the postcranial elements now
known for T. belgica indicate that it was very small (Table 2).
Compared with Microcebus berthae, the smallest living primate
(24e38 g; Rasolarison et al., 2000), Teilhardinas talar length,
calcaneal length, and proximal phalanx length are all smaller; mid-
trochlear talar width, calcaneal width, and calcaneocuboid width
are similar to or slightly larger; while the rst metatarsal, distal
femur, and middle phalanx measures are all larger. Compared with
the slightly larger mouse lemurs, Microcebus griseorufus or
Table 1
List of new postcranial elements of T. belgica.
Catalog number Identication Preservation
IRSNB Vert 26857-01 Talus Complete
IRSNB Vert 26857-02 Talus Broken
IRSNB M1237 Calcaneus Complete
IRSNB Vert 26857-03 Calcaneus Broken
IRSNB M1262 First Metatarsal Broken
IRSNB M1263 Femur Broken
IRSNB M1264 Proximal Phalanx Complete
IRSNB M1265 Proximal Phalanx Complete
IRSNB M1266 Middle Phalanx Complete
IRSNB Vert 26857-04 Middle Phalanx Broken
Figure 1. Talar and calcaneal measurement gures for Tables 3 and 6.
Figure 2. First metatarsal and phalangeal measurement gures for Tables 10 and
12e15.
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218206
Microcebus murinus (50e70 g; Génin, 2008;Rakotondranary et al.,
2011), Teilhardina is smaller or equal in absolute dimensions,
bracketing it between the smallest and medium sized mouse
lemurs.
The dimensions of Teilhardinaslimb bones are smaller than
most other currently described omomyids (e.g., Tetonius,Nannopi-
thex,Shoshonius, and Washakius; Tables 4,8,11), all of which are
estimated to be 40e100 g on the basis of tarsal size (Dagosto and
Terranova, 1992;Thalman, 1994;Dagosto et al., 1999). Based on
these comparisons and regressions of tarsal dimensions (Dagosto
and Terranova, 1992), we estimate that T. belgica weighed
between 30 and 60 g. This is reasonably consistent with estimates
based on tooth size (63 g; Conroy,1987; prosimian regression; 30 g;
Gingerich, 1981; tarsier model).
Tali
Table 3 lists the RBINS talar specimens and their measurements
(see Fig. 1), while Tables 4 and 5 provide comparative information
for other omomyids and microchoerids. The talus of T. belgica
possesses a talar neck and head that are quite elongated relative to
the talar body (Table 5;Fig. 4). The talar neck is about 50% of total
talar length (talar neck/talar length ratio, Table 5)inT. belgica,
similar to the ratio of most other omomyids. Necrolemur has
a slightly shorter neck. The elongated talar head and neck region is
partly related to the distal elongation of the calcaneus. The talar
neck in T. belgica is marked by an oblique ridge for attachments of
the anterior tibial-talo-navicular ligament, which is part of the
talonavicular joint capsule. This ridge is observable on tali of other
omomyids as well.
The talar head is oval in overall shape in T. belgica when viewed
distally (Fig. 4). The talonavicular facet is extended proximally for
a long distance along the medial side of the talar head. Compared
with its height, the talar head of T. belgica is wider (talar head width
relative to head height) than most of the other taxa in Table 5 , being
closest to Nannopithex in this ratio. In both of these taxa, the talar
head is considerably wider than the other omomyids and Necrolemur.
The talar trochlear region is relatively at with only slight
elevation of the medial talar rim relative to the lateral rim. The
medial talar rim of anaptomorphine omomyids, like that of Teil-
hardina, is rounder and extends further distally than those of
omomyines (Dagosto, 1993). T. belgica does not exhibit a squatting
facet as observed in Teilhardina brandti (Rose et al., 2011). Mid-
trochlear width as a percentage of trochlear length is greatest in
T. belgica and narrowest in Washakius and Nannopithex (Table 5).
The posterior section of the trochlear facet ends proximally, high on
the back of the talus, making a distinct oblique but horizontally
oriented facet line. This facet leaves a gap between the posterior
edge of the trochlear facet and the posterior trochlear shelf. This
pattern of trochlear facet anatomy is also observed in T. brandti (see
Fig. 10d in Rose et al., 2011). A small posterior trochlear shelf follows
below this region in T. belgica.Szalay (1976) noted a well developed
posterior trochlear shelf in one talus of T. belgica but not in Hemi-
acodon or Tetonius. In fact, all omomyid tali, including those of
T. belgica, possess a trochlear shelf although it is small compared
with those of most adapiforms (Dagosto, 1993). T. brandti exhibits
a more prominent trochlear shelf (Rose et al., 2011) relative to that
of T. belgica. The microchoerid Necrolemur possesses a more
prominent trochlear shelf than that of omomyids (Godinot and
Dagosto, 1983). The posteroplantar section of T. belgica tali
possesses a centrally positioned groove for exor bularis,
a common condition among omomyids, microcheorids, and
anthropoid primates (Beard et al., 1988;Gebo, 1988).
Both the medial and lateral facets of these tali suggest
a symmetrical tibiobular mortise like that of other omomyids and
anthropoids (Dagosto et al., 2008). The lateral talobular facet is
steep-sided with a pointed plantar process, a common condition in
other haplorhine primates (Beard et al., 1988;Gebo,1988). Posterior
to the lateral talobular facet is a well developed fossa for the
posterior talobular ligament. Compared with other omomyids and
microchoerids, Teilhardina has a fairly high talar body (relative to
mid-trochlear width). The talotibial or medial talar facet is large
relative to the medial surface, indicating a long tibial malleolus.
Posterior to the medial talotibial facet exists ridging and depres-
sions for a strong posterior talotibial ligament, a part of the deltoid
ligament.
The plantar side of the talus has a wide and elongated plantar
facet that contacts the surface of the anterior calcaneal facet. The
plantar side shows a well curved posterior facet that articulates
with the posterior calcaneal facet, a part of the subtalar joint.
Figure 3. Distal femur measurement gure for Table 11.
Table 2
Comparison of Teilhardina dimensions to those of other small extant and extinct
primates.
Teilhardina
belgica
Microcebus
berthae
Microcebus
griseorufus/
murinus
Shoshonius
cooperi
Body mass 30e60 24e38 g 50e70 g Est. 60e90 g
Talar length 4.10e4.20 4.47 5.62 5.24-5.66
Mid-trochlear width 1.78e1.86 1.69 1.86 2.01-2.09
Calcaneal length 6.81e7.39 8.94 10.15 9.76-9.82
Calcaneal width 1.96e2.70 2.25 2.56 2.97-3.53
Calcaneocuboid width 1.57e1.82 1.44 1.73 2.29-2.34
First metatarsal
proximal width
2.15 1.73 2.59
First metatarsal
proximal height
2.40 1.87 3.30
Distal femur bicondylar
width
4.20 3.79 4.47 5.05
Distal femur lateral
condylar height
4.75 3.91 4.45 5.63
Proximal phalanx length 4.40 5.00
Middle phalanx length 5.60 4.10
Comparable dimensions for other omomyids are given in Tables 4, 8 and 11.
Measurements are in mm.
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218 207
Calcanei
There are currently ten known calcanei of T. belgica (Table 6;
Fig.5), six of which are complete. Three are very similar in overall
length ranging from 6.52 to 6.98 mm, but two calcanei are
noticeably longer (7.21 and 7.39 mm; Fig. 6). This might suggest
either the presence of more than one omomyid species at Dormaal
or sexual dimorphism in body size in Teilhardina. Despite the
presence of slight but signicant dimorphism in skull length, body
mass, and pelvic dimensions in M. griseorufus (Kappeler, 1990;
Jenkins and Albrecht, 1991;St. Clair, 2007) and of body mass in
most species of Tarsius (Yustian, 2007), T-tests show no signicant
degree of dimorphism in calcaneal length or width in either of
these small extant primates (Tarsius syrichta:N¼9, length p¼0.55,
width p¼0.91; M. griseorufus:N¼15, length p¼0.39, width
p¼0.47). It thus seems unlikely that the variance exhibited by the
Teilhardina calcanei is due to sexual dimorphism.
Measures of calcaneal length variance (CV, Imax/min, R%) of the
Teilhardina sample from Dormaal are generally consistent with
single species of extant and extinct primates and are lower than in
mixed samples of primates (Table 7), indicating that a single species
hypothesis cannot be rejected (Cope, 1993). Calcaneal width,
however, shows the opposite pattern: the measures of variance are
more consistent with mixed species samples than with single
species samples. Of the ve other calcaneal measures taken, four
support a single species hypothesis and one (PACL) does not.
Although these results are not unanimous, they generally support
the interpretation that there is only a single species of Teilhardina at
Dormaal and this is consistent with interpretations of the dental
evidence (Teilhard de Chardin, 1927;Smith and Smith, 1996).
Even the largest calcaneus in this sample of T. belgica is shorter
than any of the other known omomyid specimens (Table 8).
Shoshonius cooperi and Absarokius abbotti are the closest in absolute
calcaneal length with values at 9.79 and 9.80 mm, respectively.
Even the largest calcaneus for T. belgica is smaller than calcanei
associated with S. cooperi (Dagosto et al., 1999).
First Simpson (1940) and later Szalay (1976) noted that the
calcaneus of T. belgica was distally elongated and similar to that of
Hemiacodon.Szalay (1976) concluded that this elongation is an
ancestral character of the Omomyidae. Our sixcomplete specimens
show a mean distal calcaneal elongation of 53.2% (Table 8), a value
similar to that calculated by Gebo in 1988 (53.0%). Distal elongation
relative to total calcaneal length is quite consistent across
omomyids (49.5e56.4%; Table 9), including T. belgica, and is far
shorter than the value in Tarsius or Galago (Simpson, 1940;Szalay,
1976 ;Gebo, 1988). Microchoerids have more elongated calcanei,
but none are complete enough to take a ratio.
Like the degree of distal elongation, the ratio of calcaneal width
to calcaneal length (Table 9) is also relatively consistent in omo-
myids (29.3e36.9) with T. belgica at the high end of this range. In
contrast, the more greatly elongated distal calcaneus of micro-
choerids and tarsiers result in a narrow width ratio. Both of these
bony features clearly represent family level adaptations that are
distinct relative to the Omomyidae.
Szalay (1976) further noted the presence of a distinct calca-
neocuboid pivot in omomyid calcanei, including that of T. belgica,in
contrast to Tarsius, which has a attened joint surface. The calca-
neocuboid joint of T. belgica as well as that of other omomyid and
adapiform primates is wide and tall with the pivot region located
plantarly, resembling a fan. This condition is certainly primitive for
early Eocene primates. Calcaneocuboid width to height ratios
shows T. belgica to possess the lowest ratio and therefore the nar-
rowest calcaneocuboid joint among the taxa compared in Table 9.
In contrast, Shoshonius and Hemiacodon display the widest calca-
neocuboid joints.
The anterior calcaneal facet is very round distally but proceeds
anteriorly in a narrower, thinning and pointed facet proximally. The
plantar surface of the talar neck lies upon this anterior joint. Below
this joint surface lies the sustentaculum tali, a bony shelf that is
deeply grooved on its plantar surface allowing the long extrinsic
exor tendons to proceed anteriorly toward their attachment sites
on the toes.
The posterior calcaneal facet (pcf) or the posterior astragalo-
calcaneal facet (pac) is anteroposteriorly long like that of Tetonius or
Arapahovius in contrast to more shortened facets observed in
Hemiacodon or Necrolemur. The posterior astragalocalcaneal facet
width to length ratio (Table 9)ofT. belgica is relatively high like that
of Omomys and Hemiacodon, and contrasts with the lower values
exhibited by Tetonius and Shoshonius. The ratio of posterior astra-
galocalcaneal facet length relative to total calcaneal length (Table 9)
shows T. belgica to possess the longest relative joint facet of these
comparative omomyids.
Szalay (1976) stated that no peroneal tubercle is present in
calcanei of omomyids. We disagree, but note that this structure is
much less developed in omomyids than in plesiadapiforms and
adapisoriculids (Smith et al., 2010). The widest aspect of the
Table 3
Talar measurements (mm).
Talar length Talar width Neck length Length of trochlea Mid-trochlear width Lateral height Head width Head height Neck angle
IRSNB M1235 4.23 2.51 2.01 2.09 1.86 2.30 1.89 1.27 29
IRSNB Vert 26857-01 4.10 2.51 1.97 2.04 1.78 2.06 1.16 32
IRSNB Vert 26857-02 2.32 2.00 1.68 1.24 30
Table 4
Talar comparative data.
Length Width Neck length Length of trochlea Mid-trochlear width Lateral height Head width Head height Neck angle
Teilhardina belgica n ¼2e3 4.17 2.45 1.99 2.07 1.82 2.18 1.79 1.22 30
Nannopithex abderhaldeni n ¼1 4.61 2.71 2.46 3.34 2.20 2.62 2.68 1.90 28
Tetonius homunculus n ¼3 5.41 2.90 2.96 2.78 2.07 2.75 2.00 1.15 30
Shoshonius cooperi n ¼3 5.52 2.80 2.89 2.69 2.05 2.37 2.00 1.75 30
Washakius insignis n ¼1 5.84 3.07 3.06 3.42 2.26 2.66 1.95 1.77 30
Absarokius abbotti n ¼1 6.15 3.30 3.20 3.20 2.25 28
Necrolemur zitteli n ¼1 6.65 3.47 2.99 3.49 2.87 3.16 2.76 2.46 30
Omomys carteri n ¼7 8.24 4.85 4.39 4.50 3.67 4.01 3.21 2.68 25
Hemiacodon gracilis (n¼22) 8.45 4.71 4.27 4.30 3.53 3.72 3.20 2.65 25
Taxa are ordered by mean talar length (shortest to longest). Omomys carteri material is restricted to the Covert Bridger C quarry specimens.
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218208
peroneal tubercle is situated distal to the posterior calcaneal facet
in T. belgica but this region can be more proximally positioned in
some omomyids (e.g., Hemiacodon). The peroneal tubercle of
T. belgica is more distally positioned relative to other omomyids like
Shoshonius.
The plantar surface of the T. belgica calcanei exhibits several
features of note. The posteromedial region of the calcanei attrib-
uted to T. belgica is indented with a depression that curves rst
laterally then medially in plantar view. This grooved region is the
likely attachment site of abductor digiti quinti. Anterior to this
region is the anterior calcaneal process, the attachment site for the
short and long calcaneocuboid ligaments. This site lies just prox-
imal to the calcaneocuboid joint surface. This distal calcaneal region
is angled dorsally, implying a semiplantigrade or heel-elevated foot
position when moving arboreally.
The posterior region of the calcaneus (heel) appears to be more
gracile, especially among the smaller calcanei attributed to
T. belgica, relative to omomyids like Omomys or Hemiacodon. Pro-
portionally, the heel length to calcaneal length ratio (Table 9) shows
T. belgica to be similar to that of Washakius,Absarokius, and
Shoshonius, with a smaller ratio value than Omomys or Hemiacodon.
First metatarsal
IRSNB M1262 (Fig. 7) is the proximal end of a rst metatarsal.
This specimen is broken distally, lacking most of the shaft and head.
The preserved piece measures 4.45 mm in length. The dimensions
of the rst metatarsal are similar to those of the pygmy mouse
lemur but slightly smaller than Shoshonius (Table 10). Although the
IRSNB M1262 specimen is quite small, the IVPP V 13018 haplorhine
rst metatarsal from Shanghuang (China) is still smaller in all
dimensions (Gebo et al., 2008).
The entocuneiform-rst metatarsal joint surface of IRSNB
M1262 is similar to that of other omomyids (i.e., Omomys,Hemi-
acodon,Ourayia, and Shoshonius) and to the microchoerid Necro-
lemur (Simpson, 1940;Szalay and Dagosto, 1988;Dagosto et al.,
1999;Dunn et al., 2006;Dunn, 2010). The proximal joint surface
has a narrow arc and a half cylindrical-shaped articular facet
allowing a wide arc of swing for the rst digit during grasping.
Narrow joint arcs are found among omomyids and microchoerids
but contrast with the wider arcs observed in adapiform primates.
The cylindrical-shaped articular surface for the entocuneiform-rst
metatarsal joint is commonly found among non-anthropoid
primates. In T. belgica, part of this joint surface wraps around the
lateral surface of the peroneal tubercle, making an obliquely
oriented facet-like smooth surface on the proximal aspect of the
peroneal tubercle. This smooth and oblique surface is found on
other omomyid rst metatarsals as well (Gebo et al., 1999) but is
not observed among other primates in which the proximal joint is
oriented along a mediolateral axis only. The D-C/B index (Table 10)
compares the height of the proximal joint surface relative to the
width of the rst metatarsal base. The value for T. belgica lies
between other omomyids and Necrolemur. The degree of variation
among omomyids is similar to that observed in adapiforms (e.g.,
Cantius ralstoni [0.43] compared with Cantius abditus [0.52]) (Gebo
et al., 2008).
The peroneal tubercle of IRSNB M1236 is proximodistally long as
in all primitive primates (i.e., omomyids, microchoerids, tarsiers,
adapiforms, and extant tooth-comb primates). This tubercle is
dorsoplantarly tall and mediolaterally narrow, being attened on
both its medial and lateral sides. This tubercle is not reduced as in
tarsiers and is morphologically most similar to that of omomyids.
The proximal end of this tubercle is mediolaterally narrowas it is in
other omomyids in contrast to the wider construction found among
Table 5
Talar ratios in omomyids and microchoerids.
Talar head
width/height
Talar neck length/
talar length
Mid-trochlear width/
trochlear length
Lateral body height/
mid-trochlear width
Talar width/talar
length
Teilhardina belgica n ¼2e3 148.8 47.8 (47.5e48.1) 88.1 (87.3e89.0) 119.7 (115.7e123.7) 60.3 (59.3e61.2)
Shoshonius cooperi n ¼3 115.4 (101.6e137.5) 52.4 (49.6e55.1) 76.1 (72.3e79.2) 114.8 (113.4e116.2) 50.7 (50.2e51.2)
Omomys sp. n¼7 119.6 (112.0e132.0) 53.3 (50.5e58.7) 82.0 (73.4e91.1) 110.0 (92.7e119.4) 58.9 (55.8e62.7)
Hemiacodon gracilis n ¼22 122.0 (111.7e132.9) 51.2 (48.1e56.7) 76.2 (67.6e81.2) 109.0 (95.7e118.4) 56.4 (52.7e59.6)
Tetonius homunculus n ¼3 54.9 (53.9e56.7) 74.7 (66.7e78.7) 116.5 (115e118) 57.2 (54.7e59.6)
Washakius insignis n ¼1 110.2 52.4 66.1 117.7 52.6
Necrolemur zitteli n ¼1 112.2 45.0 82.2 110.1 52.2
Nannopithex abderhaldeni n ¼1 141.1 53.3 66.0 119.1 59.0
The mean and range are listed.
Figure 4. Six views of IRSNB M1235, talus of T. belgica (left to right: dorsal and plantar views; middle, medial view (top), lateral view (below); right, distal view (top), posterior or
proximal view (below)).
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218 209
microchoerids and adapiforms. The A/B tubercle width index in
Table 10 shows T. belgica to be similar to Shoshonius,Hemiacodon,
and Omomys, all possessing relatively narrower tubercles
compared with Necrolemur and the adapiform Cantius. The C/D
index compares peroneal tubercle length relative to the length of
the proximal end of the rst metatarsal (Table 10). Here T. belgica
compares best with Necrolemur,Cantius, and the IVPP specimen,
compared with the higher values recorded for the other omomyids
and Microcebus. A lower ratio value indicates a relatively shorter
tubercle and is likely the primitive character state given its
commonality among the primitive adapiform Cantius,Teilhardina,
and Necrolemur. The H/G index (Table 10) compares height and
length values of the peroneal tubercle from the side. T. belgica is
most similar to Omomys and Cantius in this ratio whereas other taxa
possess higher values. Omomys,Cantius, and Teilhardina all show
a relatively lower tubercle height.
Distal femur
The femur (IRSNB M1263) is only represented by the distal
epiphysis (Fig. 8). It is broken at the anterior aspect of the medial
patellar rim region with severe erosion along the medial epi-
condyle. This specimen is also eroded with pitting along the lateral
epicondyle and condyle and to a lesser extent along the medial
condyle. The dimensions of IRSNB M1263 (Table 11) are between
those of the pygmy mouse lemur (M. berthae) and Shoshonius.
T. belgica exhibits an anteroposteriorly tall distal femur with a high
lateral patellar rim, a condition commonly found among leaping
Table 6
Calcaneal measurements by specimen (mm).
Length Width PACL PACW CUHT CUW HeelL
IRSNB Vert 16786-03 6.98 2.43 1.68 1.22 1.73 1.57 1.77
IRSNB M1236 7.39 2.69 2.04 1.28 1.69 1.70 1.44
IRSNB M1247 6.83 2.39 1.87 1.16 1.71 1.71 1.68
IRSNB Vert 16786-04 2.70 1.69 1.39 1.82
IRSNB Vert 26857-05 7.21 2.51 2.22 1.33 1.78 1.82 1.72
IRSNB M1237 6.52 2.41 1.79 1.07 1.61 1.66 1.67
IRSNB M61 6.62 2.39 1.61 1.55
IRSNB Vert 16786-06 1.96 1.77 1.18 1.88
IRSNB Vert 26857-03 2.42 1.69
IRSNB 4387
IRNSB 4387 is a worn specimen allowing no measurements. PACL and PACW ¼the
length or width of the posterior astragalocalcaneal facet; CUHT or CUW ¼the height
or width of the calcaneocuboid facet; HeelL ¼the length of the calcaneus or heel
region proximal to the posterior astragalocalcaneal facet.
Figure 5. Six views of IRSNB M1236, calcaneus of T. belgica (left column: distal (top) and proximal (below) views; left to right: dorsal, lateral, medial, and plantar views).
Figure 6. Size variation among four calcanei of T. belgica (dorsal and plantar views; left to right: A, IRSNB M1237; B, IRSNB M61; C, IRSNB M1247; and D, IRSNB M1236).
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218210
prosimian primates (Napier and Walker, 1967;McArdle, 1981;
Anemone, 1990;Dagosto, 1993). A ratio of lateral condylar height
relative to bicondylar width (LCH/BW, Table 11) shows T. belgica to
be very similar to Shoshonius,Hemiacodon,Omomys, the Shang-
huang haplorhine IVPP V13017, and to Ourayia (Dunn, 2010). The
patellar groove is narrow like that of other omomyids (PG/BW,
Table 11) in contrast to the wider grooves observed in tarsiers and
microchoerids. The popliteal groove is observable along the lateral
epicondylar region and the intercondylar notch is high. The inter-
condylar gap width relative to bicondylar width ratio (IG/BW,
Table 11)ofT. belgica is closest to that of Shoshonius and this value
lies in the middle relative to other omomyid IG/BW ratio values.
Proximal phalanges
Two proximal phalanges and two middle phalanges were
recovered from the miscellaneous bone boxes from Dormaal. Their
linear measurements are listed in Table 12. The smaller of the
proximal phalanges (IRSNB M1264, Fig. 9) is tentatively allocated to
the pollex on the basis of size and its proximal basal anatomy. The
larger specimen, IRSNB M1265 (Fig. 10), is most likely a proximal
phalanx for the hallux given its larger overall size and the prox-
imodorsal fossa common to pedal proximal phalanges. This allo-
cation also makes sense given their respective length to basal width
ratios (AB/KL, Tables 13 and 14). In Tarsius and the two galago
species, proximal pollical phalanges are more elongated than their
hallucal counterparts (Tables 13 and 14), and this pattern is found in
the Teilhardina specimens as well. The AB/KL ratio in Table 13 shows
Teilhardina to possess an elongated pollical phalanx, above the
range of the two galago species and within the range of T. syrichta,
a long ngered primate. Overall, the ratio values for both the pol-
lical and hallucal specimens are similar to the values calculated for
Tarsius,Galagoides, and Galago (Tables 13 and 14).
Both of the proximal phalanges are slightly curved ante-
roposteriorly and wide at the proximal and distal articular regions.
Their shafts are similar in width along their whole length (Table 12).
The proximal phalanges possess a single articular surface for the
metacarpal or metatarsal heads proximally and a wide distal facet
for the middle phalanx. The proximal joint surface is relatively at
in both. They contrast in proximal joint height, being taller in IRSNB
M1264, and quite at in IRSNB M1265. The IRSNB M1264 specimen
also has a distinct plantar tubercle that is absent on the IRSNB
M1265 proximal phalanx. Both appear to be fairly robust bones
implying strong digital musculature associated with grasping.
The proximal phalangeal distal heads are broad and cylindrical
along the distal edge. They contrast in that the IRSNB M1264
specimen is slightly indented or notched along this same region.
The collateral ligament impressions are more prevalent in the distal
head of the IRSNB M1264 specimen as well.
The plantar surfaces of both proximal phalanges are quite
grooved centrally with elevated medial and lateral rims for prom-
inent digital exor tendons that lie between. The elevated rims
provide prominent attachment sites for the exor retinacular
sheaths to keep these tendons within their brous tunnels. Flexor
Table 7
Measures of variance in calcaneal length and width in Teilhardina and other small
primates.
Taxon NCalcaneal length Calcaneal width
CV I
max/min
R%CVI
max/min
R%
Teilhardina belgica
Dormaal 6/9 4.6 113.3 12.6 8.5 137.8 30.3
Single
species
Tarsius syrichta
Different islands 12 3.4 112.9 12.1 6.3 118.6 16.9
Mindanao only 3 0.9 101.6 1.6 8.2 114.5 13.8
Tarsius bancanus 8 4.9 116.8 15.3 8.9 131.4 26.2
Microcebus griseorufus
Amboasary 14 3.9 112.5 11.8 6.1 121.7 19.6
Omomys
Covert quarry 5/3 6.6 119.1 17.5 1.7 103.1 3.0
Mixed
species
Tarsius 20 5.0 121.7 19.8 8.3 131.4 27.7
Microcebus 18 7.1 131.7 26.4 6.4 127.3 23.8
Bridger B omomyids 21/48 9.4 173.8 49.2 8.5 142.2 36.3
Imax/min is the percentage of the maximum to minimum observed value; R% is the
range expressed as a percentage of the mean. In samples with N<10, the CV was
calculated using the correction for small samples (Sokal and Rohlf, 1995).
Table 8
Calcaneal comparative data.
Length Width PACL PACW CUHT CUW HeelL Distal
length
Teilhardina belgica n ¼4 6.93 2.43 1.82 1.23 1.70 1.69 1.69 3.69
Absarokius abbotti n ¼1 9.80 2.87 1.95 1.16 1.54 2.28 5.53
Shoshonius cooperi n ¼2 9.79 3.25 2.23 1.25 1.55 2.32 2.22 5.20
Nannopithex
abderhaldeni n ¼1
3.27 2.06 1.31 2.18
Tetonius homunculus
n¼2
3.34 2.61 1.27 2.10
Washakius insignis n ¼2 10.50 3.35 2.45 1.42 1.90 2.12 2.51 5.57
Necrolemur zitteli n ¼3 3.52 3.17 1.97 3.20
Hemiacodon gracilis
n¼12
16.06 5.06 3.14 2.17 2.69 3.64 4.43 8.53
Omomys carteri n ¼6 15.90 5.14 3.21 2.17 3.15 3.74 4.09 8.44
Taxa are arranged by mean calcaneal width (smallest to largest).
Table 9
Mean and range for calcaneal ratios.
Distal length/Cal L CalW/CalL Heel L/CalL PacL/CalL PacW/PacL CuW/CuHt
Teilhardina belgica 53.2 35.7 23.7 27.0 66.6 99.3
n¼3e5 52.0e55.1 34.8e36.9 19.5e25.4 24.1e32.6 59.8e82.2 90.8e103.1
Shoshonius cooperi 53.1 33.2 22.6 22.7 56.1 149.9
n¼2 52.9e53.4 30.2e36.2 21.9e23.4 22.3e23.1 51.8e60.4 146.8e152.9
Omomys sp. 53.0 32.0 25.7 20.3 68.0 118.8
n¼6 51.2e56.0 30.0e33.2 22.7e28.4 17.4e24.0 59.0e73.4 112.6e127.9
Hemiacodon gracilis 52.3 31.6 28.7 20.2 68.3 134.2
n¼12 49.5e54.3 30.4e32.1 26.8e30.2 18.3e21.7 61.9e77.7 128.1e149.6
Tetonius homunculus
n¼2
x x x x 46.4 x
Absarokius abbotti
n¼1
56.4 29.3 23.3 19.9 59.5 x
Washakius insignis
n¼2
53.0 32.3 23.7 25.6 58.1 111.6
52.4e53.6
Necrolemur zitteli
n¼3
x x x x 62.6 x
54.3e69.0
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218 211
sheath grooving is strongest in IRSNB M1265, the likely hallucal
specimen, and this specimen also has a wider proximal base.
In terms of linear measurements (Table 12) and ratios (Tables 13
and 14), the proximal phalanges of Teilhardina are similar to each
other and this pattern generally holds for the few living primates
we have measured. They differ in that the IRSNB M1265 specimen
has an absolutely and relatively wider head and base (KL/IJ and KL/
UV ratios in Table 14). The IRSNB M1264 specimen is relatively
longer and taller as shown by its higher values for the UV/ST (base
height to mid-shaft height) and the AB/KL (phalangeal length to
base width) ratios (Table 13).
Compared with other small primates (T. syrichta,Galago sene-
galensis, and Galagoides demidovii;Table 13), the proximal pollical
phalanx of Teilhardina has a higher UV/ST ratio (base height to mid-
shaft height), suggesting a relatively taller basal shaft. In contrast,
the KL/IJ (base width to lower shaft width) and KL/UV (base width
to base shaft height) ratios are low, suggesting that this phalanx of
Teilhardina had a narrower basal width compared with those of
living taxa (Table 13). The other ratios are similar.
Likewise, the proximal hallucalphalanx of Teilhardina is very
similar in shape to that of T. syrichta,G. demidovii and
G. senegalensis. Only two ratios, the UV/ST (base height to mid-shaft
height) and the KL/IJ (base width to lower shaft width) ratios are
signicantly lower than in the living primates (Table 14). As in the
pollical comparisons, base width and height appear to be more
distinctive in Teilhardina.
Overall, both of the rst digit proximal phalanges of Teilhardina
are robust, suggesting large muscle attachments and forceful
grasping. The pollicalspecimen is long, as in Tarsius, suggesting
elongated manual digits.
Middle phalanges
The middle phalanges are quite elongated and gracile relative to
the more thick-shafted proximal phalanges (Table 15;Fig. 11),
suggesting that they are from digits 2e5 and not 1. The most
complete specimen (IRNSB M1266) is longer than either of the two
proximal phalanges (Table 12), suggesting that it derives from the
third or fourth digit (Table 14). We have been unable to identify
characters or sets of characters that consistently sort manual from
pedal middle phalanges in living primates so we cannot denitively
allocate these two specimens to the hand or foot on the basis of
shape. The one exception to this pattern is the AB/KL length to base
ratio in T. syrichta, which distinguishes the longer and thinner
phalanges of the tarsier hand from those of the foot (Table 15). This
index is also high in Teilhardina, suggesting an allocation to the
hand. IRNSB M1266 is absolutely much longer (5.56 mm) than the
middle phalanx of the presumably similarly sized M. berthae
(4.10 mm in length for a third digit manual and 4.15 mm for a third
digit pedal). It also exceeds the length of G. demidovii manual (4.97)
or pedal third digits (5.03) and falls within the size range of Tarsius
syrichtaspedal middle phalanx (digit three mean ¼5.97 mm). If
this phalanx is indeed from the third digit of the hand as suggested
by the AB/KL index, Teilhardina may have had manual digits as
elongated as the very specialized ngers of Tarsius (Figs. 12 and 13
A). If this phalanx is from the third or forth digit of the foot, it is still
Figure 7. IRSNB M1262, rst metatarsal of T. belgica (from left to right: proximal, dorsal and plantar views (top); medial and lateral views (below)).
Table 10
First metatarsal length measurements (mm) and ratios.
Teilhardina belgica Shoshonius cooperi Hemiacodon gracilis Omomys carteri Necrolemur zitteli IVPP V 13018 Microcebus berthae Cantius ralstoni
A 0.70 0.86 1.18 1.19 1.60 0.65 0.71 2.10
B 2.15 2.59 4.16 3.61 3.60 1.75 1.73 5.65
C 1.25 2.03 4.26 3.15 2.50 0.90 1.20 2.84
D 2.40 3.30 6.28 5.28 4.55 1.80 1.87 5.27
E 14.40 12.11 13.00 6.15 5.27
F 16.25 14.64 14.80 6.90 6.32
G 1.70 2.36 4.59 3.26 2.72 0.90 0.94 2.90
H 1.50 2.23 4.50 2.89 2.60 0.85 1.09 2.52
A/B 0.33 0.33 0.28 0.33 0.44 0.37 0.41 0.37
C/D 0.52 0.62 0.68 0.60 0.54 0.50 0.64 0.54
E/F 0.89 0.83 0.88 0.89 0.83
H/G 0.88 0.95 0.98 0.89 0.96 0.94 1.16 0.87
D-C/B 0.53 0.49 0.49 0.59 0.57 0.51 0.39 0.43
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218212
elongated compared with extant primates, though not nearly as
strongly (Fig. 13B and C). Other phalangeal ratios of T. belgica are
similar to those of Tarsius or Galago and do not appear overtly
unusual.
The distal joint surface of the two middle phalanges of T. belgica
is very cylindrical in shape and is very at across the most distal
edge. On the dorsal side, there is a round fossa for articulation with
the distal phalanx during extreme extension, a feature that can be
found on middle phalanges of living primates as well. This fossa is
not present on the proximal phalangeal specimensof T. belgica,asis
the case in living primates as well. The shafts of the Teilhardina
specimens have greater anteroposterior curvature than the prox-
imal phalanges, another similarity to living primates. The exor
sheath ridges and grooves run along the proximal two-thirds of the
middle phalangeal shaft. The distal third of the shaft is quite
elongated relative to other extant primates. The proximal joint
surface of these two middle phalanges possesses two facets sepa-
rated by a central ridge allowing contact with their corresponding
proximal phalangeal distal joint surface. The plantar base is wider
than that of the dorsal region (Fig. 11). These features are similar to
other primates.
Discussion
Function
The new postcranial elements of T. belgica are consistent with
the allocation of the previously known tarsal elements, the talus
and calcaneus, to this species by Teilhard de Chardin (1927) and
Szalay (1976). As has been inferred before (Szalay, 1976;Gebo,1988;
Dagosto, 1993), the talus and calcaneus of T. belgica have features
(i.e., an elongated distal calcaneus, a long talar neck, and a tall talar
trochlea) indicative of leaping. The tall knee with a high lateral
patellar rim is consistent with this conclusion. This suite of bony
hindlimb features is characteristic of other omomyids as well
(Dagosto et al., 1999). None of these features areso specialized as to
indicate vertical clinging or leaping as specialized as in Tarsius or
small galagos (i.e., the vertical clingers and leapers; see Stern and
Oxnard, 1973;Anemone, 1990;Dagosto et al., 1999). Rather they
indicate a more generalized leaper-quadruped behavioral pattern
typical of most extant strepsirhine primates (Dagosto et al., 1999).
We can also infer from the posterior calcaneal facet, the calcaneo-
cuboid joint surface, the talar head, the proximal rst metatarsal
joint, the long peroneal tubercle on the rst metatarsal, the robust
hallucal phalanx, and perhaps the long middle phalanges that
T. belgica had excellent foot mobility and grasping capabilities.
There is no doubt that this fossil species was a capable climbing
primate. Perhaps the most unusual aspect of the body of T. belgica is
its digit length, possibly being elongated to a degree seen only in
the manual phalanges of tarsiers. Unfortunately, evidence for digit
proportions in other omomyids is lacking and this prevents us from
determining whether elongated digits (whether manual or pedal)
were typical of all omomyids or all tarsiiforms or whether this is an
autapomorphic condition uniquely found in Teilhardina.
The recent article by Rose et al. (2011) documents additional
elements for the genus Teilhardina including a navicular and distal
phalanges. The navicular is elongated and thus consistent func-
tionally with the elongated distal calcaneus of Teilhardina. The
attened distal phalanges conrm the presence of nails, a hallmark
Figure 8. Three views of the IRSNB M1263distal femur of T. belgica (left to right views:
medial, distal and lateral).
Table 11
Distal femoral measurements (mm) and ratios.
Microcebus
berthae
Teilhardina
belgica (IRSNB
M1263)
Shoshonius
cooperi (CM
69764)
Hemiacodon
gracilis (AMNH
12613)
IVPP V13017 Omomys
carteri (n¼3)
Distal bicondylar width (BW) 3.79 4.20 5.05 8.10 8.35 8.45
Lateral condyle distal height (LCH) 3.91 4.75 5.63 10.15 9.54 9.27
Medial condyle distal height (MCH) 5.02 8.85 9.20 8.63
Patellar groove width (PG) 1.13 1.25 1.58 1.89 2.44 2.26
Medial condyle width (MCW) 1.27 1.25 1.71 2.70 2.20 2.59
Lateral condyle width (LCW) 1.42 1.50 3.16 2.73 2.71
Intercondyle gap width (IG) 1.30 1.43 1.96 3.06 3.07
LCH/BW 1.03 1.13 1.12 1.16 1.14 1.10
MCH/BW 0.99 1.09 1.10 1.02
PG/BW 0.30 0.30 0.31 0.24 0.29 0.27
MCW/LCW 0.89 0.83 0.85 0.81 0.96
IG/BW 0.31 0.28 0.24 0.37 0.36
Table 12
Measurements (mm) for individual phalanges of T. belgica.
Linear
measurements
Proximal
pollical
phalanx IRSNB
M1264
Proximal
hallucal
phalanx IRSNB
M1265
Middle
phalanx
IRSNB Vert
26857-04
Middle
phalanx
IRSNB
M1266
AB (L) 4.40 4.45 5.60
CD (DHW) 0.75 1.00 0.50 0.50
EF (PHW) 1.00 1.40 0.75 0.80
GH (MSW) 0.75 0.80 0.40 0.70
IJ (PSW) 1.10 1.15 0.70 0.85
KL (BW) 1.25 1.50 1.15
MN (HL) 0.75 0.75 0.55 0.60
OP (HH) 0.70 0.70 0.40 0.55
QR (DSH) 0.40 0.50 0.25 0.40
ST (MSH) 0.60 0.70 0.35 0.50
UV (BH) 1.25 1.15 0.85
() ¼approximate measurement equivalents from Hamrick et al. (1995).
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218 213
adaptation of all primates, in the oldest known primate from North
America.
Tarsiers are distinctive among living primates in having
enlarged nger pads, long ngers and hands, and an arched nger
posture (Clark, 1936,1959;Hill, 1955;Day and Iliffe, 1975;Jouffroy
and Lessertisseur, 1979;Niemitz, 1984a,b;Jouffroy et al., 1993;
Anemone and Nachman, 2003;Lemelin and Jungers, 2007). Tarsiers
use their long ngers to pin down prey items on the forest oor
before they grasp them (DLG and MD, Personal observation; see
also Niemitz, 1984b;Crompton and Andau, 1986). If the long middle
phalanges of Teilhardina imply equivalently long ngers, they may
have played a similar functional role in prey capture.
The highly expanded disk-like pads of tarsier digits are associ-
ated with reduced triangular-shaped nails, and phalangeal heads
that are round in contrast to the more arrow-head shape typical of
other primates, including other omomyids (see Dagosto, 1988).
Galagos, while not having the enlarged digital pads of tarsiers, also
exhibit rounded phalangeal digital tufts, although they are much
smaller than those of tarsiers. The distal phalanges of Teilhardina
described and illustrated in Rose et al. (2011) have expanded
circular apical tufts similar to those of extant tarsiers, suggesting
that perhaps Teilhardina also had highly expanded disk-like digital
pads with reduced triangular-shaped nails. Since this particular
feature of tarsiers has been suggested to be related to their frequent
use of smooth vertical supports (Clark, 1959;Day and Iliffe, 1975;
Niemitz, 1984a;Anemone and Nachman, 2003), the use of this type
of substrate may also have been part of the Teilhardina niche. We
note, however, that there are also several differences between the
Figure 9. Five views of the IRSNB M1264 proximal digit 1 phalanx of T. belgica (left
column: proximal joint surface (top) with an angled view below; dorsal, plantar, and
medial views to the right).
Figure 10. Five views of the IRSNB M1265 proximal digit 1 phalanx of T. belgica (left
column: proximal joint surface (top) with an angled view below; dorsal, plantar, and
medial views to the right).
Table 13
Proximal pollical phalangeal mean ratio and range comparisons.
Teilhardina
belgica
Tarsius syrichta Galagoides demidovii Galago
senegalensis
Proximal
pollical
phalanx
N¼1N¼9N¼4N¼9
EF/CD 1.33 1.53 1.43 1.46
1.39e1.80 1.28e1.56 1.25e1.61
OP/AB 0.16 0.17 0.20 0.21
0.14e0.20 0.17e0.22 0.18e0.27
GH/ST 1.25 1.18 0.99 0.95
1.00e1.25 0.89e1.23 0.83e1.10
QR/ST 0.67 0.78 0.76 0.72
0.59e1.00 0.60e0.88 0.65e0.78
UV/ST 2.08 1.50 1.61 1.49
1.39e1.75 1.47e1.92 1.39e1.67
KL/IJ 1.14 1.42 1.91 1.72
1.14e1.85 1.75e2.00 1.54e1.90
GH/AB 0.17 0.18 0.17 0.18
0.13e0.22 0.17e0.18 0.16e0.22
ST/AB 0.14 0.15 0.18 0.20
0.13e0.18 0.14e0.20 0.17e0.22
KL/UV 1.00 1.36 1.31 1.29
1.07e1.60 1.25e1.36 1.19e1.43
OP/MN 0.93 0.95 1.12 1.13
0.83e1.00 1.06e1.18 0.90e1.50
AB/KL 3.52 3.23 2.68 2.69
2.16e4.18 2.62e2.79 2.50e2.88
Table 14
Proximal hallucal phalangeal mean ratio and range comparisons.
Teilhardina
belgica
Tarsius
syrichta
Galagoides
demidovii
Galago
senegalensis
Proximal
hallucal
phalanx
N¼1N¼12 N¼4N¼8
EF/CD 1.40 1.47 1.37 1.48
1.21e1.68 1.25e1.45 1.29e1.76
OP/AB 0.16 0.19 0.22 0.22
0.16e0.22 0.19e0.24 0.19e0.26
GH/ST 1.14 1.12 1.03 1.11
0.96e1.26 0.83e1.10 1.00e1.27
QR/ST 0.71 0.72 0.68 0.69
0.62e0.86 0.58e0.85 0.54e0.80
UV/ST 1.44 1.56 1.62 1.57
1.41e1.70 1.46e1.73 1.44e1.76
KL/IJ 1.30 1.54 1.74 1.61
1.40e1.67 1.60e1.88 1.37e1.71
GH/AB 0.18 0.18 0.20 0.22
0.16e0.22 0.20e0.21 0.19e0.27
ST/AB 0.16 0.16 0.20 0.20
0.15e0.20 0.18e0.24 0.17e0.22
KL/UV 1.30 1.38 1.38 1.50
1.26e1.67 1.29e1.50 1.37e1.67
OP/MN 0.93 0.93 1.07 1.03
0.75e1.07 1.00e1.14 0.91e1.15
AB/KL 2.97 2.87 2.26 2.18
2.20e3.08 2.19e2.32 1.75e2.81
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218214
phalanges of Teilhardina and Tarsius. For example, in Teilhardina the
basal articular surface is mediolaterally broader and deeper with
better developed medial and lateral basal tubercles and the
phalangeal shaft is relatively longer.
Phylogeny
Teilhardina is one of the most primitive of all known fossil
primates from the early Eocene as demonstrated by several detailed
cladistic analyses (Ni et al., 2004;Smith et al., 2006;Beard, 2008;
Rose et al., 2011). Based primarily on dental characters, the most
recent study (Rose et al., 2011) places T. asiatica as the most basal
species. T. belgica is the sister taxon of Teilhardina magnoliana in one
analysis and of T. asiatica in a second; while in the strict consensus
T. asiatica,T. belgica, and T. magnoliana, form a polytomy with
a clade of remaining omomyids including T. brandti (Rose et al.,
2011). All of these analyses show Teilhardina,Steinius, and Teto-
nius to be basal omomyids (Rose et al., 2011). Neither the previously
known nor newly described postcranial remains of T. belgica and
T. brandti conict with these interpretations nor with several
analyses that indicate a more basal phyletic position for T. belgica or
for Teilhardina in general (see Ni et al., 2004,2010;Tornow, 2008;
and Rosenberger, 2011). The postcranium of Teilhardina appears to
be quite similar to that of other taxa within the Omomyidae. No
elements of its postcranium exhibit special resemblance to any of
the taxa suggested to be the sister group of Primates (e.g., plesia-
dapiforms, scandentians, or dermopterans; see Clark, 1959;Kay
et al., 1990;Beard, 1993;Silcox, 2007). Rather, the new post-
cranial elements described here and in Rose et al. (2011) indicate
that Teilhardina already possessed all of the hallmarkadaptations of
early Eocene primates.
Table 16 compares postcranial features in T. belgica currently
accepted on the basis of dental and postcranial features as an early
representative of the haplorhine clade (e.g., Szalay and Delson,
1979;Gunnell and Rose, 2002;Rose et al., 2011) and Cantius,an
early representative of the strepsirhine (adapiform) radiation of
primates. Cantius and Teilhardina share six of the 18 features
including full medial talobular facets (Gebo, 1986), fan-shaped
calcaneocuboid joints (Gebo, 1988), tall knees with elevated
lateral patellar rims, and robust rst metatarsals with long peroneal
tubercles. That the two earliest known members of the major
clades of primates possess these features is strong evidence that
they are primitive for the Order.
Cantius and Teilhardina differ in the degree of elongation of the
talar neck, the distal calcaneus, and phalanges. These and other
Table 15
Mean ratio values and ranges for middle phalanges (3 ¼digit three).
Teilhardina
belgica
Tarsius syrichta
3-manual
T. syrichta
3-pedal
Galago
senegalensis
3-manual
G. senegalensis
3-pedal
Middle
phalanx
N¼1e2N¼9N¼11 N¼9N¼8
EF/CD 1.55 1.65 1.47 1.37 1.40
1.25e2.00 1.17e1.87 1.14e1.76 1.15e1.88
OP/AB 0.10 0.13 0.19 0.14 0.13
0.11e0.15 0.11e0.33 0.09e0.19 0.12e0.16
GH/ST 1.27 1.08 0.98 1.27 1.15
0.83e1.43 0.77e1.11 1.0e1.53 0.80e1.5
QR/ST 0.76 0.79 0.83 0.71 0.67
0.56e1.13 0.59e1.13 0.53e0.90 0.60e0.80
UV/ST 1.70 1.53 1.53 1.76 1.69
1.11e2.00 1.18e2.00 1.41e2.38 1.40e1.87
KL/IJ 1.35 1.47 1.54 1.48 1.47
1.11e1.80 1.27e1.75 1.25e1.76 1.29e1.60
GH/AB 0.13 0.13 0.15 0.18 0.15
0.08e0.19 0.11e0.18 0.12e0.23 0.08e0.20
ST/AB 0.09 0.13 0.15 0.14 0.13
0.10e0.22 0.12e0.19 0.10e0.17 0.10e0.16
KL/UV 1.35 1.32 1.28 1.19 1.24
1.03e1.9 1.15e1.50 0.88e1.58 1.03e1.50
OP/MN 0.83 1.11 1.12 1.03 0.99
0.93e1.33 1.00e1.33 0.83e1.25 0.83e1.13
AB/KL 4.83 4.29 3.57 3.59 3.62
3.26e5.14 2.72e4.39 3.03e4.25 3.22e4.08
Figure 11. Five views of the IRSNB M1266 middle phalanx of T. belgica (left column:
proximal joint surface (top) with an angled view below; dorsal, plantar, and medial
views to the right).
Figure 12. A comparative plantar view of digit 3 manual middle phalanges for Galago
senegalensis (left), Tarsius syrichta (middle), and the IRSNB M1266 phalanx of T. belgica
(right). Base widths are approximately equal. Scale bar ¼1 mm.
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218 215
differences such as the relative size of the posterior trochlear shelf
or a wide or narrow calcaneal shape may be associated with size
differences between Teilhardina and Cantius or they may indicate
subtle differences in positional behavior.
Another set of features (e.g., the slope of the talobular facet or the
position of the exor hallucis longus groove) are indicative of the basic
division of primates into the strepsirhine and haplorhine clades(Beard
et al., 1988). Likewise, the position of the calcaneal peroneal tubercle
and the width of the patellar groove differ between these two genera
as wellas across otherprimates.The arc of jointcurvaturebetween the
rst metatarsal and the entocuneiform, however, may be phyletically
informative in diagnosing tarsiiform primates (omomyids, micro-
choerids, and tarsiids) given that adapiforms and crown group
anthropoidspossess widerjoint arcs.This differencein arc of curvature
maybeanimportantindicatorofgraspingabilitiesofearlyprimates
and presumably one of these character states should represent the
primitive primate condition. If the narrow arc condition is primitive for
primates, then both adapiforms and crown-group anthropoids inde-
pendently evolved the wide condition. If the wide condition is
primitive for primates,crown-groupanthropoids reverted to the wide
condition since the narrow arc condition is found among primitive
haplorhine primates (i.e., omomyids, microchoerids and tarsiids).
Conclusion
As one of the few early fossil primatesprimitive enough to possess
four premolars, T. belgica represents a signicant window into
primate origins.Our discoveries of a few new postcranialelements of
T. belgica and those of Rose et al. (2011) for the closely related
T. brandtienhance our knowledge of its morphologyand adaptations.
An estimated size of 30e60 g places T. belgica among the smallest of
extant and extinct primates (Gebo, 2004). Although no partial skel-
eton exists for Teilhardina, which would allowthe evaluation of more
sensitive indicators of locomotor behavior like limb indices, the
anatomy of the ankle, knee, and foot conrm that its leaping,
grasping, and climbingabilities were similar to those of other Eocene
and extant primates. The known postcranial elements of Teilhardina
are very similar tothose of other omomyids, such as the better known
Shoshonius or Hemiacodon (Simpson, 1940;Szalay, 1976;Dagosto
et al., 1999). On the basis of known postcranial anatomy, there is no
reason to exclude Teilhardina from the Omomyidae. Comparisons of
Teilhardina and Cantius postcranials illustrate that haplorhine/
strepsirhine distinctions in the tarsals were already established by
the earliest Eocene. Like Cantius,Teilhardina exhibits all of the hall-
mark postcranial features (e.g., nailed digits, opposable hallux, tall
knee, elongated tarsals) that distinguish primates from any sug-
gested sister taxa. Neither Teilhardina nor Cantius bridge the
morphological gap that exists between Eocene primates and
proposed outgroup clades such as the plesiadapiforms, scandentians,
or dermopterans, and therefore cannot resolve the current argument
as to which of these is the closest relative of Primates.
Acknowledgments
We thank the staff at the Field Museum of Natural History
(Chicago) and Dr. Annelise Folie at the Royal Belgian Institute of
Natural Sciences (Brussels, Belgium) for their help with the
collections housed therein. We add a special thank you to Eric De
Bast at the Royal Belgian Institute of Natural Sciences for realizing
the SEM pictures and Dr. Steve Goodman at the Field Museum of
Natural History for allowing access to his Microcebus collection.
This paper is a contribution to project MO/36/020, which is nan-
cially supported by the Federal Science Policy Ofce of Belgium.
Figure 13. Regressions of middle phalanx length to body mass in primates; both values natural log transformed. Circles represent anthropoids, squares represent lemuriforms, and
triangles represent tarsiers. In each graph, the solid line is the reduced major axis regression and the dotted line is the ordinary least squares regression. The position of T. belgica
(values not included in the regressions) at both 30 and 50 g is marked by the star symbol. A, the middle phalanx of the third digit of the hand (our data supplemented with values
from data from Kirk et al., 2008;r¼0.84); B, the middle phalanx of the third digit of the foot (r¼0.93); and C, the middle phalanx of the fourth digit of the foot (r¼0.91; B and C
supplemented with data from Volkov, 1903e190 4). Note that in any case, Teilhardina has relatively long middle phalanges compared with extant primates.
Table 16
Comparison of postcranial features in two primitive early Eocene primates.
Teilhardina belgica Cantius sp.
Talus
1. Talar neck length Long Short
2. Medial talotibular facet Full Full
3. Talobular facet Steep-sided Oblique
4. Posterior trochlear shelf Small Large
5. Location of exor hallucis
longus groove
Middle Lateral
Calcaneus
6. Calcaneal shape Narrow Wide
7. Distal calcaneal length Long Short
8. Calcaneocuboid joint Fan-shaped Fan-shaped
9. Peroneal tubercle position Distal to PCF Equal to PCF
Distal femur
10. Height of distal femur Tall Tall
11. Lateral patellar rim Elevated Elevated
12. Patellar groove width Narrow Wide
First metatarsal
13. First metatarsal shape Robust Robust
14. Peroneal tubercle length Long Long
15. Peroneal tubercle shape Narrow Broad
16. First metatarsal-entocuneiform
joint arc
Narrow Broad
Phalanges
17. Proximal phalangeal length Longer Long in Notharctus
18. Middle phalangeal length Longer Long in Notharctus
D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218216
Appendix
Fossil sample.
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Taxon Catalog number Element
Omomys carteri UCM 69339 Proximal phalanx
Omomys carteri UCM 69063 First metatarsal
Omomys carteri DMNH 32,300 First metatarsal
Omomys carteri DMNH 1998-179 First metatarsal
Omomys carteri UCM 68710 Distal femur
Omomys carteri UCM 67917 Distal femur
Omomys carteri UCM 57457 Distal femur
Omomys carteri UCM 67679 Calcaneus
Omomys carteri UCM 69303 Calcaneus
Omomys carteri UCM 69065 Calcaneus
Omomys carteri UCM 69678 Calcaneus
Omomys carteri UCM 68745 Calcaneus
Omomys carteri UCM 69457 Calcaneus
Omomys carteri UCM 69062 Talus
Omomys carteri UCM 69061 Talus
Omomys carteri UCM 67872 Talus
Omomys carteri UCM 67873 Talus
Omomys carteri UCM 89338 Talus
Omomys carteri UCM 57458 Talus
Omomys carteri UCM69400 Talus
Shoshonius cooperi CM 69756 Talus
Shoshonius cooperi CM 67297 Talus
Shoshonius cooperi CM 67298 Talus
Shoshonius cooperi CM 67299 Calcaneus
Shoshonius cooperi CM 69765 Calcaneus
Shoshonius cooperi CM 69764 Femur
Shoshonius cooperi CM 69754 First metatarsal
Tetonius homunculus AMNH 88817 Talus
Tetonius homunculus AMNH 88818 Talus
Tetonius homunculus AMNH 88819 Talus
Tetonius homunculus AMNH 88820 Calcaneus
Tetonius homunculus AMNH 88821 Calcaneus
Tetonius homunculus AMNH 88823 Calcaneus
Washakius insignis AMNH 97288 Talus
Washakius insignis AMNH 88824 Calcaneus
Washakius insignis AMNH 88826 Calcaneus
Hemiacodon gracilis AMNH 12613 Hindlimb
Hemiacodon gracilis AMNH 126627 Talus
Hemiacodon gracilis AMNH 12613A Talus
Hemiacodon gracilis AMNH 29183 Talus
Hemiacodon gracilis AMNH 29184 Talus
Hemiacodon gracilis AMNH 29153 Talus
Hemiacodon gracilis AMNH 29152 Talus
Hemiacodon gracilis AMNH 29188 Talus
Hemiacodon gracilis YPM 30461 Talus
Hemiacodon gracilis YPM 24479 Talus
Hemiacodon gracilis YPM 24461 Talus
Hemiacodon gracilis YPM 24476 Talus
Hemiacodon gracilis YPM 24466 Talus
Hemiacodon gracilis YPM 30457 Talus
Hemiacodon gracilis YPM-PU 10509 Talus
Hemiacodon gracilis YPM 39981 Talus
Hemiacodon gracilis YPM 39982 Talus
Hemiacodon gracilis YPM 39983 Talus
Hemiacodon gracilis YPM 24464 Talus
Hemiacodon gracilis YPM 39988 Talus
Hemiacodon gracilis YPM 39992 Talus
Hemiacodon gracilis YPM 44510 Talus
Hemiacodon gracilis AMNH 12046 Calcaneus
Hemiacodon gracilis AMNH 126626 Calcaneus
Hemiacodon gracilis AMNH 126625 Calcaneus
Hemiacodon gracilis YPM 24475 Calcaneus
Hemiacodon gracilis YPM 24460 Calcaneus
Hemiacodon gracilis YPM 39984 Calcaneus
Hemiacodon gracilis YPM 30458 Calcaneus
Hemiacodon gracilis YPM 30459 Calcaneus
Hemiacodon gracilis UW 5052 Calcaneus
Hemiacodon gracilis CM 34804 Calcaneus
Hemiacodon gracilis CM 34805 Calcaneus
Absarokius abbotti UCM 62681 Talus
Absarokius abbotti UCM 58904 Calcaneus
Appendix (continued)
Taxon Catalog number Element
Necrolemur zitteli Zurich NR-8761 Calcaneus
Necrolemur zitteli Zurich NR-8762 Calcaneus
Necrolemur zitteli Zurich A/V 637 Calcaneus
Necrolemur zitteli BFI 876 Talus
Necrolemur zitteli unnumbered First metatarsal
Nannopithex abderhaldeni GMH-Leo I-4231 Hindlimb
Cantius ralstoni UM 75756 First metatarsal
Cantius ralstoni USGS 21759 Talus
Cantius ralstoni UM 21447 Calcaneus
Cantius trigonodus USGS 5900 Hindlimb
Notharctus tenebrosus AMNH 11474 Skeleton
Notharctus tenebrosus AMNH 11478 Skeleton
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D.L. Gebo et al. / Journal of Human Evolution 63 (2012) 205e218218
... This article focuses on a single lineage of Eocene primates from the Willwood Formation, the Tetonius-Pseudotetonius (T-P) lineage. Taxonomically, the T-P lineage represents a group of Eocene omomyiform primates that are basal members of the Anaptomorphinae within the fossil primate family Omomyidae, which also includes the Omomyinae (Szalay and Delson 1979;Gunnell and Rose 2002;Gebo et al. 2012). Omomyidae is one of three haplorrhine families included within Tarsiiformes, the other two families being Microchoeridae and Tarsiidae (Szalay and Delson 1979;Gunnell and Rose 2002;Gebo et al. 2012). ...
... Taxonomically, the T-P lineage represents a group of Eocene omomyiform primates that are basal members of the Anaptomorphinae within the fossil primate family Omomyidae, which also includes the Omomyinae (Szalay and Delson 1979;Gunnell and Rose 2002;Gebo et al. 2012). Omomyidae is one of three haplorrhine families included within Tarsiiformes, the other two families being Microchoeridae and Tarsiidae (Szalay and Delson 1979;Gunnell and Rose 2002;Gebo et al. 2012). The taxonomic framework used here follows that of Szalay and Delson (1979) and Gunnell and Rose (2002) except for Microchoeridae being raised to the level of family. ...
... Omomyiforms have been extensively studied since originally described by Trouessart (1879). However, most studies have focused on systematics and phylogenetic relationships of omomyiforms (Tornow 2008;Gunnell et al. 2008), on the evolutionary relationships between omomyiforms and other primate families (Bown and Rose 1987;Bloch et al. 2007;Silcox et al. 2008;Seiffert et al. 2009;Williams et al. 2010), or adaptations in cranial and postcranial morphology (Bown and Rose 1987;Dagosto and Schmid 1996;Gebo et al. 2012). ...
Article
Full-text available
The Tetonius-Pseudotetonius (T-P) transition is an often-cited example of phyletic gradualism, but rates of evolution and the roles of neutral and adaptive processes across this lineage remain unclear. Linking Tetonius and Pseudotetonius, two omomyid primates, are a series of stratigraphic and morphologic intermediates revealing trends suggestive of a functional and developmental reorganization of the dentition. Notable changes involved the P3, which reduced size and became indistinguishable from the canine and I2, and the P4, which became a robust tall-cusped tooth comparable to a molar. We test two hypotheses: (1) neutral evolution can explain the observed phenotypic differences in the lineage, and (2) P4 lost developmental association with P3 and became integrated with the molars. First, we calculate the rate of evolutionary differentiation, based on the ratio between inter- and intra-species variation in length and width of P3, P4, M1, and M2 teeth, between lineage segments and over the entire lineage. We test the second hypothesis by comparing bivariate correlations between teeth within individual lineage segments. As the lineage evolved, correlations between P3 and the molars diminished, whereas correlations between P4 and the molars increased. We found evidence of varying degrees of stabilizing selection in the lengths and widths of all cheek teeth and evidence of neutral evolution in the width of P4. These results support a trend towards P4 becoming integrated into the molar morphogenetic field, and demonstrates that morphological rates of evolution, and consequently the degree of selective pressures, vary through time and between teeth.
... Sargis and colleagues proposed that the powerful pedal grasping capacities would have evolved even earlier, in the ancestor of the plesiadapoid-euprimate clade. And recent analyses started to question these scenarios, suggesting that leaping specialization would have been more anterior, and that grasping specialization for small branches was delayed Boyer and Seiffert 2013;Yapuncich et al. 2017;Boyer et al. 2013b;Gebo et al. 2012Gebo et al. , 2015. ...
... The past decades have been contented with a large amount of morphological-based studies of extant and extinct primate lineages aiming at verifying these hypotheses. Recent studies started to question the small branch niche early environment and proposed that features reflecting the small branch niche, prehensile foot proportions and nails on the lateral digits may well have been acquired after crown primates began to radiate, e.g. in parallel in different euprimate lineages, and that leaping specializations and large vertical substrates use were more anterior (Boyer et al., 2013aBoyer & Seiffert, 2013;Cartmill, 1974a;Gebo et al., 2012Gebo et al., , 2015Ni et al., 2013;Yapuncich et al., 2017). ...
... I thus measured two other plesiadapiforms from northwestern Wyoming (US): the carpolestid Carpolestes simpsoni from the SC-62 limestone locality (body mass around 100g, (Bloch & Gingerich, 1998)), and the micromomyid Dryomomys szalayi from the SC-327 limestone locality, with an estimated body mass around 30g (Bloch et al., , 2016. I measured also two early Eocene Euprimates: Teilhardina belgica from the Dormaal locality (western Belgium) with estimated body mass of 30-60g (Gebo et al., 2012(Gebo et al., , 2015, and Archicebus achilles from the Jingzhou locality (Hubei Province, China) with estimated body mass of 20-30g (Ni et al., 2013). Finally, I measured two Euprimates from the Middle Eocene sediments of Messel (near Darmstadt, Germany): Darwinius masillae with estimated body mass of 660g (Franzen et al., 2009) and Europolemur kelleri (Franzen & Frey, 1993;Hamrick, 2001b;Koenigswald, 1979;Von Koenigswald et al., 2012) with estimated body mass of 1,5 Kg (Boyer et al., 2013b;Von Koenigswald et al., 2012). ...
Thesis
Primate origins are subject to important controversies. The initial radiation of first Primates and their precise relationships within Euarchontans (the clade including Primates, Scandentians, Dermopterans, and Plesiadapiformes) are still debated. Moreover, the functional and evolutionary interpretation of some of the morphological characters that define Primates is still uncertain. Among them are the acquisition of manual and pedal prehensile abilities, with a specialized grasping foot bearing an opposable hallux, and nails instead of claws on the distal phalanges. Thus, the ancestral morphotype of Primates is under active investigation, despite the consensus on the arboreality and small size of our early ancestors. This PhD dissertation aimed at revisiting some blurry aspects of primate origins focusing on hand and foot grasping mechanisms, through an interdisciplinary approach blending ethology, biomechanics, comparative morphology and phylogenetics. A reappraisal of the genus Plesiadapis (Plesiadapiformes) led to question a recent hypothesis on early Primates’ phylogeny. In addition, a quantitative analysis of manual and pedal postures relatively to substrate type used during locomotion, followed by a morphological study of hand and foot metapodials and phalanges were also conducted on series of primate and non-primate species. The results were analyzed in an integrative way to relate morphological features to functional attributes, along with assessing their phylogenetic importance. Among many results, this work allowed proposing alternative hypotheses regarding two key characters of primates, the primary function of nails: more linked to sensitivity than to a mechanical advantage; and the environmental scenario that may have driven the evolution of hallucal grasping capabilities: small vertical substrates instead of the fine branch niche. Moreover, in an effort to better understand biomechanical constraints at play during arboreal locomotion, a novel spatially-resolved force sensor was created, which has potential applications in various fields such as robotics.
... 1924), as well as to infer positional behaviors in fossil taxa (Boyer and Seiffert, 2013;Boyer, Seiffert, & Simons, 2010;Boyer, Yapuncich, Butler, Dunn, & Seiffert, 2015;Dagosto, 1983;Dunn et al., 2016;Gebo and Simons, 1987;Gebo, 1988;Gebo, Dagosto, Beard, & Ni, 2008;Gebo, Dagosto, Beard, Qi, & Wang, 2000;Gebo, Dagosto, & Rose, 1991;Gebo, Smith, & Dagosto, 2012;Marig o, Roig, Seiffert, Moy a-Sol a, Marivaux et al., 2010;Marivaux et al., 2011;Seiffert and Simons, 2001;Seiffert, Costeur, & Boyer, 2015;). ...
... Additionally, comparative studies of fossil primate hands (Boyer et al., 2013bGebo et al., 2015) suggest that the ancestral euprimate likely had exceptionally elongate fingers similar to modern tarsiers or Daubentonia, which would facilitate clinging to, and grasping of, proportionally larger supports, rather than small, terminal branches. Finally, Gebo et al. (2012) ...
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Objective: On the talus, the position and depth of the groove for the flexor hallucis longus tendon have been used to infer phylogenetic affinities and positional behaviors of fossil primates. This study quantifies aspects of the flexor hallucis longus groove (FHLG) to test if: (1) a lateral FHLG is a derived strepsirrhine feature, (2) a lateral FHLG reflects inverted and abducted foot postures, and (3) a deeper FHLG indicates a larger muscle. Methods: We used linear measurements of microCT-generated models from a sample of euarchontans (n?=?378 specimens, 125 species) to quantify FHLG position and depth. Data are analyzed with ANOVA, Ordinary and Phylogenetic Generalized Least Squares, and Bayesian Ancestral State Reconstruction (ASR). Results: Extant strepsirrhines, adapiforms, plesiadapiforms, dermopterans, and Ptilocercus exhibit lateral FHLGs. Extant anthropoids, subfossil lemurs, and Tupaia have medial FHLGs. FHLGs of omomyiforms and basal fossil anthropoids are intermediate between those of strepsirrhines and extant anthropoids. FHLG position has few correlations with pedal inversion features. Relative FHLG depth is not significantly correlated with body mass. ASRs support a directional model for FHLG position and a random walk model for FHLG depth. Conclusions: The prevalence of lateral FHLGs in many non-euprimates suggests a lateral FHLG is not a derived strepsirrhine feature. The lack of correlations with pedal inversion features suggests a lateral FHLG is not a sufficient indicator of strepsirrhine-like foot postures. Instead, a lateral FHLG may reduce the risk of tendon displacement in abducted foot postures on large diameter supports. A deep FHLG does not indicate a larger muscle, but likely reduces bowstringing during plantarflexion.
... In this regard, it represents a vital source of information about intraspecific dental variability in the oldest omomyiform genus (e.g., Gingerich, 1977). Isolated postcrania of this species described from Dormaal provide the principal source of information about Teilhardina positional behaviors and allow for refinements of body mass predictions typically based on dentition alone (Gebo et al., 2012(Gebo et al., , 2015Boyer et al., 2013Boyer et al., , 2015Yapuncich et al., 2017;Dagosto et al., 2018). Beard, 2008 Holotype CM 70435, isolated R M 2 . ...
Article
Omomyiform primates are among the most basal fossil haplorhines, with the oldest classified in the genus Teilhardina and known contemporaneously from Asia, Europe, and North America during the Paleocene-Eocene Thermal Maximum (PETM) ~56 mya. Characterization of morphology in this genus has been limited by small sample sizes and fragmentary fossils. A new dental sample (n = 163) of the North American species Teilhardina brandti from PETM strata of the Bighorn Basin, Wyoming, documents previously unknown morphology and variation, prompting the need for a systematic revision of the genus. The P4 of T. brandti expresses a range of variation that encompasses that of the recently named, slightly younger North American species ‘ Teilhardina gingerichi ,’ which is here synonymized with T. brandti . A new partial dentary preserving the alveoli for P1-2 demonstrates that T. brandti variably expresses an unreduced, centrally-located P1, and in this regard is similar to that of T. asiatica from China. This observation, coupled with further documentation of variability in P1 alveolar size, position, and presence in the European type species T. belgica , indicates that the original diagnosis of T. asiatica is insufficient at distinguishing this species from either T. belgica or T. brandti . Likewise, the basal omomyiform ‘ Archicebus achilles ’ requires revision to be distinguished from Teilhardina . Results from a phylogenetic analysis of 1890 characters scored for omomyiforms, adapiforms, and other euarchontan mammals produces a novel clade including T. magnoliana , T. brandti , T. asiatica , and T. belgica to the exclusion of two species previously referred to Teilhardina , which are here classified in a new genus ( Bownomomys americanus and Bownomomys crassidens ). While hypotheses of relationships and inferred biogeographic patterns among species of Teilhardina could change with the discovery of more complete fossils, the results of these analyses indicate a similar probability that the genus originated in either Asia or North America.
... These fossils were collected as part of an ongoing effort led by crews from the University of Florida and Duke University to increase representation of small vertebrate fossils from sites bracketing and sampling the Paleocene-Eocene Thermal Maximum (PETM) in the Bighorn Basin of Wyoming (Wing et al., 2005;Chester et al., 2010;Secord et al., 2012;Baczynski et al., 2013;Bourque et al., 2015). Washakie Basin grooming phalanges Sorting through screen-wash concentrate from Wa3 and Wa5 localities of the Washakie Basin, D.M.B. recovered an additional three specimens (UCMP 217999, 218000, 218344) Table S1) and a total of 395 postcranial specimens attributable to omomyiforms based on their small size and comparisons to previously identified postcranial material of omomyiforms (Szalay, 1976;Savage and Waters, 1978;Dagosto, 1988;Gebo, 1988;Covert and Hamrick, 1993;Dagosto et al., 1999;Hamrick, 1999;Anemone and Covert, 2000;Dunn et al., 2006;Gebo et al., 2012), adapids (Gregory, 1920;Hamrick and Alexander, 1996), and extant primates. ...
Article
Euprimates are unusual among mammals in having fingers and toes with flat nails. While it seems clear that the ancestral stock from which euprimates evolved had claw-bearing digits, the available fossil record has not yet contributed a detailed understanding of the transition from claws to nails. This study helps clarify the evolutionary history of the second pedal digit with fossils representing the distal phalanx of digit two (dpII), and has broader implications for other digits. Among extant primates, the keratinized structure on the pedal dpII widely varies in form. Extant strepsirrhines and tarsiers have narrow, distally tapering, dorsally inclined nails (termed a 'grooming claws' for their use in autogrooming), while extant anthropoids have more typical nails that are wider and lack distal tapering or dorsal inclination. At least two fossil primate species thought to be stem members of the Strepsirrhini appear to have had grooming claws, yet reconstructions of the ancestral euprimate condition based on direct evidence from the fossil record are ambiguous due to inadequate fossil evidence for the earliest haplorhines. Seven recently discovered, isolated distal phalanges from four early Eocene localities in Wyoming (USA) closely resemble those of the pedal dpII in extant prosimians. On the basis of faunal associations, size, and morphology, these specimens are recognized as the grooming phalanges of five genera of haplorhine primates, including one of the oldest known euprimates (∼56 Ma), Teilhardina brandti. Both the phylogenetic distribution and antiquity of primate grooming phalanges now strongly suggest that ancestral euprimates had grooming claws, that these structures were modified from a primitive claw rather than a flat nail, and that the evolutionary loss of 'grooming claws' represents an apomorphy for crown anthropoids.
... Like their extant relatives, Eocene euprimates had digital pads with nails rather than claws, and a hind foot modified into a grasping organ with a divergent hallux. Whilst plesaidapiforms apparently engaged in more deliberate quadrupedalism , the limbs of earliest euprimate (Omomyiformes, Adapiformes, and Eosimiidae; Figure 4) were modified for more arboreal leaping, and above-branch quadrupedalism (Anemone & Covert, 2000;Dagosto, Gebo, & Beard, 1999;Gebo, Dagosto, Beard, & Qi, 2001;Gebo, Smith, & Dagosto, 2012;Ni et al., 2013;Rose & Walker, 1985). Rapid movement in earliest euprimates is not a reason to reject the visual-predation scenario as Gebo (2004) suggests. ...
... Despite this general diversity, the European fossil record is still scarce and poorly known. However, several recent works have reported new early Eocene fossil material that sheds light on the evolution of the first primates that inhabited the European continent (e.g., Gebo et al., 2012;Hooker, 2012;Marig o et al., 2012Marig o et al., , 2014. Particularly, regarding adapiforms, new material of the genus Agerinia has been lately documented from several early Eocene localities in the Ager Basin (Lleida province, NE Spain), including some additional remains of the previously known species Agerinia roselli from Les Saleres (Femenias-Gual et al., 2016b) and two new species, Agerinia smithorum from Casa Retjo-1 (Femenias-Gual et al., 2016a) and Agerinia marandati from Masia de l'Hereuet . ...
Article
New material attributed to Agerinia smithorum from Casa Retjo-1 (early Eocene, NE Iberian Peninsula), consisting of 13 isolated teeth and a fragment of calcaneus, is studied in this work. These fossils allow the first description of the calcaneus and the upper premolars for the genus Agerinia, as well as the first description of the P2 and M2 for A. smithorum. The newly recovered lower teeth are virtually identical to the holotype of A. smithorum and are clearly distinguishable from the other species of Agerinia. The upper teeth also show clear differences with Agerinia marandati. The morphology of the calcaneal remains reveals that A. smithorum practiced a moderately active arboreal quadrupedal mode of locomotion, showing less leaping proclivity than notharctines but more than asiadapids. All the morphological features observed in the described material reinforce the hypothesis of a single lineage consisting of the species A. smithorum, A. marandati, and Agerinia roselli. Furthermore, the phylogenetic analysis developed in this work, which incorporates the newly described remains of A. smithorum, maintains the position of Agerinia as closely related to sivaladapids and asiadapids.
... Recent, comprehensive analyses of ankle (Boyer and Seiffert, 2013;Boyer et al., 2013aBoyer et al., , 2015bYapuncich et al., 2017) and hand (Boyer et al., 2013b(Boyer et al., , 2016b anatomy of primates, as well as certain early fossil specimens (Gebo et al., 2012(Gebo et al., , 2015Ni et al., 2013) have begun to question the primacy of specialized grasping in the ancestral primate lineage, as they have pointed to the possibility that acrobatic leaping in the trees was emphasized first. Under this scenario the more generalized anatomy of euprimates from Vastan mine, India would represent specializations that postdate the initial radiation of euprimates from their common ancestor. ...
Article
The fossil record of early primates is largely comprised of dentitions. While teeth can indicate phylogenetic relationships and dietary preferences, they say little about hypotheses pertaining to the positional behavior or substrate preference of the ancestral crown primate. Here we report the discovery of a talus bone of the dentally primitive fossil euprimate Donrussellia provincialis. Our comparisons and analyses indicate that this talus is more primitive than that of other euprimates. It lacks features exclusive to strepsirrhines, like a large medial tibial facet and a sloping fibular facet. It also lacks the medially positioned flexor-fibularis groove of extant haplorhines. In these respects, the talus of D. provincialis comes surprisingly close to that of the pen-tailed treeshrew, Ptilocercus lowii, and extinct plesiadapiforms for which tali are known. However, it differs from P. lowii and is more like other early euprimates in exhibiting an expanded posterior trochlear shelf and deep talar body. In overall form, the bone approximates more leaping reliant euprimates. The phylogenetically basal signal from the new fossil is confirmed with cladistic analyses of two different character matrices, which place D. provincialis as the most basal strepsirrhine when the new tarsal data are included. Interpreting our results in the context of other recent discoveries, we conclude that the lineage leading to the ancestral euprimate had already become somewhat leaping specialized, while certain specializations for the small branch niche came after crown primates began to radiate.
... These bones reveal that Teilhardina was arboreal, with a hindlimb that was adapted for leaping and grasping. Intriguingly, the phalanges known for T. belgica are highly elongated like those of modern tarsiers, suggesting that Teilhardina may have engaged in tarsier-like hunting behavior, in which the giant hands function like baseball mitts to pin prey to the ground (Gebo, Smith, and Dagosto 2012). However, the development of molar shearing crests in Teilhardina is surprisingly weak in comparison to modern primates such as tarsiers that feed extensively on insects and other live animal prey. ...
Chapter
One of the oldest known fossil primates, Teilhardina is a primitive representative of the haplorhine clade that also includes living tarsiers and anthropoids. Species of Teilhardina have been described from earliest Eocene sites in Asia, North America, and Europe. A prominent episode of global warming known as the Paleocene–Eocene Thermal Maximum (PETM) allowed Teilhardina and other mammals to disperse rapidly across high-latitude corridors such as Beringia to achieve their broad geographic distribution. With an adult body mass of only approximately 30–60 grams, Teilhardina resembled the smallest living mouse lemurs in size. Teilhardina shows an interesting mosaic of anatomical features, including very primitive teeth and a hindlimb that was adapted for leaping and grasping. Its skull had relatively small orbits, suggesting a diurnal activity pattern. Isolated phalanges (finger bones) of Teilhardina show that it had very long fingers like those of living tarsiers.
... Recent works dealing with European Eocene primates have focused on the description of new material (Hooker, 2007;Hooker, 2012;Hooker & Harrison, 2008;Marigó, Minwer-Barakat & Moyà-Solà, 2011;Gebo, Smith & Dagosto, 2012;Gebo et al., 2015;Minwer-Barakat et al., 2013;Femenias-Gual et al., 2015), the revision of previous taxonomic assignations Minwer-Barakat, Marigó & Moyà-Solà, 2016;Marigó et al., 2014) and the establishment of relationships between different taxa (Smith, Rose & Gingerich, 2006;Minwer-Barakat et al., 2017), with some exceptions focused on the diet (Ramdarshan, Merceron & Marivaux, 2012), the locomotor behaviour (Marigó et al., 2016) and the endocranial anatomy (Ramdarshan & Orliac, 2016) of several species. However, only a few contributions have been published regarding European primates from the early Eocene, recently including the revision of Agerinia roselli from Les Saleres and the description of the new species Agerinia smithorum from Casa Retjo-1 (Femenias-Gual et al., 2016a andFemenias-Gual et al., 2016b, respectively). ...
Article
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Background The Eocene was the warmest epoch of the Cenozoic and recorded the appearance of several orders of modern mammals, including the first occurrence of Euprimates. During the Eocene, Euprimates were mainly represented by two groups, adapiforms and omomyiforms, which reached great abundance and diversity in the Northern Hemisphere. Despite this relative abundance, the record of early Eocene primates from the European continent is still scarce and poorly known, preventing the observation of clear morphological trends in the evolution of the group and the establishment of phylogenetic relationships among different lineages. However, knowledge about the early Eocene primates from the Iberian Peninsula has been recently increased through the description of new material of the genus Agerinia from several fossil sites from Northeastern Spain. Methods Here we present the first detailed study of the euprimate material from the locality of Masia de l’Hereuet (early Eocene, NE Spain). The described remains consist of one fragment of mandible and 15 isolated teeth. This work provides detailed descriptions, accurate measurements, high-resolution figures and thorough comparisons with other species of Agerinia as well with other Eurasian notharctids. Furthermore, the position of the different species of Agerinia has been tested with two phylogenetic analyses. Results The new material from Masia de l’Hereuet shows several traits that were previously unknown for the genus Agerinia, such as the morphology of the upper and lower fourth deciduous premolars and the P2, and the unfused mandible. Moreover, this material clearly differs from the other described species of Agerinia, A. roselli and A. smithorum, thus allowing the erection of the new species Agerinia marandati. The phylogenetic analyses place the three species of Agerinia in a single clade, in which A. smithorum is the most primitive species of this genus. Discussion The morphology of the upper molars reinforces the distinction of Agerinia from other notharctids like Periconodon. The analysis of the three described species of the genus, A. smithorum, A. marandati and A. roselli, reveals a progressive change in several morphological traits such as the number of roots and the position of the P1 and P2, the molarization of the P4, the reduction of the paraconid on the lower molars and the displacement of the mental foramina. These gradual modifications allow for the interpretation that these three species, described from the early Eocene of the Iberian Peninsula, are part of a single evolutionary lineage. The stratigraphical position of Masia de l’Hereuet and Casa Retjo-1 (type locality of A. smithorum) and the phylogenetic analyses developed in this work support this hypothesis.
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In biology, there is currently a debate being waged about the basic principles of doing taxonomy (e.g., Benton, 2000; Cantino and De Queiroz, 2000; De Queiroz, 1994, 1997; De Queiroz and Gauthier 1990, 1992, 1994; Lee, 1996; Lidén and Oxelman, 1996; Lide7acute;n et al., 1997; Moore, 1998; Nixon and Carpenter, 2000; Pennisi, 1996; Schander and Thollesson, 1995). This debate stems from the common opinion that taxonomy should reflect evolution in some manner, combined with a disagreement about the practical details of how to do this. Although some authors have provided suggestions for making the Linnean system of taxonomy work within the context of a cladistic approach to phylogeny reconstruction (e.g., McKenna and Bell, 1997; Nixon and Carpenter, 2000; Wiley, 1981), others have advocated scrapping the entire Linnean system (De Queiroz, 1994; De Queiroz and Gauthier, 1990, 1992, 1994; Griffiths, 1976), culminating in the dissemination by way of the Internet of a new code for phylogenetic nomenclature, the Phylocode (Cantino and De Queiroz, 2000). Although this document does not yet include guidelines for species taxa, and in spite of the fact that the Phylocode has not yet been "activated" by its authors (as of May, 2003; but see below), there are nonetheless a growing number of instances of the principles codified by this system being applied in real taxonomic practice (e.g., the redefinitions of Mammalia by Rowe, 1988). As such, even if the Phylocode is never adopted or accepted in full, it can still be considered to represent many current ideas about the practicalities of doing taxonomy. In light of these debates a reconsideration of the meaning, content, and status of the taxon name "Primates" seems timely. Anthropologists have sometimes been criticized for ignoring taxonomic principles common to other areas of Biology (e.g., Mayr, 1950; Simpson, 1963), and the fact that the intense debates over taxonomic practice that have been waged in the biological literature in recent years are only rarely reflected in the contemporary anthropological literature suggests that this problem is ongoing. One of the central goals in understanding primate origins must be forming an understanding of where the group lies in relation to non-primate groups, since only against that comparative background can the relative uniqueness of primate features be fully understood. Without such an understanding it is impossible to create plausible adaptive scenarios for why changes occurred in the early evolution of the group. In light of this, it is clear that anthropologists cannot work in a vacuum from current evolutionary and taxonomic practice as applied to other groups of mammals. Thus, it seems prudent to consider how Primates would stand in the context of the new system if the Phylocode were enacted, and how our common conceptions of what this term means could be dealt with in this framework. Even if the Phylocode is never accepted by all, it is worth considering the relative merits of the philosophical position that it represents. This has particular relevance in relation to the inclusion or exclusion of plesiadapiforms from the order Primates, since a determination of whether or not this cluster of extinct forms can be designated as primates depends not only on the supported pattern of relationships but also the taxonomic philosophy being applied. Finally it must be asked whether or not these disagreements over taxonomic approach influence the way in which we do, and should, ask questions about primate origins.
Article
Data obtained during recent excavations and analysis of fossil mammal collections from the Dormaal site have permitted assembly of a dozen cheek teeth of Diacodexis. The study of this material and the comparison with other primitive Diacodexis species of the early Eocene from Europe, North America and Asia made it possible to define the new species, Diacodexis gigasei. A phylogenetic position for this new species is proposed.
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Recognition of species in the fossil record is a critical issue for students of primate evolution. Since dental remains provide the largest samples for assessing intra- and interspecific variation, dental variation in fossil mammals is frequently compared with data for recent related species when investigating systematic diversity (Gingerich, 1974, 1979; Gingerich and Shoeninger, 1979; Kay, 1982a,b; Kay and Simons, 1983; Kelley, 1986; Kimbel and White, 1988; Martin and Andrews, 1984; Simpson, 1941a, and many others). The coefficient of variation (CV) has been the most frequently used statistic in a majority of these studies.