New postcranial elements for the earliest Eocene fossil primate Teilhardina belgica
Daniel L. Gebo
, Thierry Smith
, Marian Dagosto
Department of Anthropology, Northern Illinois University, DeKalb, IL 60115, United States
Direction Earth and History of Life, Royal Belgian Institute of Natural Sciences, B-1000, Brussels, Belgium
Department of Cell and Molecular Biology, Feinberg School of Medicine, Northwestern University, Chicago, IL 60611, United States
Received 1 July 2011
Accepted 14 March 2012
Available online 15 June 2012
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 proﬁle 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 ﬁngers similar to those of living tarsiers.
Ó2012 Elsevier Ltd. All rights reserved.
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
ﬁgured 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 ﬁrst 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 ﬂuvio-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,
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.
E-mail addresses: firstname.lastname@example.org (D.L. Gebo), email@example.com
(T. Smith), firstname.lastname@example.org (M. Dagosto).
Contents lists available at SciVerse ScienceDirect
Journal of Human Evolution
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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 ﬁeld 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 ﬂuviatile 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
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
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), Teilhardina’s 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 ﬁrst metatarsal, distal
femur, and middle phalanx measures are all larger. Compared with
the slightly larger mouse lemurs, Microcebus griseorufus or
List of new postcranial elements of T. belgica.
Catalog number Identiﬁcation 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 ﬁgures for Tables 3 and 6.
Figure 2. First metatarsal and phalangeal measurement ﬁgures for Tables 10 and
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
The dimensions of Teilhardina’slimb 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).
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 ﬂat 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 ﬂexor ﬁbularis,
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 tibioﬁbular mortise like that of other omomyids and
anthropoids (Dagosto et al., 2008). The lateral taloﬁbular 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 taloﬁbular facet is a well developed fossa for the
posterior taloﬁbular 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
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 ﬁgure for Table 11.
Comparison of Teilhardina dimensions to those of other small extant and extinct
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
2.15 1.73 2.59
2.40 1.87 3.30
Distal femur bicondylar
4.20 3.79 4.47 5.05
Distal femur lateral
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
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 signiﬁcant 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 signiﬁcant
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 ﬁve 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 ﬂattened 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-
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
ﬂexor 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
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
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
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
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 ﬁrst
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.
IRSNB M1262 (Fig. 7) is the proximal end of a ﬁrst 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 ﬁrst 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
ﬁrst metatarsal from Shanghuang (China) is still smaller in all
dimensions (Gebo et al., 2008).
The entocuneiform-ﬁrst 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 ﬁrst 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-ﬁrst
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 ﬁrst 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 ﬁrst 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 ﬂattened 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
Talar ratios in omomyids and microchoerids.
Talar neck length/
Lateral body height/
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 ﬁrst 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.
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
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
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.
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 ﬁngered 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 ﬂat
in both. They contrast in proximal joint height, being taller in IRSNB
M1264, and quite ﬂat 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 ﬂexor tendons that lie between. The elevated rims
provide prominent attachment sites for the ﬂexor retinacular
sheaths to keep these tendons within their ﬁbrous tunnels. Flexor
Measures of variance in calcaneal length and width in Teilhardina and other small
Taxon NCalcaneal length Calcaneal width
Dormaal 6/9 4.6 113.3 12.6 8.5 137.8 30.3
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
Amboasary 14 3.9 112.5 11.8 6.1 121.7 19.6
Covert quarry 5/3 6.6 119.1 17.5 1.7 103.1 3.0
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).
Calcaneal comparative data.
Length Width PACL PACW CUHT CUW HeelL Distal
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
abderhaldeni n ¼1
3.27 2.06 1.31 2.18
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
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).
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
x x x x 46.4 x
56.4 29.3 23.3 19.9 59.5 x
53.0 32.3 23.7 25.6 58.1 111.6
x x x x 62.6 x
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 ‘hallucal’phalanx 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
signiﬁcantly 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 ﬁrst digit proximal phalanges of Teilhardina
are robust, suggesting large muscle attachments and forceful
grasping. The ‘pollical’specimen is long, as in Tarsius, suggesting
elongated manual digits.
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 deﬁnitively
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
syrichta’spedal 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 ﬁngers 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, ﬁrst metatarsal of T. belgica (from left to right: proximal, dorsal and plantar views (top); medial and lateral views (below)).
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
The distal joint surface of the two middle phalanges of T. belgica
is very cylindrical in shape and is very ﬂat 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 ﬂexor
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
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 ﬁrst metatarsal
joint, the long peroneal tubercle on the ﬁrst 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
ﬂattened distal phalanges conﬁrm 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).
Distal femoral measurements (mm) and ratios.
IVPP V13017 Omomys
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
Measurements (mm) for individual phalanges of T. belgica.
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
Tarsiers are distinctive among living primates in having
enlarged ﬁnger pads, long ﬁngers and hands, and an arched ﬁnger
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 ﬁngers to pin down prey items on the forest ﬂoor
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 ﬁngers, 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).
Proximal pollical phalangeal mean ratio and range comparisons.
Tarsius syrichta Galagoides demidovii Galago
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
Proximal hallucal phalangeal mean ratio and range comparisons.
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.
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 conﬂict 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 taloﬁbular facets (Gebo, 1986), fan-shaped
calcaneocuboid joints (Gebo, 1988), tall knees with elevated
lateral patellar rims, and robust ﬁrst 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
Mean ratio values and ranges for middle phalanges (3 ¼digit three).
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 taloﬁbular facet or the
position of the ﬂexor 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
ﬁrst 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
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).
As one of the few early fossil primatesprimitive enough to possess
four premolars, T. belgica represents a signiﬁcant 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 conﬁrm 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.
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 ﬁnan-
cially supported by the Federal Science Policy Ofﬁce 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.
Comparison of postcranial features in two primitive early Eocene primates.
Teilhardina belgica Cantius sp.
1. Talar neck length Long Short
2. Medial talotibular facet Full Full
3. Taloﬁbular facet Steep-sided Oblique
4. Posterior trochlear shelf Small Large
5. Location of ﬂexor hallucis
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
10. Height of distal femur Tall Tall
11. Lateral patellar rim Elevated Elevated
12. Patellar groove width Narrow Wide
13. First metatarsal shape Robust Robust
14. Peroneal tubercle length Long Long
15. Peroneal tubercle shape Narrow Broad
16. First metatarsal-entocuneiform
17. Proximal phalangeal length Longer Long in Notharctus
18. Middle phalangeal length Longer Long in Notharctus
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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
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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