AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 73:9-31(1987)

Functional Anatomy of the Tarsier Foot

DANIEL L. GEBO
Department of Anthropology, Duke Uniuersity, Durham,
North Carolina 27706

KEY WORDS
anatomy

Tarsiidae, Galaginae, Omomyidae, Locomotion, Foot

ABSTRACT
Tarsiers possess a very odd musculoskeletal foot anatomy that
goes beyond their acknowledged specialized leaping adaptations. Tarsius has
evolved a fundamentally different method of bone rotation to achieve a n
inverted foot position during grasping and has developed a n unusual muscular
system for holding onto vertical supports. Although galagos and tarsiers possess elongated foot bones a s adaptations for leaping, galagos utilize many more
types of movements, have specialized osteological surfaces for inversion, and
have a more common type of muscle development in the foot and leg than
tarsiers possess. Likewise, the Omomyidae, the ancestral lineage of Tarsius,
exhibit a lack of morphological similarity with Tarsius in the known foot
joints.
Tarsier foot anatomy is far more astonishing than the widely acknowleged leaping adaptations of the elongated calcaneus and
navicular (Morton, 1924; Hafferl, 1932;
Clark, 1959; Schultz, 1963; Hall-Craggs,
1966; Napier and Walker, 1967; Martin, 1972;
Szalay, 1976; Jouffroy and Lessertisseur,
1979; Jouffroy et al., 1984). Tarsiers have
evolved a fundamentally different method of
bone rotation to achieve a n inverted foot position and have developed a n unusual muscular system for holding on to vertical
supports. This paper will discuss foot function in tarsiers by examining foot postures
and locomotion, and tarsier musculoskeletal
anatomy, which includes the crucial information on joint structure and function. Two
questions are particularly important for this
paper: 1)How are tarsiers different from galagos in foot anatomy and function since both
have elongated calcanei and naviculars; and
2) how does Tarsius foot anatomy compare
with the Omomyidae, the ancestral lineage
of tarsiers?
MATERIALS AND METHODS

The biggest problem in developing a project on prosimians is related to the access and
use of these rare materials. Duke University
has a Iarge and diverse collection of animals,
skeletons, and cadaver specimens a t the

0 1987 ALAN R. LISS, INC.

Duke Primate Center. Two species of tarsiers, Tarsius bancanus and Tarsius syrichta,
and four species of galagos, Galago demide
vii, Galago senegalensis, Galago garnettii,
and Galago crassicaudatus, are present at
the Duke Primate Center.
Four of the six species noted above at the
Duke Primate Center were observed and
movement was recorded in terms of frequency of locomotor bouts. Caging was
judged to be appropriate for movement studies after consideration of the following variables: 1)the size of the animal; 2) the number
of animals within the cage; 3) the arrangement of the supports within the cage; and 4)
the amount of space for freedom of movement. Each cage had a variety of horizontal,
oblique, or vertical supports which were arranged for movement diversity. Observations were recorded on focal animals when
the animals were active and moving following the technique of Fleagle (1976, 1978;
Fleagle and Mittermeier, 1980; Mittermeier,
1978). All animals were observed according
to their artificial daylnight light schedule for
evenings. Only adults were observed. More
Received J u n e 30, 1986; revision accepted December 22, 1986.
Address reprint requests to D. Gebo, Dept. ofCe11 Biology and
Anatomy, The Johns Hopkins University, School of Medicine,
Baltimore, MD 21205.

10

D.L. GEBO

details of the methodology are explained in

Gebo (1986).
Three species of Galago and one species of
Tarsius were dissected at the Duke Primate
Center. All cadavers were animals which had
been frozen immediately following death. The
relationships of foot musculature, ligaments,
and retinacula were noted. All muscles were
dissected away from the leg and foot, tendons
cut, and excess fascia material removed.
Next, the muscles were immersed in water,
blotted dry, and then weighed to the nearest
0.01 gm. All muscle weights were divided by
the total extrinsic or intrinsic musculature
weight of the leg or foot for each animal. This
gives a relative weight figure for each muscle
that is comparable among different-sized animals and among species. Tuttle (1972) has
discussed previous quantitative studies of
muscle mass and the advantages of muscle
ratios for intertaxonal comparisons. Although wet muscle weights may not be as
accurate as dry muscle weights, the quickness and ease of the wet muscle technique,
and the use of relative weights in the analysis rather than the actual wet muscle
weights, offer a valuable and empirically justifiable approach. Care should be exercised,
however, in interpreting the functional significance of muscle weights because of the
small sample sizes. The total extrinsic and
intrinsic muscle weights are provided below
so that the raw muscle weights may be calculated from the relative weight values provided in the text.
Species

G. crassicaudatus
G. senegalensis
G. garnettii
T syrichta

2
1
2
2

Extrinsic Intrinsic

(PI

(PI

19.17
1.96
14.36
1.14

2.39
0.38
1.55
0.31

Xeroradiographs were taken at the North

Carolina Veterinary School by Earl Del
Santo. Feet were taped into positions of plantarflexion, dorsiflexion, inversion, and eversion. All xeroradiographs were taken on
cadavers of G. senegalensis, G. crassicaudatus, and II: syrichta. The xeroradiographs provided excellent views of joint and bone
positions within the context of a n intact foot.
Besides the xeroradiographs, individual foot
bones and ligamentous preparations were examined and compared in order to understand
form and function across taxa.

\\

Fig. 1. Normal foot position for Tarsius on a vertical

support.

Tarsier and galago behavioral differences

Foot positions: Tarsiers and galagos, like
other prosimians, utilize two basic types of
foot postures with some variation owing to
the nature of the support. The basic prosimian foot position on a horizontal arboreal support has the big toe on the opposite side of
the support from the other lateral digits. The
distal part of the foot beginning at the calcaneo-cuboid joint and the navicular-cuneiform
joints is dorsiflexed, leaving the heel elevated above the support (Fig. 1).The digits
curl around and grasp the support. The foot
is inverted and slightly abducted (Fig. 1).
Weight is borne by the plantar surfaces of
the navicular, entocuneiform and the cuboid.
On a vertical support, the foot assumes a
more abducted (peroneal) position. When a
foot is placed on the ground, the foot assumes
a more everted and adducted position with
the heel still elevated above the ground's
surface.
Tarsiers show a modification of the vertical
foot position in their digits. The metatarsalphalangeal joint is hyperextended, the interphalangeal joint between the proximal and
the middle phalanges is flexed, and the interphalangeal joint between the middle and the
terminal phalanges is extended. These joint
positions place the digits into a n elevated
arch which is used to place more pressure on
the large digital pads (Fig. 2).
The second type of foot posture, the suspension grasp, is used when a prosimian hangs
below a branch using only its hind feet. This

TARSIER FOOT ANATOMY

Fig. 2. Elevated arch position for distal, middle, and

proximal phalanges in tarsiers.

foot position has the distal phalanges of the

second, third, fourth, and fifth digits hooked
above the horizontal support. The big toe is
flexed but does not participate in the grasp.
Tarsiers do not utilize this second type of foot
posture since they never suspend their body
below a support using only their hind feet as
galagos do.
Movement: Tarsiids are specialized leapers
and frequent climbers but are more confined
in the variety of movements compared to galagos (Gebo, 1986).Tarsiers never suspend below a branch using only two feet, nor do they
bridge across two supports. Galagos, on the
other hand, utilize many more types of movements in greater frequencies (Table 1). Galagos employ two movement gradients, one for
leaping and one for quadrupedalism. The
smallest galago, G. demidouii, and the largest, G. crassicaudatus, emphasize more quadrupedal tendencies, while the other galago
species are more frequent leapers.

Tarsiids
Tarsius syrichta is a vertical clinger and
leaper. Leaps make up 61% of all movements
(Table 1). Most other movements are rare in
I: syrichta with the exception of climbing. I:
syrichta will move using quadrupedalism,
hang suspended below a n oblique branch
after a leap, and will on occasion bimanually
pull itself up upon a branch. Tarsiers never
suspend below a horizontal branch, nor was
T syrichta observed to bridge across supports. 1: syrichta climbs downward tailfirst
and will slide down bamboo by letting go of
its grasp and sliding downwards. This sequence of sliding can be repeated consecutively several times. l? syrichta climbs
upwards with a side-to-side waddle. I: syrzchta will hold onto a vertical support with
only its two hands and stretch its legs out

11

12

D.L. GEBO

nique, 1977).Table 1shows that G. demidouii

is predominantly a leaping species with
quadrupedalism and climbing being the next
two most frequent movements. Quick rather
than deliberate quadrupedal movements are
made by G. demidovii. G. demidovii is more
suspensory than G. senegalensis a t the Duke
Primate Center (Table 1).Walker (1979) recorded movement frequencies for G. demide
uii which compare well with my observations
(Table 1). G. demidovii climbs downward
headfirst, performs the cantilevering suspensory movement, and moves on the ground
using quadrupedalism or by bipedal hops.
Galago senegalensis inhabits the lower forest levels (Crompton, 1984) and is a lightning-quick power leaper. Long fast leaps are
common from vertical supports. Crompton
(1984) states that G. senegalensis vertically
clings 50% of the time. Climbing is the only
other movement G. senegalensis performs
with any frequency above 5% at the Duke
Primate Center (Table 1). G. senegalensis
normally climbs downward headfirst but will
on occasion climb tailfirst. Crompton (1984)
records movement frequency data for G. senegalensis, and his study has a much greater
frequency for quadrupedalism than my study,
in which quadrupedalism is fairly rare (Table 1).Cromptons frequency for quadrupedalism in G. senegalensis is comparable to G.
demidovii. G. senegalensis prefers to hop bipedally on the ground. Suspensory movements are not common, although G.
senegalensis does cantilever. When cantilevering, galagos will fully stretch their body
away from the vertical support. The body is
extended outward like a n accordian with the
knees fully extended and the arms stretched
outward to grab a n insect. Only the hind feet
are attached to the vertical support. The cantilevering position is like a flag which extends away from the vertical flagpole. G.
senegalensis uses vertical supports 16%, horizontal supports 27%, and oblique supports
55% of the time (Crompton, 1984).
G. alleni prefers medium-diameter supports from 1 to 5 cm and uses vertical supports 81% of the time (Charles-Dominique,
1977). G. alleni prefers to leap from vertical
support to vertical support. This species is an
infrequent user of quadrupedalism, although
G. alleni moves using: auadruoedalism or biGalagines
pedal hopping when &-the g k n d (CharlesGalago demidovii uses vertical supports Dominique, 1977).
Galago crassicaudatus prefers to move us48%, oblique supports 30%, and horizontal
supports 22% of- the time (Charles-Domi- ing quadrupedalism and climbing before

without grasping the support and rub its legs

across the vertical support. Tarsiers, holding
on only by their hind feet and using the basic
prosimian foot posture, will lean out from a
vertical support to catch insects with their
hands. This is different than cantilevering
since the knees are never fully extended, nor
is the body extended very far away from the
vertical support.
Niemitz (1984) has discussed the movements of 7: bancanus, which are similar to 7:
syrichta. My observations differ somewhat
from this view. Table 1indicates that 7: bancanus does not utilize the more varied movements of T syrichta. 7: bancanus prefers to
leap and climb, and will stretch out to grab
crickets. No quadrupedalism or bipedalism
was ever observed for T bancanus. T bancanus is less active than 7: syrichta in terms
of overall movement activity and prefers to
vertically cling while at rest. T syrichta, on
the other hand, rests on horizontal supports
most often. Niemitz (1984) notes one occurrence ih which ?: bancanus moved by means
of below-branch quadrupedalism. Niemitz
(1984) also describes a movement called bipedal climbing in which T bancanus leaps
upward and grasps hold of the same support,
thereby moving up the support. 7: syrichta
performs this same movement, but I scored
this movement as a leap, rather than a
climbing sequence in Table 1.
Tarsius spectrum is stated by MacKinnon
and MacKinnon (1980) to be less specialized
than 7: bancanus in that it climbs, scurries,
runs, leaps, and engages in loris-like clambering more often than I: bancanus. 7: spec
trum leaps 63%, climbs 14%, and hop/runs
8%of the time (MacKinnon and MacKinnon,
1980). These frequencies are similar to my
observations for T. syrichta (Table 1).Leaps
of 5-6 m in length have been recorded for
this species (MacKinnon and MacKinnon,
1980). ?I spectrum prefers to rest on oblique
or horizontal supports. T spectrum is more
active at many different heights (0-9 m) than
is T bancanus (MacKinnon and MacKinnon,
1980). T spectrum prefers to move on vertical
and larger-diameter supports (MacKinnon
and MacKinnon, 1980). Use of larger-diameter supports is perhaps related to the pointy
nails of this species.

13

TARSIER FOOT ANATOMY

leaping (Crompton, 1984) (Table 1). Leaping

is about half as frequent a s it is for G. sene
galensis but is still a considerable amount
for G. crassicaudatus. Richochetal leaping is
rare for G. crassicaudatus. G. crassicaudatus
moves using quadrupedalism or hops when
on the ground (Crompton, 1984). Bridging is
more common in G. crassicaudatus than in
G. dernidovii or G. senegalensis. Suspension
by four limbs and cantilevering are movements which G. crassicaudatus can perform
but uses rarely. G. crassicaudatus prefers
oblique supports 63% of the time; then horizontal supports, 31%; and lastly, vertical supports, 4% (Crompton, 1984).
Galago garnettii prefers horizontal and
larger-diameter supports than G. crassicaudatus (Carolyn Harcourt, pers. comm.). G.
garnettii moves with a greater use of hopping, richochetral leaping, and crouch leaping than G. crassicaudatus (Carolyn Harcourt, pers. comm.).
Galago elegantulus moves in the higher
layers of the forest and rarely comes to the
ground (Charles-Dominique, 1977).Its keeled
nails allow it to use large vertical trunks for
gum feeding. G. elegantulus prefers oblique
supports, 51% of the time; the vertical supports, 27%; and finally horizontal supports,
22% (Charles-Dominique, 1977). G. elegantulus climbs downward headfirst. Leaps are
common. Branch-running quadrupedalism
has been noted most often for this species'
movement preference (Charles-Dominique,
1977).
Galago and Tarsier foot musculature
Extrinsic foot musculature: Relative muscle weights for the extrinsic musculature
show differences between galagos and tarsiers which reflect in part the movement differences noted above. First, tarsiers possess
a larger gastrocnemius and slightly larger
soleus and plantaris muscles (Table 2). The
greater emphasis of the superficial flexor
musculature over the deep flexor muscula-

ture illustrates the greater emphasis in musculature associated with leaping over
musculature associated with grasping and
climbing adaptations (Table 3). This does not
mean than muscles associated with grasping
or climbing do not exist or do not function in
ta-siids. Clearly, these muscles exist, but
they are not as well developed in tarsiids as
they are in galagines.
The flexor hallucis longus and the flexor
digitorum longus are much larger for galagines than tarsiids (Table 2). The deep flexors
correlate with a more varied movement repertoire. If we combine the three superficial
flexors-the gastrocnemius, the soleus, and
the plantaris-and compare their relative
weight values to the three deep flexors-the
flexor hallucis longus, the flexor digitorum
longus, and the posterior tibialis-galagos
and tarsiids show opposite patterns of muscle
development (Table 3). Tarsiers have about
twice the amount of superficial flexors to deep
flexors, which is probably correlated with
their strong tendency for leaping. Galagines
have about equal amounts of both groups of
flexor musculature, with a bit greater weight
values for the superficial flexors.
Galagines are very different in the packaging of their lower leg musculature in comparison to tarsiids. Galagines are in fact
much more similar to lemurids and cheirogaleids (Gebo, 1986) since the muscles follow
the same relative weight ranking-gastrocnemius (0.20), flexor hallucis longus (0.171,

T A B L E 3. Relative weight values for certain leg

muscles of galagines (GI and tarsiids (Ti

Superficial flexors'

Deep flexors'
Total

0.33
0.29
0.62

0.42
0.22
0.64

'Superficial flexors include gastrocnemius, soleus, and plantaris

muscles.
2Deep flexors include flexor hallucis longus, flexor digitorum
longus, and the posterior tibialis muscles.

T A B L E 2. Extrinsic relative muscle weights for galagines (GI and tarsiids (T)','

FHL

FDL

PT

AT

EHL

EDL

PL

PB

0.20
0.26

0.06

0.07

0.08

0.08

0.17
0.13

0.09
0.05

0.03
0.04

0.15
0.16

0.04
0.05

0.02
0.04

0.14
0.06

0.04
0.04

'Abbreviations: G = gastrocnemius; S = soleus; P = plantaris; FHL = flexor hallucis longus; FDL = flexor digitorurn
longus; PT = posterior tibialis; AT = anterior tibialis; EHL = extensor hallucis longus; EDL = extensor digitorum
longus; PL = peroneus longus; PB = peroneus brevis.
2n = 2 specimens of Galago crassicaudatus; 2 specimens of Galago garnettii; 1 specimen of Galago senegalensis; 2
specimens of Tarsius syrichta.

14

D.L. GEBO

TABLE 4. Extrinsic and intrinsic relative muscle

weights by species'

S
P
FHL
FDL
PT
AT
EHL
EDL
PL
PB
EDBR
FDBR
AB I
AB V
AB OSSIS
FL I
FL V
OPP
AD HALL
CONTRA
LUM
DOR INTER
PLAN INTER
n

GC

GP

Gs

Ts

0.18
0.11
0.07
0.17
0.10
0.04
0.12
0.03
0.02
0.13
0.03
0.12
0.06
0.04
0.11
0.04
0.12
0.04

0.18
0.05
0.06

0.24
0.03
0.07
0.13

0.26
0.08
0.08
0.13
0.05

0.20

0.09
0.03
0.18
0.03
0.02
0.14
0.03

0.07
0.03
0.13

0.14
0.05
0.03
0.10
0.04
0.12
0.03

0.22
0.05
0.03
0.09
0.07
2

0.23
0.04
0.03
0.09
0.10
2

0.05
0.03
0.16
0.06
0.08
0.08
0.05
0.11
0.05
0.11
0.03
0.21
0.05
0.05
0.11
0.08
1

0.04
0.16
0.05
0.04
0.06
0.04
0.03
0.06
0.13
0.13
0.03
0.13
0.06
-

0.19
0.06
0.03
0.06
0.06
2

'All muscle abbreviations are listed below. The species

abbreviations are the same as in Table 5. Extrinsic foot
musculature: G = gastrocnemius; S = soleus; P = plantaris;
FHL = flexor hallucis longus; FDL = flexor digitorum longus;
PT = posterior tibialis; AT = anterior tibialis; EHL = extensor
hallucis longus; EDL = extensor digitorum longus; PL =
peroneus longus; PB = peroneus brevis. Intrinsic foot
musculature: EDBR = extensor digitorum brevis; FDBR = flexor
digitorum brevis; AB I = abductor hallucis; AB V = abductor
digiti quinti; AB OSSIS = abductor ossis metatarsi quinti; FL I
= flexor hallucis brevis; FL V = flexor digiti quinti; OPP =
opponens digiti quinti; AD HALL = adductor hallucis; CONTRA
= contrabentes; LUM = lumbricals; DOR INTER = dorsal
interossei; PLAN INTER = plantar interossei.

TABLE 5. Ratios of muscle weight values'

Superficial flexors
Deep flexors
Extensors
Deep flexors
Intrinsic musculature
Extrinsic musculature

Gc

Gg

Gs

Ts

1.33

1.00

1.70

2.33

0.19

0.17

0.40

0.50

0.12

0.11

0.19

0.27

'Abbreviations: Gc = Galago crassicaudatus; Gg = Galago

gamettii; Gs = Galago senegalensis; T s = Tarsius syrichta.

anterior tibialis (0.151, peroneus longus (0.141,

and flexor digitorum longus (0.09) (Table 2).
Galagos show the importance of their leaping musculature followed by the inverting
grasping musculature of the foot.
Galagirie extrinsic muscle weights stand
out for four muscles: the gastrocnemius, the
flexor hallucis longus, the anterior tibialis,

and the peroneus longus. Among the galagines, Galago senegalensis possesses a much
larger gastrocnemius than does Galago crassicaudatus and Galago garnettii but smaller
flexor hallucis longus and flexor digitorum
longus relative weight values (Table 4).Galago crassicaudatus has very large soleus
muscle, while Galago garnettii has the largest flexor hallucis longus of the three species.
The anterior tibialis is important for inversion and dorsiflexion and this muscle is large
for both tarsiids and galagines. The peroneus
longus muscle adducts the great toe during a
grasp and is especially important when utilizing more gravity-burdened postures on
oblique and vertical supports. The peroneus
longus muscle is very large in galagos but
tiny in tarsiids (Table 4).
The superficial flexorsldeep flexor ratio
(Table 5) shows Galago senegalensis with the
highest ratio value; Galago crassicaudatus is
intermediate; and Galago garnettii is last for
the galagines. The first two species are well
above 1.0 for this ratio, whereas Galago garnettii has essentially equal amounts of superficial and deep flexors. Tarsius has the
highest ratio of all prosimians for this ratio
(Gebo, 1986).
The extensorsldeep flexors ratio ranks Galago senegalensis second (0.40) to Tarsius
(0.50) at the high end of the ratio, which
perhaps supports the notion of extreme dorsiflexion of the foot for the use of vertical
supports. The other two galago species, G.
garnettii and G. crassicaudatus, are much
lower for this ratio and are similar to lorisines (Gebo, 1986),which are at the low end
of this ratio. The extensorldeep flexor ratio
supports the limited use of vertical supports
by the large-bodied galagines and their preference for horizontal substrates.
The larger-bodied galagos, Galago crassicaudatus and Galago garnettii, are more similar in their functional associations than they
are to Galago senegalensis. This is perhaps
due to the different nature of support usage
by these two groupings and in their movement preferences. The large-bodied galagines use horizontal supports and move
quadrupedally far more often than the smallbodied galagines which like vertical supports
and prefer to leap most of the time.
The top five muscle rankings for tarsiid
muscle development in the leg are as follows-gastrocnemius (0.261, anterior tibialis
(0.161, flexor hallucis longus (0.131, soleus and
plantaris (0.081, and peroneus longus (0.06)
(Table 2). The increase in the anterior tibialis

TARSIER FOOT ANATOMY

15

In Tarsius, the contrahentes musculature

is supportive of extreme dorsiflexion at the
ankle and an inverted foot in the vertically inserts well up the medial and lateral phaclinging posture, while a small flexor digito- langeal sides. Day and Iliffe (1975) discuss
rum longus (Table 4) indicates a decline in this unique insertion pattern for the hand of
suspensory postures (Gebo, 1986). Tarsiers Tarsius. This elaborate muscle arrangement
never hang by their feet or use suspensory allows Tarsius to hyperextend the metatarmovements very often, and the flexor digito- sal-phalangeal joint and flex the interphalrum longus muscle is tiny in tarsiers com- angeal joint of the middle and terminal
pared to galagos. The flexor hallucis longus phalanges. This puts pressure on the large
is an important muscle in Tarsius, but has toe pads, which helps the foot hold onto slick
the lowest value compared to other prosim- vertical surfaces like bamboo.
Tarsiers rank well above the other species
ian families (Gebo, 1986). Even if we add the
values for the flexor hallucis longus and the in comparing their total intrinsic muscle
flexor digitorum longus together, Tarsius is weight to their total extrinsic muscle weight.
still well below the other prosimian families This ratio is 0.27 for Tarsius and Tarsius is
(tarsiids =0.18; galagines and lemurids followed by Galago senegalensis (0.19), then
=0.26; cheirogaleids=0.28; indriids=0.34, G. crassicaudatus (0.12), and finally G. garnettii (0.11) (Table 5). The increase in intrinand lorisines = 0.43) Gebo, 1986).
How does a tarsier which grasps and clings sic foot musculature seems like a likely
to vertical supports, where the requirements adaptation for the use of vertical supports.
for holding onto a support (i.e., resisting
Retinacula
gravity) are the greatest, manage like a vertically clinging galago to maintain its hold
There are three basic bands of retinacula
with such small grasping musculature? Tar- for the prosimian leg and foot-the flexor resiers have evolved a different adaptive grasp- tinaculum, the peroneal retinacula, and the
ing strategy than other prosimians. Tarsiers extensor retinacula. The flexor retinaculum
have greatly increased their intrinsic grasp- extends from the medial calcaneus t o the
ing musculature. The intrinsidextrinsic distal tibia, where it joins the posterior part
muscle ratio shows tarsiers to have a value of the superior tibia1 extensor band (Fig. 3).
of 0.27 (Table 5). Galago senegalensis has a This posterior band holds the tendon of the
higher intrinsidextrinsic muscle ratio than posterior tibialis close to the tibia, while the
most other prosimians, and this value is ex- flexor retinaculum keeps the flexor digitoactly equal to the value for Propithecus (0.19) rum longus tendon, which runs behind the
(Gebo, 1986). Tarsius, Propithecus, and Gal- posterior tibialis, held firmly in place. The
ago senegalensis all utilize vertical supports tendon of flexor hallucis longus is also held
more often than the other prosimian species. by the widest part of the flexor retinaculum
An increase in intrinsic foot musculature band.
which resists the greater forces produced
The peroneal retinacula are divided into
from the interaction of body mass and grav- superior and inferior bands (Fig. 3). The suity when grasping a vertical support is a perior peroneal retinaculum wraps around
likely adaptation for these three species, with the lateral side of the fibula, and this band
tarsiers being more extreme in this adapta- holds the tendons of the peroneal musculation than the others. In addition, tarsiers ture firmly in place. The inferior peroneal
have an elaborate contrahentes musculature retinaculum is attached at the peroneal tuwhich functions to increase frictional pres- bercle of the calcaneus. This retinaculum has
sure on their enlarged toe pads.
two separate loops: one for peroneus brevis,
Intrinsic musculature: Galagines have in- peroneus digiti quarti, and peroneus digiti,
trinsic relative muscle weights that are sim- and one for peroneus longus. The tendons of
ilar to tarsiids. Both utilize the I-V grasp, peroneus longus and peroneus brevis do not
which emphasizes musculature which in- touch. The peroneal retinacula and the flexor
serts on the first and fifth digits (Gebo, 1986). retinaculum are similar in galagines and
Tarsiids have a much greater relative weight tarsiids as they are in all prosimians (Gebo,
value for abductor hallucis and slightly 1986).
The extensor retinacula have a more intrigreater values for flexor hallucis and flexor
digiti quinti than galagines (Table 4). Gala- cate organization to hold the extensor tengines have a much larger extensor digitorurn dons close to the lower leg and foot. The
brevis muscle and larger interossei muscles extensor retinacula are a complex of fibrous
bands
with different Datterns for different
than tarsiers possess.
~~.
~~

16

D.L. GEBO

Fig. 3 . Extensor retinacular band patterns. 1: Basic

pattern, common to lemurids, cheirogaleids, and lorisines. 2: Indriid extensor retinacular pattern. 3: Tarsiid
extensor retinacular pattern. 4: Galagine extensor retinacular pattern. Bands A-E are described in the text.

Band C is presented diagrammatically and should run

behind E as it is represented in panel 3. AT = anterior
tibialis and extensor hallucis longus tendons; EDL =
extensor digitorum longus tendon.

TARSIER FOOT ANATOMY

prosimian families. The most common extensor retinacula pattern is apparent for lemurids, cheirogaleids, and lorisines. Three main
and two smaller bands are observed at the
distal lower leg and foot for these three
groups (Fig. 3). Band A, the superior tibial
retinaculum, originates along the medial and
anterior sides of the distal tibia. Band B, the
superior fibular-tibia1band, begins higher up
on the lateral side of the fibula and runs at
an oblique angle until it meets with band A
at the tibia. The extensor digitorum longus
tendon is held close to the fibula by band B,
while this tendon runs above the C band at
the talar region. Band C, the inferior tibialcalcaneal retinaculum, which is a continuation of band B in most species, runs at an
oblique angle from the intersection area of
bands A and B, across and behind the lateral
talofibular facet to the calcaneus (Fig. 3). At
the anterior distal tibia, where bands A and
B meet, band B curves downward into band
C. Bands B and C form a triangular loop of
attachment with the point of the triangle at
the anterior tibia and the ends splayed wide
apart at the fibula and the calcaneus. Some
species show bands B and C to be distinctly
separate bands which are fused to band A,
where band B is the top and band C is the
bottom of the retinacular loop holding the
anterior tibialis and the extensor hallucis
longus tendons (Fig. 3). This arrangement
forms the working end of the triangular loop
in a different manner but is functionally similar. Band A is fused to the medial side of
this loop.
The two smaller retinacular bands originate from the anterior calcaneus before the
posterior calcaneal facet. Band D is a little
calcaneal loop which fuses with band E on
the lateral calcaneal side. Bands D and E
form the roof above the extensor digitorum
longus tendon which runs along the lateral
talofibular facet. Band E attaches at two separate points on the talus, and the extensor
digitorum longus tendon runs between these
two points of attachment. The floor of this
retinacular arrangement is made by band C
as it runs behind and below the extensor
digitorum longus tendon t o its calcaneal insertion (Fig. 3). Perodicticus shows a slight
variation wgere the D band is fused to the C
band along the normal calcaneal attachment
area for band D.
Indriids possess an extensor retinacular assembly very similar to the above but have
added an extra band (A') above the superior
extensor A and B bands (Fig. 3). The A' band

17

may be a functional response to indriid use

of vertical supports and their prolonged use
of a dorsiflexed foot posture.
In tarsiers, the A and B bands have combined into a broad oblique band. There is also
a smaller B' band from the fibula to the A-B
band (Fig. 3). The C band is separate but
connected to the A-B band. The anterior tibialis and extensor hallucis longus tendons
are held in a similar triangular loop as described above. The broad A-B band in Tarsius may help to hold the extensor tendons
in place better during extreme dorsiflexion
of the ankle. This unique A-B band supports
the notion of increased use of ankle dorsiflexion for tarsiers as previously noted in the
extensor musculature section.
Galagos are very different in the way they
hold the anterior tibialis and the extensor
hallucis longus tendons in place. Galagos
have an extra C' band, which forms the bottom part of a calcaneal loop made by bands
C and C' (Fig. 3). This loop originates from
the lateral calcaneus, extends to the anterior
distal tibia, and curves back toward the lateral calcaneus, where it inserts proximal to
its origin. The C and C' bands attach behind
the lateral talofibular facet on the lateral

Fig. 4. Diagram of the navicular-cuneiform articulations in Tarsius illustrating the navicular-mesocuneiform ligament (black rectangle) in tarsiers. N =
navicular; En = entocuneiform; M = mesocuneiform; Ec
= ectocuneiform; and 1 = first metatarsal.

18

D.L. GEBO

calcaneus. The tendon of extensor digitorum

longus runs above the C and C bands. Band
B does not participate in holding the anterior
tibialis or extensor hallucis longus tendons
in place. All three species of galagos possess
this calcaneal loop rather than the triangular loop of other prosimian feet. There seems
to be no apparent functional reason for this
difference, although the calcaneal loop could
perhaps be more efficient in reducing tension. Galagos, unlike indriids and tarsiids,
do not have any extra extensor bands above
the A band, but most species are frequent
users of vertical supports where extreme ankle dorsiflexion occurs. This novel retinacular loop, like the extra indriid and tarsiid
extensor bands, could perhaps function in a
similar way. Galago species, which are infre-

quent users of vertical supports (e.g., G. crassicaudatus), have the same retinacular pattern as those species that are. Thus, the retinacular evidence may indicate that Galago
evolved from a species that utilized vertical
supports often.

Ligaments
The ligaments in the galago and tarsier
foot are the same with the exception of the
distal navicular-mesocuneiform ligament
(Gebo, 1986).Tarsiers are the only prosimian
to possess this ligament. This ligament is
responsible for keeping the distal navicular
associated with the mesocuneiform, since
these bones are not in articulation in tarsiers
(Fig. 4).
Galago and Tarsier osteological comparisons
Talus: Galagines have flat talar heads,
which look squared off from a dorsal view,
while tarsiids have a triangular-shaped talar
head, which is very unusual when compared

Fig. 5. Tarsier joint anatomy. a: Distal view of talar

head shape (arrow points to medial side). b: Anterior
plantar talar facet (arrow points toward the bony ridge
which separates this facet into two tiers): c. Distal view
of the flat calcaneal-cuboid joint surface of the calcaneus.
d Dorsal view of the cuboid illustrating the unusual
cuboid shape in tarsiers and the lack of a proximal cuboid process. e: Shape of the navicular-cuneiform joint
surface (arrow points to ball-like entocuneiform facet on
the navicular). E Plantar view of the distal navicular
facet outline illustrating the reduced mesocuneiform
facet (arrow). Compare a-f with Figure 6.

Fig. 6. Galagine joint anatomy. a: Distal view of talar

head shape (arrow points t o lateral side). b: Continuous
anterior plantar talar facet. c . Distal view of calcanealcuboid joint surface for the calcaneus (arrow points to
calcaneal pit or pivot area). d. Dorsal view of the cuboid
(arrow points to cuboid process). e. Shape of the navicular-cuneiform joint surface. f. Plantar view of the distal
navicular facet outline. Note the prominent mesocuneiform facet (arrow). Compare with Figure 5.

19

TARSIER FOOT ANATOMY

to other prosimians (Figs. 5, 6). Seen from

above, the tarsier talar head comes to a
rounded point on the dorsal medial side. Tarsiers have a talar head which is more developed on the medial side, and this contrasts
dramatically with other prosimians. This opposite side of development is in response to
the horizontal position of the talus in the
tarsier foot (Jouffroy et al., 1984) and navicular rotation at the talonavicular joint.
A midline through the talar head forms an
angle to the talar body. Talar heads tilt in a
medial-lateral direction in prosimians with
galagines elevating the medial side, while
tarsiids have a horizontal facing talar head
with neither side elevated.
The talar neck is long in galagines and
short in tarsiids (Table 6). I: spectrum has a
longer relative talar neck length compared
to I: bancanus or I: syrichta. Tarsiers possess
the relatively shortest talar necks of all prosimians (Gebo, 1986). Galagines and tarsiids
possess fairly straight talar necks with little
medial deviation.
On the plantar side of the talar neck is the
anterior facet for the calcaneus and the
spring ligament. This facet is roughly divided into a lateral half, which articulates
with the calcaneal facet, and a medial half,
which rests on the spring ligament. Small
ridges separate the two halves. The anterior
plantar talar facet extends well proximal toward the posterior facet but stops at the
groove for the interosseous ligaments. In galagines, the anterior plantar talar facet extends to and blends into the posterior talar
facet on the lateral side and the interosseous
groove is a circular pit. Tarsiers possess an
unusual anterior plantar talar facet where
the proximal and distal halves are separated
by a ridge (Fig. 5).The distal half of this facet
is raised above the proximal half. This twotiered talar facet likely functions for talar
stability while the navicular rotates over the
talar head during inversion in tarsiers. Since
the tarsier talus moves very little during inversion, there is no need to have a smooth
joint surface between the talus and the anterior calcaneal facet.
The tibia and the fibula rest upon the trochlea of the talar body. The medial malleolus
of the tibia and the lateral malleolus of the
fibula form a two-sided prong which fits over
the medial and lateral sides of the talar body.
Together, the tibia, the fibula, and the talus
form the talocrural joint, which is mainly
responsible for dorsi and plantarflexion. The
curvature of the medial talar rim results in

TABLE 6. Relative lengths of the talar neck (TNL),

cuboid (CL) and navicular (NL) in galagines and
tarsiids

TNL
MTW

Galago alleni
Galago crassicaudatus
Galago demidouii
Galago garnettii
Galago senegalensis
Galag-oelegantulus
Tarsius bancanus
Tarsius syrichta
Tarsius spectrum

CL
MTW

NL
MTW

1.68
1.86
1.77

3.88
7.56
4.12
6.71
4.39

1.22
1.14
1.15
1.12
1.18

2.19

1.09

1.95

0.79
0.84

1.11

5.70

1.01

All length values are divided by midtalar trochlear width (MTW)

for comparisons across taxa.

some abduction during dorsiflexion as well

as adduction during plantarflexion (Hafferl,
1932; Barnett and Napier, 1952; Lewis,
1980a). Tarsiers have a very wedge-shaped
trochlea compared to the more parallel trochlear rims of galagines. The talocrural joint
is not as tight a joint in the medial-lateral
direction as it is in the anteroposterior direction. This looseness allows the foot as a
whole to be abducted and adducted at the
talocrural joint relative t o a stable tibia-fibula position. The medial side of the talus,
which articulates with the tibia, is extensive
and extends to the plantar edge for both galagines and tarsiids. The lateral talar side,
which articulates with the fibula, extends
obliquely outward from the talar body in prosimians except for Tarsius, where the lateral
talar side is vertically steep. In galagines the
lateral talar side is more vertical than other
prosimians but different than tarsiers in
appearance.
In tarsiids and galagines, the talar body is
high in the dorsoplantar plane. Functionally,
talar body height may relate to leaping animals possessing a more tightly curved talar
body. A higher and tighter curvature provides a quicker pivot at the talocrural joint
for leapers (Hall-Craggs, 1965).
The position of the flexor hallucis longus
groove is on the lateral side of the posterior
trochlear facet of the talus for all prosimians
except tarsiers, which have this groove in the
middle below the posterior trochlear facet.
These two different flexor positions reflect
the two spatial positions the talus occupies
in galagines and tarsiids, where the tarsier
talus is positioned in a more horizontal set.
Navicular: The position of the navicular in
a prosimian foot is an oblique one where the
entocuneiform facet of the navicular rests on

20

D.L. GEBO

Fig. 7. Photograph of the navicular-cuneiform joint

surface in the same anatomical position for Tarsius (A)
and Galago (B). a = entocuneiform facet; b = mesocuneiform facet; and c = ectocuneiform facet. Note the

especially different mesocuneiform facets, the plantar

(bottom) and lateral (left) shapes of the distal navicular
joint surface, and the size of the entocuneiform facet
relative to the total joint surface.

the plantar side of the foot and the navicularectocuneiform facet lies dorsally, adjacent to
the cuboid. The navicular angles laterally
away from the first digit. The shape of the
proximal navicular mirrors the shape of the
talar head. Tarsiers possess a sharp medial
bend at the medial side of this rounded surface, reflecting their triangular-shaped talar

head. The proximal extension of the navicular tuberosity is longer in tarsiers and galagos than in other prosimians.
The distal surface of the navicular is one of
the most distinctive regions of the foot in
identifying prosimian families. The distal
navicular is composed of three joint surfaces,
one for each of the three cuneiforms. All three

TARSIER FOOT ANATOMY

facets are stacked upon each other. The facet

for the entocuneiform is either round or oval
in shape and touches the mesocuneiform
facet dorsally in most prosimians. The mesocuneiform facet comes in many shapes from
squared to rounded and borders both the entocuneiform and ectocuneiform facets. The
ectocuneiform facet is very variable in shape
and is the most dorsal and lateral of all of
the distal navicular facets. The distal navicular does not permit much joint mobility,
especially rotational movements. The reason
the navicular-cuneiform joints do not permit
very much mobility is due to the confluent
movements of the navicular and the distal
part of the foot during inversion and eversion. At the navicular-cuneiform joints, the
ectocuneiform protrudes well forward proximally above the level of the other cuneiform
facets in galagines and prevents a very confluent joint surface for mobility. Tarsiers are
an exception to this. Tarsiids have the most
mobile navicular-cuneiformjoint of all prosimians and have moved the mesocuneiform into
a position which does not normally articulate
with the navicular (Fig. 5). This allows the
navicular to rotate in the cup-shaped entocuneiform facet and the curved surface of the
ectocuneiform facet of the navicular with far
greater freedom. Figure 7 pictures the distal navicular facet shapes for Galago and

Tarsius.
Galagines have probably the most distinctive distal navicular of all strepsirhines. The
distal navicular is composed of three prongs,
one for each cuneiform facet (Fig. 6). There
exists a distinct gap between the entocuneiform facet and the mesocuneiform facet (Hafferl, 1932).In the smaller galagines, this gap
is more like a groove which separates the
entocuneiform and mesocuneiform facets.
The entocuneiform facet extends prominently away (distally) from the ectocuneiform facet. The mesocuneiform facet
protrudes distally and is equal in distance
with the distal projection of the entocuneiform facet. The ectocuneiform facet has a
very dished surface and is similar to that of
tarsiers. This surface is more closed, however, than in tarsiers and both differ in comparison between a very open and a more
closed c in shape. The ectocuneiform facet
has also markedly shifted laterally in galagines. The distal navicular three-prong configuration in galagines prohibits rotational
mobility. Only dorsiflexion has more than a
few degrees of mobility at this joint (Gebo,
1986).

21

Tarsiers have a very distinctive distal navicular which is related to their unusual
manner of foot rotation. The entocuneiform
facet on the navicular is enlarged and balIlike in tarsiers (Hafferl, 1932).The ectocuneiform facet is very large and has a very dishedout surface. The mesocuneiform is very small
in tarsiers and does not articulate with the
mesocuneiform except during extreme plantarflexion. A ligament connects the midnavicular area to the mesocuneiform at the
navicular-mesocuneiform joint (Fig. 4). The
distal navicular of tarsiers is constructed for
greater mobility than it is in other prosimians.
Calcaneus: The normal foot position of a
prosimian places the cuboid in a dorsiflexed
position relative t o the calcaneus. Because of
this, the anterior calcaneus has been bent
dorsally to accommodate the articulation of
the anterior calcaneus with the cuboid. This
position elevates the heel of the calcaneus off
the substrate. I call this anterior calcaneal
bend dorsal tilting, since the anterior calcaneal bend places the heel dorsal to the
substrate surface. Dorsal tilting is evident in
all prosimian calcanei, except Tarsius.
The calcaneal-cuboidjoint, which makes up
the transverse tarsal joint along with the
talonavicular joint, is probably the most significant joint in the prosimian foot. The shape
of the cuboid facet on the anterior calcaneus
allows prosimians to invert their feet in an
extreme fashion compared to other mammals. Several authors have pointed out
clearly the derived nature of this joint for
euprimates (Hall-Craggs, 1966; Decker and
Szalay, 1974; Lewis, 1980b). Galagines possess a very dorsoplantar high cuboid facet. In
galagines, the deep calcaneal pit helps galagos to overcome the extreme calcaneal and
navicular elongation which would normally
limit mobility at this joint (Fig. 6). Galagines
also possess a prominent dorsal lateral bump
on this facet for the dorsal lateral calcanealcuboid ligament. Tarsiers possess the most
oval cuboid facet of all prosimians. The tarsier cuboid joint is also very flat with no
central pit (Hafferl, 1932; Hall-Craggs, 1966)
(Fig. 5). The dorsal lateral direction of this
facet is much longer in tarsiers than in other
prosimians. The flatness and lack of a central
pit limit rotational mobility at this joint compared to other prosimians. Joint-mobility
measures for inversion by Tarsius indicate a
lower mobility than other prosimians but still
a highly mobile joint (Gebo, 1986). Tarsier
rotational mobility is due to the unique mor-

22

D.L. GEBO

phology of the distal navicular, not the calcaneal-cuboid joint. Tarsiers are the only
prosimians which have never been observed
to hang by their two hind feet where extreme
calcaneal-cuboid rotation is required. This is
likely due to their extremely elongated calcaneus and navicular; the shape of the calcaneal-cuboid joint, where the absence of a
cuboid process cannot be locked into a nonexistent calcaneal pit; and their unique
method of rotation at the navicular-cuneiform joints.
Tarsiers have the longest anterior calcaneus of any prosimian, followed in turn by
galagos (Gebo, 1986).This anterior calcaneal
length has led to modifications at the distal
calcaneus for rotation. Galagines and tarsiids do not possess a navicular facet on the
proximal calcaneus. In galagines, a synovial
joint between the calcaneus and the navicular prevents a proximal calcaneal-navicular
articulation. Hall-Craggs (1966) has explained how this synovial joint is unique to
galagos and how it increases the strength of
the tarsus, but does not reduce mobility.
The posterior calcaneal facet is medially
convex in shape, and this shape is responsible for the helical movement of the talus
during inversion and eversion (Manter, 1941;
Decker and Szalay, 1974; Lewis, 1980b). The
posterior calcaneal facet lies in the anteriorposterior plane of the calcaneal midline and
faces medially. This is true for both galagines and tarsiids.
Both galagines and tarsiids possess a
straight plantar calcaneal edge rather than
being curved in the middle. Tarsiers bow the
plantar edge of the posterior calcaneal region
dorsally and this is strikingly different than
the straight calcaneus of galagos.
Cuboid The proximal cuboid surface in
prosimians is more or less oblong in shape
and has a prominent process which fits into
the calcaneal pit. In galagines the cuboid
process is prominent and peglike (Fig. 6).
Tarsiids have a very flat proximal cuboid
surface which does not contain a cuboid process (Fig. 5). Szalay (1976)offers two possible
explanations for the lack of a cuboid process.
One reason is that the cuboid process inhibits
extensive dorsal-plantar movements in the
distal part of the foot for leaping, and therefore a reduced cuboid pivot would serve better. Unfortunately, galago anatomy and foot
positions argue against this explanation,
since galagos possess a very prominent cuboid process and galagines also have exten-

sive dorsal-plantar movements in the distal

part of the foot. The second explanation links
the use of vertical supports with less varied
foot orientations; and thus, there is no longer
any need for a prominent cuboid process. Szalay assumes in both explanations that medial-lateral rotations have diminished for
tarsiers or that the distal part of the tarsier
foot is normally more dorsiflexed than a galago foot on a vertical support. The reason
tarsiers have a reduced cuboid process is due
to a whole series of joint changes which allow
tarsiers to rotate their feet in a very novel
way.
The relative length of the cuboid is very
short for tarsiers while it is long in galagines. The short cuboid in tarsiers allows the
calcaneal-cuboidjoint to lie more in line with
the navicular-cuneiform joints (Jouffroy et
al., 1984).This facilitates rotation at the calcaneal-cuboid joint and the navicular-cuneiform joints. Since galagines utilize the
standard method of rotation at the transverse tarsal joint, navicular and cuboid
lengths need to be more similar (Table 6).
Galagines have two cuboid facets for the
ectocuneiform, while tarsiers possess only
one. The shape and position of this facet in
tarsiers is unusual in that it is rectangular
and high in a dorsoplantar direction. This
facet is small; and its size, shape, and orientation are likely responses to the unique rotational movements of the navicular in
tarsiers, since the tarsier cuboid does not
screw into the nonexistent calcaneal pit.
Since the tarsier cuboid does not shift proximally toward the calcaneus, the cuboid does
not displace itself along the ectocuneiform facet as it does in other prosimians.
The cuboid-ectocuneiform facets are therefore shaped in a much different manner
in tarsiers.
In addition to the ectocuneiform and the
navicular facets, the cuboid articulates with
or at least abuts against the mesocuneiform
along its medial plantar border for all prosimians but tarsiers. In tarsiers, the mesocuneiform does not touch the cuboid.
The cuboid fits into a triangular notch on
the distal lateral side of the navicular in
tarsiers. There is extensive contact between
the cuboid and the navicular since the proximal cuboid curves towards the navicular. The
navicular-cuboid facet lies distal to the proximal cuboid surface and behind the extended
medial lip of the proximal cuboid facet. This
facet is Gery concave. In galagines, the cu-

TARSIER FOOT ANATOMY

23

Fig. 8. Second metatarsal notch arrangement for Galago (A) and Tarsius (B). 2 = second
metatarsal; M = mesocuneiform; E = ectocuneiform; S = calcaneal-navicular synovial joint.
Note the ectocuneiform joint surface for the second metatarsal in Tarsius (B).

boid-navicular facet is flat and faces plantad.

Cuneiforms: The three cuneiforms-the entocuneiform, the mesocuneiform, and the ectocuneiform-perform three key functions.
One, the three cuneiforms form a tightly
bound stable arch in the foot. The navicular
and the cuboid articulate with and move relative to this cuneiform unit. Two, the three
cuneiforms make up most of the plantar
transverse arch so the flexor tendons can
reach the toes. Three, the cuneiforms hold
the second metatarsal in a recessed notch of
bone. Tarsiers are noteworthy for this last
function in that they possess a unique wedgeshaped notch. This wedge-shapednotch is due
to the expanded articulation of the now
base of the second metatarsal with the ectocuneiform, rather than the mesocuneiform
(Fig. 8). The tarsier mesocuneiform articulates along the medial side of the second metatarsal with the entocuneiform.
Metatarsals and digits: The peroneal tubercle for the first metatarsal is very large
in galagines and more slender in tarsiids.
Likewise, the peroneus longus muscle, an
important muscle for adducting the hallux
during grasping, is large in galagines and
much smaller for tarsiids.
GaIagines have shorter metatarsals than
tarsiers (Schultz, 19631, and the tubercle on
the fifth metatarsal is smaller in galagines
than it is in tarsiids.

The proximal and distal ends of prosimian

metatarsals are normally dorsoplantarly
high as they are in galagines, but tarsiers
possess broader distal metatarsal ends (Hafferl, 1932).
Tarsiers are different from galagines in two
characteristics in the digits. First, tarsiers
possess a toilet claw on the third digit as well
as the second digit, and both are true claws
in tarsiers (Clark, 1936).Second, tarsiers possess very large digital pads which help to
increase toe friction when grasping supports.

Osteological summary
Although galagines and tarsiids share several osteological features for leaping (e.g.,
high talar body height, straight talar neck, a
straight plantar calcaneal edge, and an elongated calcaneus and navicular), they differ
dramatically in the shape of their joints
which function to rotate the foot into inversion. Table 7 lists the unique features of galagines and tarsiids. Most of these features in
Table 7 are involved in joint rotations for
inversion and eversion, but tarsiids and galagines do differ in other features which are
not related to joint rotation. These features
are the position of the flexor hallucis groove
on the talus, the shape of the lateral talar
facet, the shape of the second metatarsal
notch, the size of the tubercles on the first
and fifth metatarsals, distal phalangeal

24

D.L. GEBO

TABLE 7. Galwine and tarsiid osteological characteristics

Galagines
Unique features
1. Calcaneal-navicular synovial joint
2. Shape of talar head is squared (dorsal view)
3. Shape of calcaneal-cuboidjoint with dorsal lateral tubercle (Fig.6)
4. Anterior plantar talar facet extends to posterior facet
5 . Three-prong distal navicular (Fig. 7)
6. Round distal metatarsal ends (side view)
Shared features with tarsiids
1. Elongate calcaneus and navicular
2. Straight talar neck
3. Dorsoplantar high talar body
4. Lack of a navicular facet on the calcaneus (proximal calcaneus)
5 . Straight plantar calcaneal edge
Tarsiids
Unique features
1. Medial development of the talar head
2. Triangular shaped talar head
3. Two-tiered anterior plantar talar facet
4. Horizontally facing talar head relative to talar body
5 . Tiny lateral talofibular process on the talus
6 . Flexor groove is in the midline of the posterior talar trochlea
7. Short talar neck
8. Distal navicular shape and mesocuneiform ligament (Figs. 4 and
7)
9. Medial bend in proximal navicular
10. Flat calcaneal-cuboid joint
11. Dorsal bowing of the posterior calcaneal plantar edge
12. Short cuboid
13. Shape of the cuboid (Fig. 5 )
14. Lack of a cuboid process on the proximal cuboid surface
15. Wedge-shaped second metatarsal notch (Fig. 8)
16. Large fifth metatarsal tubercle
17. Angle of the distal phalangeal facet to the phalangeal shaft
18. Toilet claw on digit three

Fig. 9. Xeroradiograph of an inverted foot position for

G. crassicaudatus (medial view). Arrows and letters point
to the following bone positions and observable bony charaeteristics: a = rotated position of the calcaneus showing
a plantar view of the calcaneus; b = dorsal position of

the proximal navicular process relative to the talar head;

c = rotated position of the entocuneiform facet on the
distal navicular; d = rotated cuboid position which shows
the plantar groove for the peroneus longus tendon; and
e = position of the calcaneal-navicular synovial joint.

TARSIER FOOT ANATOMY

shape, posterior calcaneal bowing, and the

toilet claw on the third digit. The important
point to remember from Table 7 is that just
about every joint in the tarsier foot has been
remodeled; the following section will explain
how these remodeled joints function in the
tarsier foot.

Joint function
Inversion for galagines and all other prosimians: As the invertor musculature (anterior tibialis, posterior tibialis, flexor hallucis
longus, and flexor digitorum longus) contracts, it exerts a pull on the medial part of
the plantar aspect of the foot. This pull rotates the navicular in a lateral direction, with
the result that the navicular comes to lie on
top of (dorsal to) the cuboid rather than on its
medial aspect (Fig. 9). This is the beginning
movement of inversion contra Decker and
Szalay (1974) and Lewis (1980b).Meanwhile,
the long flexor tendons are at the same time
pulling the distal part of the foot toward the
talus and the calcaneus at the transverse
tarsal joint. This movement forces the cuboid
process to screw into the calcaneal pit at the
calcaneal-cuboidjoint. As the cuboid screws
into the calcaneus, the cuboid is moving
proximally and slides closer to the talonavicular joint, which places the calcaneal-cuboid
joint more in line with the talonavicular joint
for greater ease of rotation. The proximal
displacement of the cuboid allows the cuboid
to also slide along the cuboid-navicular and
the cuboid-ectocuneiform facets in a proximal direction. After the cuboid has moved
proximally and has been screwed into the
calcaneus, the ectocuneiform protrudes distally away from the cuboid (Fig. 9).
The long flexor tendons, which run medially through and behind the posterior talar
tubercles and alongside of and below the sustentaculum tali, upon contraction, rotate the
calcaneus laterally (Fig. 9). The short calcaneal-cuboid plantar ligament is also responsible for rotating the calcaneus into an
inverted position, since this ligament keeps
the calcaneal-cuboid joint in close association. In other words, rotating the cuboid during inversion necessitates some calcaneal
rotation as well. The long flexors and the
short calcaneal-cuboid plantar ligament are
essential for calcaneal inversion, since no
muscles attach to the calcaneus to bring
about this rotation. In inversion, both the
calcaneus and the cuboid are rotated laterally, with the cuboid rotating laterally out-

25

ward and below the calcaneus in a much

greater degree. So much so that the peroneus
longus groove on the plantar side of the cuboid is in clear view after rotation (Fig. 9).
As the calcaneus rotates laterally, the talus
shifts backward (proximal) along the posterior calcaneal facet. The talar neck is lifted
above and medially away from the sustentaculum tali and toward the spring ligament.
This tightens ligament cervicis. The posterior part of the anterior plantar talar facet
locks with the posterior part of the anterior
calcaneal facet. The talar head shifts medially and dorsally to correspond with the lateral direction of movement by the navicular.
The talar head and neck point ahead in a
proximal-distal line of direction during inversion with the navicular above the rim of
the talar head. (Fig. 9).
Manter (1941) discussed the screw-like
character of the subtalar joint in Homo where
he noted the medial curvature and the proximal-distal orientation of the posterior calcaneal facet. The arc of the posterior calcaneal
facet allows the talus to roll forward or backward along the calcaneus in a helical motion.
Manter (1941) noted that the subtalar and
transverse tarsal joints are interdependent
when rotated during inversion and eversion.
The navicular-cuneiform and the intercuneiform joints are not very mobile and are
essentially unimportant for inversion or eversion foot rotations. The navicular-cuneiform joints permit some plantar and
dorsiflexion and a little medial-lateral twisting but are in general restrictive in their
joint mobility, with the exception of dorsiflexion, which has the greatest freedom of movement of the possible directions of movement.
The intercuneiform joints for the entocuneiform, mesocuneiform, and ectocuneiform
are firmly held in place by their ligaments.
Together these three cuneiforms hold the second metatarsal in a notch between them, and
all form the transverse arch plantarly. Very
tiny sliding movements are possible between
the intercuneiform joints during inversion
but in general the cuneiforms are firmly held
in place and are essentially nonmobile joints.
Navicular rotation in tarsiers: The extremely elongated calcaneus and navicular
in tarsiers, an adaptation for leaping, has led
to modifications in the foot to maintain the
inverted position of the foot when grasping a
support. The calcaneal-cuboid joint, where
most rotational ability comes from in the foot
of prosimians, is very flat in tarsiers and
therefore far less mobile. The tarsier talar

26

D.L. GEBO

foot. Note the following: the position of the proximal

navicular process (a); the position of the entocuneiform
facet for the navicular 03); and the calcaneal-navicular

spatial separation (c). Features a and b are in the same

position in Galugo (A) and Tarsius (B), but feature c is
much larger in Tarsius (B). Compare this figure with
Figure 11to note bone position changes after rotation of
the foot into an inverted foot position.

head is pointed, triangular in appearance

rather than round, and is more developed on
the medial side, rather than the lateral side
as in other prosimians. The anterior plantar
talar facet is not continuous but two-tiered,
with the distal part elevated above the proximal part (Fig. 5). The talar head shape and
the anterior plantar facet show morphologies
that reflect limited mobility as well. Thus,
the transverse tarsal joint in tarsiers is not
as mobile as in other prosimians.
Like tarsiers, galagos have a very elongated calcaneus and navicular, but the galagine transverse tarsal joint is similar to that
of other prosimians. Hall-Craggs (1966) has
described how galagos rotate the navicular
around the calcaneus, which acts as a stable
axis. The navicular is held firmly in place at
the navicular-cuneiformjoints, and strength
is added by the synovial joint between the
calcaneus and the navicular (Figs. 10,111.

Unlike galagos, tarsiers rotate the navicular at both the proximal and distal ends,
instead of the calcaneus and proximal navicular, to achieve an inverted foot position.
The calcaneus in tarsiers rotates only
slightly. In galagos, the synovialjoints keeps
the calcaneus and the navicular closely associated, but in tarsiers there exists a large
and evenly spaced gap between the calcaneus and the navicular (Fig. 10). This gap
allows ample space for navicular rotation.
Distally, the tarsier navicular allows for
far greater rotational movements than in
galagos. The distal navicular articulates with
all three cuneiforms in galagos, but in tarsiers, the distal navicular does not articulate
with the mesocuneiform except during plantarflexion, which prevents dislocation of the
navicular-cuneiform joints. Tarsiers possess
a small ligament which bridges this spatial
gap and connects the navicular and the me-

Fig. 10. Xeroradiographs of the foot of G. senegalensis

(A) and 71 syrichtu (B). Dorsal view of a plantarflexed

TARSIER FOOT ANATOMY

27

Fig. 11. Xeroradiographs of the foot of G. senegalensis

Notice in Tarsius (B) that the navicular has rotated

position. Note the position of the proximal navicular

process (a), the position of the entocuneiform facet of the
navicular (b), the position of the plantar calcaneal edge
(c) in A and the medial calcaneal side in B, and the loss
of the calcaneal-navicular spatial gap in B but not in A.

(A). The distal entocuneiform facet (b) of the navicular

in Tarsius (B) is not visible from this medial view, since
it lies in a more lateral position. Likewise, the proximal
navicular process (a) also lies in a more lateral position
compared to A.

socuneiform. The distal navicular in tarsiers

has developed the entocuneiform facet into a
very round ball which fits into the cup-shaped
entocuneiform facet as a ball-and-socket type
of joint for greater rotation. The navicularectocuneiform joint normally locks up in prosimians, especially galagos, so that this joint
does not permit very much motion except for
dorsiflexion. In tarsiers, the navicular-ectocuneiform joint allows a far greater range of
rotational movement (Fig. 11).If the distal
navicular in tarsiers has been modified for
rotational movements, the proximal navicular must move as well (Fig. 11).There is no
calcaneal-navicular synovial joint in tarsiers
to interfere with this rotation as there would

be in galagos if galagos utilized navicular

rotation. Since the calcaneus does not move
very much, the talus must also remain fairly
stable in its position relative to the navicular
(Fig. 11).The tarsier talus sits in a much
more horizontal position (Jouffroy et al., 1984)
compared to other prosimians, and this also
facilitates navicular rotation.
Manipulation of galago and tarsier cadavers confirms this calcaneal movement difference between the two genera. As the anterior
tibialis contracts for inversion, the entocuneiform and the distal foot rotate laterally.
The cuboid rotates (slips) laterally but since
there is no cuboid peg in tarsiers, the cuboid
does not insert into the calcaneal pit as in

(A)and 7: syrichta (B). Medial view of a n inverted foot proximally and distally farther than it has in Galago

28

D.L. GEBO

other prosimians (Figs. 10, 11). The cuboid

and ectocuneiform maintain their articulation since the cuboid does not screw into the
calcaneus. The tarsier navicular rotates with
the distal part of the foot and the cuboid until
the distal part of the navicular locks on the
calcaneus and the cuboid. This rotation is
greater than 90" (Fig. 11). The shape of the
proximal cuboid, which curves medially, and
the position of the navicular-cuboid facet,
which lies behind (proximal)this curvature,
are important components in allowing this
extreme rotation by the navicular. As the
navicular rotates past the normal 90"
amount of movement, the proximal cuboid,
which normally fits securely into the excavated notch of the lateral navicular, slides
along the navicular-cuboid facet until only
the most medial point of the medially curving proximal cuboid surface articulates with
the navicular (Fig. 11). This sliding by the
cuboid allows the navicular to rotate farther
than would ordinarily be possible.
As the navicular rotates laterally, the talar
head remains fairly fixed in place and the
navicular rotates across the medial side of
the talar head to the lateral side. This movement explains why the medial side of the
tarsier talar head is more developed and
rounded dorsally than the talar heads of
other prosimian species.
Tarsier navicular rotation is an adaptation
for small-bodied specialized leapers which
must also invert their feet for grasping
curved surfaces. Extreme elongation of the
calcaneus and navicular places a mechanical
strain on these distal elements especially if
they are to maintain their rotational ability.
In galagos, a synovial joint evolved to
strengthen the elongated calcaneus and navicular (Hall-Craggs, 1966). A synovial joint
allows galagos to have the structural support
which is needed, and at the same time, have
the flexibility to invert their feet for grasping. Galagos have also evolved a calcanealcuboid joint with a deep calcaneal pit and its
corresponding peglike cuboid process. This
enhanced calcaneal-cuboidjoint and the calcaneal-navicular synovial joint allows galagos to invert their feet adequately with such
elongated tarsals. Tarsiers, on the other
hand, solved these same mechanical problems in a different manner by changing the
critical joints involved in rotation movements. Thus, navicular rotation in tarsiers is
functionally equivalent to the synovial joint
and calcaneal-cuboidjoint of galagos.

The leaping adaptations in the tarsier foot

have led to modifications for inversion in tarsiers. The proximal and distal ends of the
navicular rather than the calcaneal-cuboid
joint account for most of the rotational ability in the tarsier foot. Jouffroy et al. (1984)
had some inkling of this unusual distal navicular region in tarsiers when they described this part of the foot as a transverse
side to side joint, based on the shortness of
the cuboid and the same transverse level of
the distal navicular and calcaneus. Unfortunately, their analysis did not focus on the
critical joint morphology, and their data led
them to infer slight or lack of rotational
movement at the distal navicular. The tarsier talar head and neck, the calcaneal-cuboid and the navicular-cuboid joints, and
especially the navicular have all been
uniquely modified to maintain the critical
inverted foot posture which is crucial for primate grasping feet. Tarsier foot rotation and
the anatomy associated with it evolved only
once in the order and is a truly unusual primate phenomenon.

Omomyid-tarsiid comparisons
When tarsier foot anatomy is compared
with the known omomyid foot bones, it is
clear that omomyids are not very similar to
modern-day Tarsius (Table 8). Calcaneal
length is more cheirogaleid-like for the omomyids (Szalay, 1976)than the extreme length
of Tarsius with the exception of Necrolemur.
Yet, the talus of Necrolemur is unlike that of
Tarsius (Godinot and Dagosto, 1984). The
talar plantar facet is not two-tiered as it is in
tarsiers, and the talar head is oval in Necre
lemur rather than triangular in appearance
as it is in tarsiers. The oval talar head and
the continuous talar plantar facet are features found in Teilhardina, Tetonius, Washakius, Arapahovius, and Hemiacodon as well.
A cuboid process on the proximal cuboid is
known for Hemiacodon, Tetonius, and Arapahovius (Simpson, 1940;Savage and Waters,
19781, while a calcaneal pit or pivot on the
distal calcaneus has been documented in
Teilhardina, Tetonius, Washakius, and Hemiacodon (Szalay, 1976). Thus, calcaneal-cuboid rotation during inversion for the known
omomyids is like that of most other primates
and unlike Tarsius.
The known distal navicular joint surface of
Hemiacodon and Arapahovius shows the mesocuneiform facet to articulate with the mesocuneiform, and the entocuneiform facet is

See Figure 5

1. Anterior calcaneal elongation

2. Navicular length
3. Cuboid length
4. Talar neck length
5. Talar head shape
6. Talar head-side of development
7. Anterior talar plantar facet
8. Trochlear shape
9. Trochlear depth
10. Talar body height
11. Flexor groove position-talus
12. Proximal navicular shape
13. Distal navicular shape
14. Mesocuneiform articulates winavicular
15. Distal navicular groove for calcaneus
16. Cuboid facet position on navicular
17. Cuboid-ectocuneiform facets
18. Shape of distal cuboid
19. Calcaneal-cuboidjoint
20. Entocuneiform facet for first
metatarsal
21. First metatarsal tubercle
22. Distal metatarsal shape
23. Distal phalangeal angle
24. Second metatarsal notch
25. Large fifth metatarsal tubercle
26. Posterior calcaneal facet height
relative to dorsal heel height
27. Posterior calcaneal facet
28. Posterior calcaneal bowing
29. Peroneal tubercle placement
30. Anterior calcaneal facet
31. Plantar tubercle calcaneus
32. Cuboid shape
33. Anterior calcaneal dorsal tilting
34. Calcaneal-navicular facet
35. Posterior trochlear shelf
36. Medial trochlear curvature
37. Talofibular facet

Equal
Tightly curved
No
Mid-calcaneus
Curved
Moderate
Rectangular
Yes
Yes
Slight
No
Steep

Large
Round
Oblique

Long
Long
Moderate
Long
Oval
Medialilateral
Continuous
Parallel
Moderate
Moderate to high
Midline
Oval
Normal pattern
Yes
No
Below ectocuneiform
2
Square
Pivot type
Stops at lateral edge

Omomyids

TABLE 8. Omomyid characteristics compared to Tarsius

Steep

No

Absent
Absent

No

Moderately curved
Yes
Same
Straight
Small
Odd'

Broad
Very angled
Ectocuneiform base
Yes
Equal

L-shaped
Flat
Continuous
Moderate

Yes
Same

No

Very long
Very long
Short
Short
Triangular
Medial
Two-tiered
Wedged
Deep
High
Midline
Medial bend
Unique'

Tarsius

30

D.L. GEBO

not ball-like in appearance. Both features are

unlike Tarsius and similar to other extant
prosimians. Therefore, navicular rotation did
not occur in these omomyids where the relevant foot evidence is known.
The first metatarsal of Hemiacodon has a
very robust peroneal tubercle (Simpson,
1940),which is similar to that of galagines
but not that of tarsiids. Herniacodon undoubtedly possessed a large peroneus longus
muscle which adducted the hallux forcefully
during a grasp. Peroneus longus emphasis is
a different method of muscle development for
grasping a support than the method tarsiers
utilize.
To sum up the known omomyid foot evidence is to show that the calcaneal-cuboid
joint, the subtalar joint, the navicular-cuneiform joint, and the first metatarsal peroneal
tubercle in omomyids lack any morphological similarity with tarsiers. Thus, the species
of Tarsiidae are much more specialized in
their foot anatomy than were their ancestors,
the Omomyidae, which are more similar to
other extant prosimians in terms of joint
function and grasping supports.
CONCLUSIONS

Galagos can be separated into two main

movement gradients, one for leaping and the
other for quadrupedalism. G. senegalensis
and G. alleni are specialized leapers, while
the smallest galago, G. demidouii, and the
largest, G. crassicaudatus, emphasize more
quadrupedal tendencies. Both quadrupedalism and leaping are common for G. elegantulus, which has clawlike nails and utilizes
large-diameter tree trunks. Galagos utilize
many more types of movements than do tarsiers. Galagos possess the I-V grasp, which
concentrates intrinsic musculature along the
first and fifth digits. The extrinsic leaping
musculature of galagos is slightly better developed than their flexor musculature. The
deep flexor musculature is better developed
in galagines than tarsiers since galagines
utilize many more types of movements. The
galago peroneus longus is well developed for
adducting the hallux during a grasp. Galagos possess an unusual extensor retinacular
loop compared t o other prosimians.
Galago foot osteology represents a foot
adapted for leaping. This includes the galago
species which often move by quadrupedalism. Galagos have evolved a calcaneal-navicular synovial joint t o strengthen their very
elongated calcaneus and navicular. The dis-

tal navicular is unusual in that the three

distal facets are in a three-prong arrangement and there is a large gap between the
entocuneiform facet and the other two distal
facets. The three-prong navicular arrangement limits rotational movements. Galagines, like other prosimians, utilize a specific
calcaneal-cuboid,talonavicular, and subtalar
joint anatomy to invert their feet.
Tarsier extrinsic foot musculature is well
developed for leaping with twice the amount
of superficial flexors to deep flexor musculature. Tarsiers have a small peroneus longus
muscle, and the deep flexor musculature is
less developed for grasping than in other prosimians. The method tarsiers use to hold on
t o vertical supports differs from other prosimians. Tarsiers possess more intrinsic foot
musculature relative to extrinsic foot musculature and a unique contrahentes arrangement that allows them to arch the middle
and proximal phalanges to put more pressure on the large digital pads for grasping.
Thus, tarsiers rely more on their intrinsic
flexors than the deep extrinsic flexor musculature or the peroneus longus.
Tarsiers, like indriids, have extra extensor
retinacular bands. Only tarsiers possess a
ligament between the distal navicular and
the mesocuneiform, since these two bones do
not normally articulate. Tarsier foot osteology is very unusual among prosimians and
relates t o their specialized leaping and their
unique method of foot rotation. Unlike other
prosimians, tarsiers are able to rotate the
navicular at both the proximal and distal
ends rather than rotating the proximal navicular and calcaneus to achieve foot inversion. The talar head and neck, calcanealcuboid joint, and especially the navicularcuneiform joint contribute to this novel type
of rotation. This novel method of foot rotation, coupled with the specializations for
leaping, has dramatically remodeled the
bones in the tarsier foot, and thus Tarsius is
very different anatomically from its ancestral lineage, the Omomyidae.
ACKNOWLEDGMENTS

I would like to thank E.L. Simons, P. Chatrath, and P. Wright at the Duke Primate
Center for permission, access, and help
throughout this study. A special thanks for
the many helpful discussions must go to T.
Rasmussen and M. Dagosto. I would also like
to thank K. Glander, W. Hylander, R. Kay,
and R. MacPhee for writing improvements.

TARSIER FOOT ANATOMY

31

M. McKenna and G. Musser a t the American Gebo, DL (1986)The Anatomy of the Prosimian Foot and
its Application to the Primate Fossil Record. Ph.D.
Museum of Natural History and R. Thoringthesis, Duke University.
ton at the US. National Museum were also
Godinot, M, and Dagosto, M (1985) The astragalus of
helpful in allowing access to the many osteoNecrolemur (Primates, Microchoerinae). J. Paleon.
logical and paleontological specimens in their
57:1321-1324.
care.
Hafferl, A (1932)Bau und Funktion des AffenfuBes. Ein
NOTE ADDED IN PROOF

Crompton and Andaus recent article in

Primates, Volume 27:337-355 (1986), presents field data on the locomotor behavior of
Tarsius bancanus. 7: bancanus leaps 66.1%,
climbs 27.6%, quadrupedally walks 3.2%,
hops 2.3% and cantilevers 0.4% of the time.
Their field observations further document 71
banacus preference for vertical clinging postures as well as the absence of quadrupedal
suspensory postures or movements. Crompton and Andau (1986, p. 337) believe that
Tarsius bancanus locomotion is similar to
but more specialized than that of Tarsius
spectrum. Thus, these new field observations add support to the tarsier observations
made in a captive setting at the Duke Primate Center.
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