The Anatomy of the Forelimb in the Anteater (Tarnandua)

and Its Functional Implications

BRUCE K. TAYLOR
Department of Cell Biology, The University of Texas, Health Science Center at Dallas,
Dallas, Texas 75235

ABSTRACT The forelimbs of anteaters play a major role in obtainment of

food, defense, and locomotion. The greatly enlarged claws on the manus are
used for ripping open insect nests and insect-infested wood; the claws also serve
as the animals' only defensive weapons, since they lack teeth. Specialization of
the claws for these functions has also had a substantial effect on the ways in
which the forelimb is used for posture and locomotion.
Modifications of the forelimb in the anteater Tamandua include the follow-
ing. Attachments of the medial head of triceps are rearranged so as to greatly
increase capability for powerful flexion of the claws. Ability to flex the elbow
and to retract the humerus is also augmented; these movements would assist
digital flexion in applying traction with the claws to material being torn away
during food procurement. This traction can be supplemented by a variety of
powerful side-to-side andlor twisting movements of the hand, brought about
primarily by axial rotation of the upper arm and forearm. The digital joints are
reinforced to resist the deviational and torsional loading to which the digits
would be subjected during such movements.
The morphological modifications of the forelimb in Tamandua are discussed
in terms of how they affect the mechanical capabilities of the limb, what func-
tions the limb is best designed to perform, how they may relate to what little is
known about the specialized behavior of this animal, and what behavioral pre-
dictions may be made based on mechanical design.

This paper deals with the very distinctive
anatomical specializations of the forelimb of
the anteater Tamundua. This animal is inter-
mediate in size between the giant anteater
Myrmecophaga and the little silky anteater
Cyclopes. Unlike the other genera (Myrmeco-
phuga being almost wholly terrestrial and Cy-
clopes being almost wholly arboreal), Tuman-
d u a moves freely between terrestrial and ar-
boreal environments.
The specializations shown by the forelimb of
Tamandua reflect more than one major bio-
logical role. In addition to its locomotor func-
tions, the forelimb is used as an organ for ob-
taining food and for defense. Tamandua, like
the other two myrmecophagid genera, has
slender mandibles, small jaw muscles, an ex-
tremely small mouth gape, and no teeth.
These facts reflect a very specialized diet and
method of obtaining food and also t h a t the
mouth is of no use in defense.
J. MORPH. (1978)157: 347-368.
The feeding strategy of Tamandua involves
utilization of social insects nesting in great
numbers (Azara, 1838; Cabrera, '29; Walker
et al., '64). Penetration of insect nests or in-
sect-infested wood is accomplished by ripping
them open with greatly enlarged central fore-
claws (figs. lC, 2C, 3C). The chief role of the
forelimb in this type of feeding behavior in-
volves forcefully pulling the claws toward the
body.
Since anteaters do not utilize burrows and
cannot use their jaws for protection, they have
developed defensive behavior which involves
use of their enlarged claws. When provoked,
their defensive posture involves erecting the
body on the hindlimbs and holding the fore-
claws out on either side of the head ready to
strike a t an adversary (Azara, 1838; Cabrera,
'29).
It is the purpose of this paper to examine the
anatomy of the forelimb in Tamandua in
347
348 BRUCE K TAYLOR
terms of: (11 t h e identification of those fea-
tures which distinguish i t from t h e limbs of
other (non-myrmecophagid)mammals; (2) its
mechanical capabilities as determined by
morphology (especially as these capabilities
relate to distinguishing features); (3) iden-
tifying those biological roles to which t h e
structure is most closely matched; (4) how
limb morphology does or may relate to what is
known about the specialized behavior of this
animal; and (5) what behavioral predictions
may be made based on mechanical design.
Both Maynard Smith and Savage ('56) and
Abbrrviutrons
AV. Acromioii process of scapula
ADLT, Acromiodeltoid
ALS, Anterior divLsion, levator scapulae
A N C N . Anconeus
ATHP. Acromiotrapezius
BIC, Biceps
BICL, Long head of biceps
BICS, Short head of hiceps
BRCH, Brachialis
BRD. Brachioradialis
AL, C,oraco-acromial ligament
AP. Capitulum of humerus
CDLT, Clavodeltoid
CEPH. Cephalohumeralis
CORB. Coracobrachialis
CR?, Coraroid process of scapula
DBCL. Deep division, long head of hiceps
DBRD, Deep head, brachioradialis
IIEL. Delto-epicondylar ligament
DEPI, Dorso~epitr1,chlearis
DLT. Deltoid
U P Distal phalanx
DPR, Delto-pectoral ridge
B, Deltoid tubercle
K. Extensor carpi radialis
U, Extensor carpi ulnaris
C, Extensor digitorum coinmunis
EFDP. Epicondylar head. flexor
digitorum profundus
EOMP. Extensor ossis metacarpi pollicis
EPAN, Epitrochleo-anconeus
EXIN, Extensor indicis
FCR, Flexor carpi radialis
FCU. Flexor carpi ulnaris
FUI'. Flexor digitorum profundus
FDS, Flexor digitorum sublimis
GTB, Greater tuberosity of humerus
IBM, Distal border of medial epicondyle
INFR, Infraspinatus
L, Lateral epicondyle of h u m e r u s
LATD, Latissimus dorsi
LN. Lunar
LS. Levator scapulae
LTB, Lesser tuberosity of humerus
M, Medial epicondyle of humerus
Scale is 1 cm in all figures.
Hildebrand ('74) mentioned anteaters within
the context of more general discussions on
fossorial adaptations in mammals. Yalden
('66b. '72) included T a m a n d u a and Myr-
mecophaga in his studies on the functional
anatomy of t h e mammalian carpus. However,
with t h e exception of these aut,hors, no previ-
ous work has paid diligent attention to the
mechanical capabilities of forelimb morpholo-
gy and how they may relate to behavior in
anteaters. The anatomy of these animals is
reasonably well known from the work of a
number of nineteenth cent,ury anatomists (see
MA, Metacromion process of scapula
MC, Metacarpal
MN, Magnum
MI', Middle phalanx
OLEC. Olecranon process of ulna
PABD, Pectoralis ahdominis
PALM, Palmaris longus
PI, Pisiform
PLS, Posterior division, levator scapulae
PMAJ. Pectoralis major
PMIN, Pectoralis minor
PP, Proximal phalanx
PPR, Posterior process of spine of scapula
PR. Pectoral ridge
PROQ, Pronator quadratus
PROT, Pronator teres
PS, Postscapular fossa
RFDP. Radial head, flexor d i g t o r u m profundus
RH, Rhomhoideus
RTB, Radial tuberosity
S, Scaphoid
SBCL. Superficial division, long head of hiceps
SBRD, Superficial head, brachioradialis
SCF, Supracoracoid foramen of scapula
SCN. Supracoracoid notch of scapula
SCR. Supracondyloid ridge of humerus
SDLT, Spinodeltoid
SER, Serratus magnus
S P , Primary spine of scapula
S S P , Secondary spine of scapula
SUBS, Subscapularis
S U P N , Supinator
SUPR, Supraspinatus
TD. Trapezoid
TM, Trapezium
TMAJ, Teres major
TMIN. Teres minor
TR. Trochlea of humerus
TRIC, Triceps
TRL, Lateral head of triceps
TRM, Medial head of triceps
TRP, Trapezius
TRS, Scapular head of triceps
UFDP, Ulnar head, flexor digitorum profundus
UN. Unciform
 FORELIMB ANATOMY I N TAMANDUA 349

INFR
TMAJ
PS
SSP )c-c 'MIN

./- (DLT,

ANCN,, PMAJ

Fig. 1 Forelimb skeleton in lateral view: (A) Didelphis; (B) muscle attachment areas in Dzdelphis; (C)
Tamandua; (D) muscle attachment areas in Tamandua.

Windle and Parsons, 1899, for a review), but
little, if any, attention was paid to function by
any of those workers.
MATERIALS A N D METHODS
delphis marsupialis, was chosen as a repre-
sentative generalized mammal. In forelimb
morphology, it is relatively unspecialized, and
in terms -of posture and- movements of t h e
limb, i t Drobablv
" amroximates the Drimitive
I -

In order to provide a standard of com- therlan condition as well as any l i v k g mam-

parison, t h e North American opossum, Di- mal and certainly better than cursorial forms
350 BRUCE K. TAYLOR
such as Felis (Jenkins, '71). Since the opossum
is similar in size to Tamandua, comparisons
a r e f a c i l i t a t e d by m i n i m i z i n g possible
allometric differences. Comparisons were also
made with Dasypus to see how the specializa-
t i o n s of t h e a n t e a t e r d i f f e r from t h e
stereotyped digging adaptations of armadillos.
Skeletal material and wet specimens for dis-
section were obtained from various museum
and university collections. Dissections were
performed on four specimens of Tamandua,
two of Didelphis, and two of Dasypus, using
the muscle staining technique described by
Bock and Shear ('72). Skeletal material from
the dissected animals was supplemented with
museum skeletons, making a total of nine
skeletons of Tamandua, five of Didelphis, and
eight of Dasypus.. Only adult skeletal mate-
rial was used.
The forelimb muscles of one specimen each
of Tamandua and Didelphis were weighed.
Wet muscles were trimmed of excess connec-
tive tissue, blotted to remove excess preserva-
tive, and weighed to the nearest 0.01 gm;
these values were rounded to t h e nearest 0.1
gm. These weights were not taken for t h e pur-
pose of a thorough quantitative study, which
would have required much larger samples
than were available. For the purpose of this
study, the emphasis is on differences which
are large and obvious. Subtle differences, par-
ticularly with the limited data presented here,
cannot by themselves be regarded as having
any real biological meaning. The muscle
weights are presented in table 1. Some obser-
vations were also made of forelimb use in zoo
specimens of Tamandua.
RESULTS AND DISCUSSION
Skeletal design and significant features
of joint surfaces
The forelimb skeleton of T a m a n d u a is char-
acterized by the following features.
The scapula is expanded in width, primarily
due to development of a postscapular fossa
(PS: fig. 1 0 , which is separated from the in-
fraspinous fossa by a secondary spine (SSP).
The primary spine (SP: fig. 1C) of the
scapula is quite prominent, ending in a long
acromion process (AC) t h a t arches forward
and medially, rather high above the shoulder
joint (compare with the short acromion in
Didelphis, fig. 1A). The upward direction of
the acromion in T a m a n d u a distinguishes i t
from other forms with a long acromion, such
as armadillos (e.g., Dasypus, fig. 2A), in which
the acromion curves medially but does not ex-
tend high above the shoulder joint.
The deltoid tubercle (DTB: figs. 1C, 4A,B) of
the humerus is quite large and projects later-
ally; compare with Didelphis (figs. lA, 5A,B).
The distal end of the humerus is very wide
mediolaterally, mostly due to the large medial
epicondyle (M: figs. 4A,B; compare with
Didelphis, figs. 5A,B).
The proximal end of the lateral supracon-
dyloid ridge (SCR: figs. 4A,B) of the humerus
is connected to the distal end of the deltoid
tubercle by a stout delto-epicondylar ligament
(DEL: figs. 4A,B). The ventrolateral aspect of
t h e deltoid tubercle, the delto-epicondylar
ligament, the supracondyloid ridge, and the
lateral epicondyle (L) all provide attachment
area for the antebrachial extensor muscles
(fig. 4A). The line of origin for this muscle
group is thus extended proximally more than
half the length of t h e humerus. In Didelphis
(fig. 5A), there is no such ligament, and since
none of the forearm extensors attach above
the supracondyloid ridge, their origin is re-
stricted to t h e distal one-fourth of the length
of the humerus, a condition typical of most
other mammals.
Compared with Didelphis (fig. 5A,B), the
humeral trochlea of T a m a n d u a (TR: figs.
4A,B) forms a relatively deep and narrow in-
tercondylar groove, and there is also a rela-
tively large capitulum (CAP) for the head of
the radius.
The carpus of T a m a n d u a has been described
in detail by Yalden ('66b, '72); my own obser-
vations are in agreement with his.
Digit I11 (fig. 2C) is much longer and more
robust in other dimensions than the others.
The metacarpals of all of the digits are similar
in length; the main difference is t h a t all but
the third (MC) are slender. The proximal pha-
lanx (PP) of digit I11 is extremely short, while
the distal phalanx (DP) is quite long, with a
very large claw. Digit V bears no claw and lies
completely buried in the palmar pad.
In t h e third digit of Tamandua, the heads of
the metacarpal (MC: fig. 14) and the middle
phalanx (MP: fig. 16) are spool-shaped. The
metacarpal is also equipped with a very large
median keel (larger than the condyles on
either side of it) which runs between the flex-
or and extensor extremes of the distal joint
surface. The base of the proximal phalanx
(PP: fig. 14) has three deep concavities into
 FORELIMB ANATOMY IN TAMANDUA 351

PS

MA

AC

GTB

PR

n"
OLEC
DT B

I"

MN

C
Fig. 2 (A) D Q S Y ~ Uforelimb
S, skeleton in lateral view; (B) Didelphis, extensor aspect o f carpus and digits;
(C) Tamandua, extensor aspect of carpus and digits.

which fit the two condyles and median keel of
the metacarpal. The head of the middle pha-
lanx has no median keel, but fits into a deep
socket in the base of the distal phalanx (DP:
fig. 16). In contrast, the head of the proximal
phalanx (PP: fig. 15) and the base of the mid-
dle phalanx (MP: fig. 15) have been modified
so that there is a complex array of flat facets
which virtually eliminates all movement a t
the proximal interphalangeal joint. There are
well developed collateral ligaments a t the
sides of each joint.
Correlation o f myology with skeletal
design and its functional
implications
Movements of digits
Modification of triceps and its effects
on digital flexion
Perhaps t h e most conspicuous feature
which distinguishes the forelimb of T a m a n -
d u a from those of non-anteaters is the unique
arrangement of the medial head (TRM: fig.
6C) of triceps. Unlike the condition common to
352 BRUCE K . TAYLOR

L SBCL
Flg 3 Forelimb skeleton in medial mew (A) Dzdelphzs, (B) muscle attachment areas in Dzdelphbs, (C)
T a m a n d u a , (D)muscle attachment areas in T a m a n d u a

all other groups of mammals, in Tamandua
t h e medial head of triceps (52%of t h e total tri-
ceps musculature: see table 1) does not insert
on t h e ulna. J u s t proximal to t h e elbow joint, a
tendon develops on t h e deep surface of the
muscle. This tendon, accompanied by fleshy
fibers superficial to it, passes under the distal
border (IBM: figs. 4A,B) of t h e medial epicon-
dyle, deep to epitrochleo-anconeus (EPAN:
fig. 6D). Upon passing into t h e forearm, the
tendon thickens; this portion of the tendon
slides proximally past t h e distal border of
 FORELIMB ANATOMY IN TAMANDUA 353
the medial epicondyle upon contraction of the tendon serving these muscles then passes
muscle. The tendon abruptly narrows, a t through the distal forearm and into the hand.
which point fleshy fibers of medial triceps end, The contribution from medial triceps serves
and the tendon becomes continuous with that mainly the third digit; insertion is onto a
of flexor digitorum profundus (fig. 8B). The prominent tubercle on the flexor aspect of the
base of the distal phalanx. This arrangement
was first described by Macalister (18751, but
it has never been analyzed functionally.

ECU SUPN FOP

M

INFR
HEAD
TMIN /

TRM LTB GTE

DLT

B
B DTB

SCR
ANCN

L
FDP TR

Fig. 4 Humerus in Tamandua, with muscle attach- Fig. 5 Humerus in Didelphis, with muscle attachment
ment areas shown in the right-hand figures; (A) ventral areas shown in the right-hand figures: (A) ventral view;
view; (B) dorsal view. (B) dorsal view.

TABLE 1

Muscle wecghts ( . )
Didelphis Tamandua Didelphis Tamandua Lhdelphis Tamandua

CEPH - 1.7 TMIN - 0.3 SBRD - 3.7

TRP 5.8 3.0 a SUBS 4.1 7.4 DBRD - 3.0
PMAJ 8.9 5.4 TMAJ 1.5 4.5 ECR 1.3 6.2
PABD 2.7 1.8 CORB 0.1 0.3 EDC :' 0.6 2.2
PMIN 3.1 1.8 1.3 0.9 ECU 0.4 2.0
LATD 10.1 13.6 0.5 - SUPN 0.2 1.1
RH 1.9 2.1 - 1.1 EXIN 0.1 0.4
SER 3.3 2.9 DBCL - 1.0 EOMP 0.4 1.2
LS 3.2 - BRCH 1.4 0.9 PROT 0.8 2.6
ALS - 3.3 TRS 4.3 6.7 FCR 0.3 1.2
PLS - 1.5 TRL 2.4 3.2 PALM 0.2 -
SDLT 0.9 1.4 TRM 1.8 10.7 FDS 0.2 2.0
ADLT 0.6 4.0 DEPI 0.8 2.4 FCU 0.9 3.5
CDLT 1.0 -a ANCN 0.1 0.7 FDP 3.1 11.7
SUPR 2.7 3.9 EPAN 0.1 1.6 PROQ 0.1 0.8
- -
INFR 2.6 2.4 BRD 0.5 - Total 74.3 132.1

' Length of scapula along spine = 52.5 mm.

Length of scapula along spine = 52.7 mm.
Includes proper extensors of d i g t s IV and V.
a Weight of clavicular portion included under cephalohumeralis
 354

Fig. 6 Musculature in medial aspect: (A) Didelphis, with superficial forearm muscles removed; (B) Didelph~s,super-
ficial forearm muscles; (C) Tamandua, with superficial forearm muscles removed; (D) Tamandua, superficial forearm
muscles.
 FORELIMB ANATOMY IN TAMANDUA 355
TRP

TRS
BRCH

c--- BRD

EDC
E C U \= P
ECR
EOHP

Fig. 7 Musculature in lateral aspect: (A) Didelphis, deep muscles; (B) Didelphis, superficial muscles; (C) Tarnan-
dua, deep muscles; (d) Tamandua, superficial muscles.
356 BRUCE K. TAYLOR

N
/- RFDP.
A TRM
IS
I I /

7
C OMMON
C
TENDON
B
Y

Fig. 8A Ventral view of humerus i n Tamandua. The approximate axis of flexion-extension of t h e elbow is
indicated by t h e dashed line. Due t o t h e concavity of t h e distal border of t h e medial epicondyle, t h e tendon of
medial triceps (shown in cross section) passes through this axis.
B Medial head of triceps in Tamandua, shown semi-diagrammatically in medial view. Attachment
areas ar e indicated for t h e three heads of flexor digitorum profundus, with which medial triceps inserts. The
approximate position of t h e axis of flexion-extension of the elbow is indicated by a dark circle.
C Anconei in Tamandua, shown in dorsal (above) and posterior (below) views of humerus and elbow
joint

The remaining heads of triceps in T a - compare Didelphis and Tamanduai. In addi-

mandua TRS, TRL: figs. 6C, 7 0 retain their
primitive attachments; i.e., they insert on t h e
olecranon process of the ulna.
The effects of medial triceps upon t h e elbow
joint must be very small in Tamandua. As can
be seen from figure 8A, t h e tendon of medial
triceps passes through or very close to t h e axis
of flexion-extension of t h e humero-ulnar joint.
It is true t h a t those fibers which pass on t h e
extensor side of this axis would tend to extend
the joint, and those fibers which pass on t h e
flexor side would tend to flex the joint. How-
ever, any effects on flexion or extension of t h e
elbow are relatively minor when compared
either with the other two heads of triceps in
Tamandua or with any of t h e heads of triceps
in other mammals.
In Tamandua, t h e major effect of the modi-
fied attachments of medial triceps is upon dig-
tion, the weight of t h e medial head of triceps
(TRM: table 1) in Tamandua is 91% that of
flexor digitorum profundus in t h e same ani-
mal; thus, shifting the insertion of medial tri-
ceps almost doubles the already large amount
of musculature involved in digital flexion.
Digital joints
The outstanding features of the digital
joints in T a m a n d u a relate to restriction of
movements. There is no significant movement
possible a t t h e proximal inter-phalangeal
joint. The metacarpo-phalangeal and distal
inter-phalangeal joints, however, both allow
one type of movement but restrict others. The
median keel on t h e metacarpal provides added
lateral stability in all joint positions, while its
smooth convex curve in FES ' allows a wide
I FES = flexor-extensor section. i.e , cross section cut from the

ital flexion. Flexor digitorum profundus in flexor to t h e extensor sides of the carpus, with the plane of section
parallel to the long axis of the wrist and hand (terminology of
this animal is a large muscle (FDP: table 1; Yalden, '721.
 FORELIMB ANATOMY IN TAMANDUA
range of flexion and extension. The side walls
of the concave socket in the base of the distal
phalanx perform the same function, i.e., pre-
cluding lateral movement while allowing a
full range of flexion-extension. Lateral and/or
torsional stresses would certainly occur dur-
ing twisting motions of the hand produced by
pronation and supination of the radius against
resistance applied to the digitk). These con-
siderations also suggest that Tamandua may
pry apart objects with the side of the claw as
well as with the flexor aspect.
In T a m a n d u a there are two joints on the
digit a t which flexion may occur: the meta-
carpo-phalangeal and distal inter-phalangeal
joints. From the amount of musculature (me-
dial triceps plus flexor digitorum profundus,
discussed above) which inserts a t the base of
the distal phalanx, i t seems clear that flexion
of the distal inter-phalangeal joint is of major
importance for tearing functions. Flexion of
the metacarpo-phalangeal joint, on the other
hand, accomplishes the following. First, it in-
creases the range of flexion-extension of the
entire digit. Second, it brings the phalanges
out of the way during quadrupedal walking, in
which the weight is borne on the heads of the
metacarpals (Yalden, '66b). Third, it swings
the phalanges around toward the palmar as-
pect of the hand, thus bringing the claws into
opposition with the palmar pad and allowing
the hand to perform grasping functions. The
ability to use the hand as a grasping organ is
of obvious importance for climbing. In addi-
tion, this ability increases the manipulative
capability of the hand; thus, for example,
pieces of termite nest may be grasped and
pulled away rather than simply pried apart.
Retraction of distal part of limb
toward body
Elbow flexion
In addition to biceps and brachialis, there
are two antebrachial muscles, brachioradialis
and extensor carpi radialis, which might play
a major role in elbow flexion in Tamandua.
The outstanding features of brachioradialis
are its large size (table 11, proximal origin on
the humerus (BRD: figs. lD, 4A), and the fact
that its superficial portion (SBRD: figs. 6D,
7D) wraps around the forearm to insert by an
aponeurosis into the deep fascia of the flexor
aspect of the forearm and wrist.
Long and short heads of extensor carpi ra-
dialis cannot be distinguished in anteaters.
Galton (18691, Humphry (18691, Macalister
357
(18751, and Windle and Parsons (1899) all des-
ignated this muscle as extensor carpi radialis
brevis, the longus being absent. The muscle
(ECR: fig. 7D) is very large in Tamandua
(table 1).It arises rather high in the arm, tak-
ing origin from the lowermost part of the tip of
the deltoid tubercle, the entire length of the
delto-epicondylar ligament, and the upper
third of the supracondyloid ridge (figs. lD,
4A). In addition, a tendon, from which fleshy
fibers arise, fans out over the proximal third of
the inferior side of the muscle belly. Insertion
is onto a prominent scar on the extensor
aspect of the middle of the third metacarpal.
The proximal shift in the origins of brachio-
radialis and extensor carpi radialis, facili-
tated by the presence of the delto-epicondylar
ligament, distinguishes Tamandua from most
other mammals, in which the origins of the
antebrachial extensors are confined to the epi-
condyle and supracondyloid ridge. These mus-
cles are thus displaced relatively further from
the elbow joint in Tamandua, which increases
their moment arms for flexion of that joint.
While brachialis is relatively small in T a m a n -
d u a (see below), its reduction is offset to some
extent by the increased size and mechanical
advantage of brachioradialis and extensor car-
pi radialis. The specimens of Tamnndua and
Didelphis for which muscles were weighed
were of nearly equal body size (in terms of
overall linear dimensions, not weight), yet in
Tamandua brachioradialis was about ten
times as massive and extensor carpi radialis
was about five times as massive as their re-
spective homologs in Didelphis (table 1).
Thus, T a m a n d u a is adapted for much more
powerful flexion of the elbow as compared
with Didelphis. Tamandua may use powerful
elbow flexion in conjunction with digital flex-
ion in order to pull material toward the body
(fig. 9B).
Shoulder retraction
The humerus can be retracted by teres
major and latissimus dorsi. In Tamandua,
teres major (TMAJ: figs. 6C, 7C) arises from
the posterior half of the postscapular fossa
(fig. 1D). Its insertion extends distally on the
humerus to about its midpoint (figs. 3D,
4A,B). Between its attachments, it is tightly
bound to latissimus dorsi. Ventral expansion
of the postscapular fossa and a more distal in-
sertion both contribute to increasing the
moment arm of teres major in Tamandua rel-
ative to the didelphid condition (compare figs.
358 BRUCE K TAYLOR

L
a
\
limb caudally; this is likely to be useful in
climbing, in attacking prey nests, and in de-
fensive behavior.
Retraction of the limb as a whole
To summarize the above, flexion of the digit
(fig. 9A) may be supplemented by retraction of
the humerus and flexion of the elbow in order
MOVEMENT to draw the flexed claw proximally (fig. 9B).
OF LIME The principal muscles involved in these move-
MOVEMENl ments in T a m a n d u a are shown diagram-
OF C L A W
matically in figure 9C, and the same muscles
A B are shown in Didelphis in figure 9D for com-
parison (latissimus dorsi and the proximal
portions of the digital flexors are omitted for
clarity). The major differences in Tamandua
are: (1) fusion of medial triceps with flexor
digitorum profundus (discussed above) ; (2) in-
creased moment arm of brachioradialis (BRD)
and extensor carpi radialis (ECR) due to more
proximal origin; (3) increased moment arm for
teres major (TMAJ) due primarily to more dis-
L D tal insertion; and (4)vastly increased size of
Fig. 9 Flexion of limb segments: (A) flexion of claw in
all of the above muscles (table I). It may be
Tarnandua; (B) retraction of humerus and flexion of predicted on the basis of these features that
elbow and wrist in Tarnandua in order to approximate t h e T a m a n d u a habitually uses digital flexion sup-
manus to the body; (C) diagram of main limb flexors in plemented by flexion of the elbow and retrac-
Tarnandua; (D) diagram of main limb flexors in Di-
delphzs.
tion of the shoulder in order to apply powerful
traction to material against which the claws
may be engaged.
9C, and 9D). Also, the muscle itself is rela- Extension of reach
tively larger in Tamandua, as can be appreci-
ated from its increased area of origin (com- Elbow extension
pare figs. 1B and 1D) and its relative weight In Tamandua, the lateral and scapular
(table 1). heads of triceps (TRL, TRS: figs. 6C, 7C) in-
Latissimus dorsi (LATD: fig. 7D) arises in sert on the olecranon process of the ulna and
Tamandua via the dorso-lumbar aponeurosis are well developed (table 1).Elbow extension
from the dorsal midline and the dorsolateral is also aided in most mammals by dorso-
body wall. The expansion of latissmus onto the epitrochlearis, which inserts into the deep
lateral body wall increases its posterior com- fascia of the extensor aspect of the proximal
ponent of pull as compared to the condition forearm and often onto the olecranon as well
seen in Didelphis (fig. 7B), in which the mus- (Jouffroy, '71). This muscle arises from the
cle arises only from the dorsal midline. The tendon of latissimus dorsi a t approximately a
firm connective tissue attachment (which 90" angle to it (DEPI: figs. 7B,D); thus its
does not occur in Didelphis) between latissi- point of origin is, in most mammals, rather
mus dorsi and teres major would limit the cap- loosely slung between the attachments of
ability of these two muscles to act indepen- latissimus to the body wall and to the hu-
dently in Tamandua. It also has the effect of merus. It follows that, upon contraction of
providing a more stable base of origin for dor- dorso-epitrochlearis,some of its force would be
so-epitrochlearis (discussed below). expended in tensing latissimus dorsi. In T a -
As a result of the above modifications of mandua, however, that portion of latissimus
teres major and latissimus dorsi, T a m a n d u a dorsi from which dorsi-epitrochlearis arises is
has a greater capability for powerful retrac- firmly bound down over the teres major fascia
tion of the shoulder than occurs in relatively a t the caudal angle of the scapula. The ef-
unspecialized mammals such as Didelphis. fect is to stabilize the origin of dorso-epi-
Retraction of the shoulder rotates the entire trochlearis, thus allowing most of its force to
 FORELIMB ANATOMY IN T A M A N D U A 359
be exerted a t its distal attachment Le., the
olecranon). (~ AXIS

Protraction of shoulder A
Protraction of the humerus in Tamandua
may be accomplished by cephalohumeralis,
acromiodeltoid, and supraspinatus. The first
of these muscles (CEPH: fig. 7D) is formed by
fusion of the primitive clavicular heads of tra-
pezius and deltoid, the clavicle being either -
tj
AXIS

absent or vestigial in Tamandua.

The mediad curvature of the acromion .
around the anterior side of the shoulder joint Fig. 10 Side view of radius, showing moment arm of
(which occurs in Tamandua but not in more pronator teres: (A) Didelphis; (B) Tamandua.
generalized mammals) has the effect of in-
creasing t h e capability of acromiodeltoid
(ADLT: fig. 7D) for protraction of the humer- fig. 6D) inserts by a broad aponeurosis onto
us, since it orients the muscle in front of the the distal half of the radius (figs. lD, 3D),
shoulder and not just to the side of it. Also, theadjacent to t h e insertion of supinator
elevation of the acromion above the shoulder (SUPN). In Didelphis (fig. 6B), the muscle in-
allows extension to occur over a wide range serts on the middle third of the radius (fig.
without interference between the acromion 3B). Pronator quadratus (PROQ: fig. 6C) in
and the shaft of the humerus (compare figs. Tamandua covers virtually the entire flexor
surface of the interosseus membrane and
1C and 2A).
adjacent surfaces of the radius and ulna; it
Supraspinatus (SUPR: fig. 7C) may also
does not cover the surfaces of the bones as ex-
assist in extension of the humerus, although it
tensively in Didelphis (figs. 3B,D). The rela-
probably primarily aids infraspinatus, teres
tively large size of these muscles in Taman-
minor, and subscapularis in maintenance of
dua (table 1) suggests that pronation of the
joint integrity. Being short-fibered and pen-
radius often occurs under significant loading
nate in most mammals, the muscle is powerful in this genus.
and a t the same time incapable of being
stretched a great deal. Indeed, the shortness of An important feature of the mechanics of
its fibers makes it efficient over only a very pronator teres is the length of its moment arm
narrow range of deviation from its optimum for axial rotation of the radius. This moment
length. These characteristics would be con- arm is increased by displacing the insertion
sistent with the interpretation that the mus- away from the axis of rotation. In Didelphis
cle is primarily adapted for maintaining this is accomplished by bowing of the radius,
shoulder joint integrity in the face of large as shown in figure 10A. In Tamandua, the
dislocating forces. moment arm for the more distal fibers is in-
creased due to the flaring out of the distal
Axial rotation of limb segments crest of the radius, as shown in figure 10B.
Forearm rotation The muscles which supinate the radius in
Tamandua are supinator, the short head of
While the range of pronation-supination in biceps (plus part of the long head), and
anteaters may be somewhat limited, it is by brachioradialis. The well-developed supinator
no means insignificant. According to Yalden (SUPN: fig. 7C) of Tamandua arises from the
('66b), the radius of the giant anteater Myr- lateral epicondyle and inserts along the prox-
mecophaga can axially rotate through a range imal two-thirds of the radius (fig. 1D). In the
of approximately 5 0 O . ' My own observations in opossum, supinator (fig. 7A) is much smaller
Tamandua also indicate a range of about 50"; than in Tamandua (table 11, and i t inserts
this compares with a range of about 90" in only along the proximal third of the radius
Didelphis. (fig. 1B).
The muscles involved in pronation of the
hand are pronator teres and pronator quad- In Myrrnecophaga, Yalden also measured approximately 40" of
axial rotation in the wrist, chiefly at the mid-carpal Jolnt, thus pro-
ratus. These muscles are both very well de- viding a total of about SO" of possible rotation of the hand resulting
veloped in Tamandua. Pronator teres (PROT: from movements occurring at or distal to the elbow.
360 BRUCE K. TAYLOR
BlCS
SBCL
BlCL
BlCS
,CORE
A benefit when simple traction is not sufficient
Fig. 11 Brachial flexors in ventral aspect, with rela-
tion of humerus shown for orientation: (A) Didelphis; (B)
Tamandua.
A distinctive feature of biceps in T a m a n d u a
is the bifurcation of t h e long head into su-
perficial and deep divisions (fig. 11B). The su-
perficial division (SBCL) fuses distally with
t h e short head (BICS); these insert via a com-
mon tendon onto t h e radial tuberosity. The
deep division of t h e long head (DBCL) fuses
distally with brachialis (BRCH); these insert
onto t h e coronoid process of t h e ulna.
Both brachialis and t h e short head of biceps
appear to be somewhat smaller in T a m u n d u a
than in Didelphis. The long head of biceps, on
t h e other hand, is considerably larger in
Tamandua (table 1).The functional signifi-
cance of t h e relative sizes of these muscles
depends on the nature of their attachments.
Brachialis always inserts on the ulna and is
thus a simple flexor of t h e elbow with no con-
current rotatory action upon t h e radius. The
nature of the insertion of biceps is variable in
mammals, but t h e primitive arrangement ap-
pears to be t h a t which is seen in Didelphis
(Howell, '37; Jouffroy, '711, i.e., in which t h e
short head inserts entirely on t h e radius and
t h e long head inserts entirely on t h e ulna. In
this case, the short head would produce elbow
flexion and concurrent supination, while t h e
long head would produce only flexion. In
Tamandua, however, t h e shifting of approx-
imately half of t h e long head over to t h e
radius allows recruitment of additional bra-
chial flexor musculature which will produce
concurrent supination.
Another muscle in T a m a n d u a which has a
substantial flexor action a t t h e elbow is
brachioradialis, this by virtue of t h e extreme
proximal position of its origin (discussed
above). This muscle also tends to produce
supination, since i t wraps around t h e medial
aspect of t h e forearm as i t proceeds distally
(fig. 6D). Thus, in Tamandua, a large portion
of the musculature which flexes t h e elbow also
produces supination of t h e forearm.
Pronation and supination would impart
twisting motions to t h e hand which may be
useful in loosening pieces of material when
t h e animal is tearing a n object apart. In addi-
tion, combined supination and elbow flexion
would twist t h e material while i t is being
pulled toward t h e body, which is likely to be of
to pull t h e material apart. The above kinds of
movements were in fact observed in a captive
individual of Tamandua which was tearing
apart a small log.
Although not directly related to pronation
and supination, there is a n additional feature
of brachioradialis in Tumanduu which merits
mention here. Whereas in t h e opossum t h e
muscle inserts entirely on the radial aspect of
t h e extensor side of t h e wrist, in Tamandua
t h e muscle continues around t h e medial
aspect of t h e forearm to insert on t h e flexor
side of t h e wrist (compare figs. 6B and 6D).
Thus in t h e anteater this muscle may be
regarded as a flexor of t h e wrist as well as a
supinator of t h e forearm and a flexor of t h e
elbow. The implication is t h a t all three of
these movements may habitually occur to-
gether. This statement does not mean, how-
ever, t h a t flexion of t h e wrist necessarily oc-
curs every time t h e elbow is flexed. The nature
of t h e attachments of extensor carpi radialis
and of brachioradialis would enable one or t h e
other of these muscles to assist t h e brachial
flexors in flexing t h e elbow while also produc-
ing t h e desired movement (either extension or
flexion) at t h e wrist. Also, combined contrac-
tion of both of these muscles could be used to
produce radial deviation of t h e hand while
stabilizing t h e wrist against both extension
and flexion.
Humeral rotation
One of t h e principal lateral rotators of t h e
humerus is t h e spinal head of deltoid (SDLT:
fig. 7D). Its origin in Tamandua (DLT: fig.
1D) is along t h e spine of t h e scapula and from
t h e caudal part of t h e infraspinous fascia as
well. One aspect of this muscle, in which T a -
m a n d u a stands out in sharp contrast to gen-
eralized mammals such as Didelphis, is its mo-
ment a r m for axial rotation of the humerus,
which is directly related to laterad displace-
ment of its insertion away from the shaft. The
laterad projection of t h e deltoid tubercle in
edentates reaches its extreme in anteaters
 FORELIMB ANATOMY IN TAMANDUA
(DTB: figs. 4A,B). This maximizes the mo-
ment arm of spinodeltoid for lateral rotation
of the humerus, because it moves the line of
action of the muscle further away from the
axis of rotation.
Among the medial rotators of the humerus,
subscapularis (SUBS: fig. 6C) stands out as
being modified for increased power in Taman-
dua, although its moment arm ( a function of
the degree of mediad projection of the lesser
tuberosity of the humerus) is not significantly
increased. In Tamandua, subscapularis has a
complex arrangement of nine distinct adja-
cent multipennate sections, seven of which
arise from the subscapularis fossa (fig. 3D). Of
the remaining two sections, one arises from
the anterior border of the scapula, the tip of
the coracoid process (CRC: fig. lC,D),and part
of the coraco-acromial ligament (CAL: fig.
7C); the other arises from the anterior half
of the postscapular fossa (fig. lD) and thus
wraps around the axillary border. The first of
these sections has a somewhat separate inser-
tion on the dorsomedial aspect of the lesser
tuberosity of the humerus. The sections aris-
ing on the medial surface of the scapula are
separated from one another by septa which a t -
tach deeply to the surface of the bone. Each
section has a centrally located tendon of inser-
tion. Distally, these central tendons fuse to
insert on the medial aspect of the lesser
tuberosity of the humerus (figs. 3D, 4A,B). In
the opossum, subscapularis (figs. 3B, 6A)
arises from the medial surface of the scapula
only. It is not subdivided. Its insertion tendon
develops as a sheet on the medial surface
which converges distally. In Tamandua, the
subdivision of this muscle into many pennate
bundles increases t h e number of muscle
fibers. Increasing the amount of fibers is fur-
ther facilitated by increased width of the
scapula and by extension of the origin of the
muscle around the axillary and anterior bor-
ders of the scapula.
All of the above features suggest that
humeral axial rotation is of major importance
in Tarnandua. This genus characteristically
walks with the elbows widely abducted. If the
elbow were fully abducted to the point that
the humerus was held perpendicular to the
body axis, then retraction of the distal part of
the limb would be accomplished by medial
rotation of the humerus and not by retraction
a t the shoulder joint. This is, in fact, the major
propulsive mechanism in the subterranean
locomotion of moles (Yalden, '66a). In a n t -
361
eaters, however, the situation is not so ex-
treme. While the elbows often are widely
abducted, this rarely occurs to the point where
the humerus is held perpendicular to the body
axis. (This statement is based primarily on
personal observation of captive animals.) In
Tamandua, then, retraction of the distal part
of the limb usually results from a combination
of retraction and medial rotation of the
humerus, while protraction of the distal part
of the limb would result from a combination
of protraction and lateral rotation of the
humerus.
In situations where the elbows are not held
out from the body (i.e., where the humerus is
maintained in, or close to, a parasagittal
plane), axial rotation of the humerus would
then produce side-to-side movements of the
hands. In tearing apart an object such as a log,
the capability for powerful side-to-sideas well
as fore- and -aft movements could be of con-
siderable benefit in that it would enable the
animal to apply forces in a variety of di-
rections so that the object may be pulled apart
in the direction of least resistance.
Thus, axial rotation of the humerus is of
probable importance to Tamandua both in
locomotion and in using the forelimbs to ex-
pose insect prey. The question arises as to
what type of rotation (medial, lateral, or both)
occurs habitually under loading. The elbow
joint of Tamandua lacks any obvious bony
modifications (e.g., flanges on the humeral
trochlea and on the trochlear notch of the
ulna) which would aid in transmitting axial
torques from the humerus to the forearm (see
Jenkins, '73, for a general discussion of the
modifications of mammalian humero-ulnar
joint surfaces which relate to asymmetrical
distributions of forces across the elbow).
Therefore, if large torques do occur in Taman-
dua, then their transmission from an axially
rotating humerus must be mediated by a mus-
cular rather than a bony mechanism. Such a
mechanism may be provided by anconeus
(ANCN: fig. 8C) and epitrochleo-anconeus
(EPAN). These muscles are oriented such that
they would tend to pull the olecranon either
laterally (anconeus) or medially (epitrochleo-
anconeus), and the result would be adduction
or abduction, respectively, of the forearm.
These muscles are relatively much larger in
Tamandua than in Didelphis (figs. 6A,D,
7A,C, and table 1). It is conceivable that in
Tamandua, by contracting isometrically,
these muscles could stabilize the elbow and
362
SER
Fig. 12 Deep extrinsic scapular muscles, lateral aspect: (A) Didelphis, scapula in place; (B) Didelphis,
scapula removed; (C) Tamandua, scapula in place; (D) Tamandua, scapula removed.
thus transmit axial torque from the humerus
to the forearm. The possibility that selection
occurred for muscular reinforcement of this
joint instead of bony reinforcement also
implies selection for increase in overall mo-
bility. The range of abduction-adduction of
the (flexed) forearm may be increased in
Tamandua by facilitating side-to-side devia-
tions a t the elbow in addition to axial rotation
of the humerus, but this question remains to
be thoroughly investigated.
Rotation of scapula
The scapula can rotate in two directions; in
defining these directions, I follow the termi-
nology of English ('76). In anterior rotation,
the vertebral border moves anteriorly on the
thorax, and the glenoid fossa thus moves
downward and caudaily. Posterior rotation is
the opposite.
The origin of rhomboideus in Tamandua
(figs. 7C, 12C,D) is extended caudally along
the dorsal midline much further than it is in
Didelphis (figs. 7A, 12A,B). As a result, there
is a greater difference in direction of the most
BRUCE K. TAYLOR
SER
RH
S ER
anterior as opposed to the most posterior fi-
bers. The anterior rotators of the scapula are
shown schematically in figures 13A,B. In the
opossum (a), rhomboideus inserts along a very
short vertebral border of the scapula (seen
here in medial view). The arrows labeled RH
represent the lines of action of its most anteri-
or and most posterior portions; the entire mus-
cle pulls the vertebral end of the scapula an-
terodorsally. In Tamandua (fig. 13B), the fi-
bers of rhomboideus insert tangentially along
a longer vertebral border which approximates
an arc of a circle; thus, the entire muscle
tends to produce anterior scapular rotation,
such that the caudal angle is moved dorsally
and the vertebral border is moved forward.
In anteaters, levator scapulae is differenti-
ated into anterior and posterior divisions. The
anterior division (ALS: fig. 12D) in Taman-
dua forms a flat sheet which passes pos-
terodorsally to insert (fig. 3D) on the dorsal
part of the flattened area on the medial aspect
of the indistinct cranial angle. The distal part
of this division is sandwiched between rhom-
boideus medially and the posterior division of
 FORELIMB ANATOMY IN TAMANDUA 363

RH
RH

ATRP
SER

C
Fig. 13 Mechanics of scapular rotation: (A) anterior rotation in Didelphis; (B) Anterior rotation in
Tamandua; (C) posterior rotation in Didelphis; (d) posterior rotation in Tamandua.

levator scapulae (PLS: fig. 12D) laterally. The
posterior division consists of a flat sheet
which passes anterodorsally to insert just be-
low the anterior division. There is some blend-
ing of fibers between the two divisions. In the
opossum, levator scapulae (LS: fig. 12B) con-
sists of five slips which blend with serratus
magnus and insert on the vertebral border of
the scapula (fig. 3B). The action of levator
scapulae in Didelphis would be to pull the ver-
tebral end of the scapula anteroventrally (fig.
13A). In Tamandua (fig. 13B), anterior rota-
tion could be assisted by both the anterior
(ALS) and posterior (PLS) divisions of levator
scapulae, which together tend to pull the cra-
nial angle ventrally. The resulting rotation
forces the glenoid (and thus the entire limb)
ventrally and caudally. Tamandua is likely to
utilize anterior scapular rotation under condi-
tions of moderate to heavy loading during
climbing, striking a t an adversary, and also
during any tearing motions in which the hand
is pulled caudally.
The muscles which produce posterior scapu-
lar rotation are acromiotrapezius and serratus
magnus. Figure 13C shows schematically the
disposition of these muscles in Didelphis. The
acromial portion of trapezius (ATRP) pulls
the acromion anterodorsally, while the poste-
rior segments of serratus magnus (SER:
heavy arrows) pull the caudal angle postero-
ventrally. The result is posterior scapular ro-
tation such that the glenoid fossa arcs anteri-
orly and dorsally. It is important to note that
the more anterior segments of serratus mag-
nus (SER: light arrows) are not oriented such
that they would function as scapular rotators;
their primary function (aided in this case by
levator scapulae and the remainder of ser-
ratus magnus) is in body support. The impor-
tant modification in Tamandua is in the na-
ture of the insertions of the individual slips of
364 BRUCE K. TAYLOR

serratus magnus. In most mammals, t h e most for applying traction with t h e claw allows
posterior slip inserts a t t h e caudal angle, and considerable maneuverability and effective
the more anterior slips insert progressively up reach. Thus, t h e claw can engage a n object
and forward along the vertebral border (e.g., powerfully regardless of limb position.
Didelphis, fig. 12B). In Tamandua, however, Capability for elbow flexion in Tamandua is
the orientation is reversed; i.e., t h e morepos- enhanced by proximal migration of the hu-
terior slips are t h e ones which extend their in- meral attachments of brachioradialis and ex-
sertions progressively up t h e vertebral border tensor carpi radialis, and by enlargement of
from the caudal angle (fig. 12D).The result of these muscles. Likewise, shoulder retraction
this difference is shown diagrammatically in is enhanced by enlargement and distal migra-
figure 13D. All of t h e slips of serratus (SER) tion of t h e insertion of teres major and by lat-
now tend to be oriented at a tangent to t h e eral and ventral migration of t h e origin of la-
scapular border, and therefore all of these tissimus dorsi.
slips (and not just t h e most caudal ones as in The capacity for powerful flexion of the dig-
Didelphis) would tend to produce posterior its in T a m a n d u a is considered here to be
rotation of the scapula. As before, this move- adapted for such activities as tearing apart in-
ment would also be aided by t h e acromial por- sect nests or insect-infested wood, during
tion of trapezius (ATRP). The result would be which t h e claws may be hooked behind a por-
protraction and elevation of the glenoid, tion of material in order to pull i t away. It is
which would presumably assume importance predicted t h a t t h e traction applied by the
in Tamandua during climbing and also during claws is often augmented by flexion of the
defensive posturing, in which t h e forelimbs elbow and wrist, and by retraction of the
are protracted, abducted, and elevated. shoulder; these movements would bring the
claws, and thus t h e material being torn away,
CONCLUSIONS AND BEHAVIORAL PREDICTIONS
closer to t h e body of t h e animal. If necessary, a
From t h e morphology of t h e digital joints in variety of side-to side or twisting movements
Tamandua, i t may be predicted t h a t they a r e may serve to further loosen t h e portion being
subject to severe side loading in all joint posi- torn away.
tions, as would be expected if the hand were T a m a n d u a appears to compensate for t h e
twisted or deviated from side to side during loss of medial triceps as a n elbow extensor by
flexion of the claw against resistance. Expec- moderately increasing t h e size of t h e remain-
tations t h a t such movements do occur are ing extensors. Extension of t h e elbow plays a n
based upon capacity to produce powerful pro- important role in quadrupedal walking. There
nation-supination as well as other rotary and are also some modifications relating to in-
side-to-side movements of t h e limb. These ob- creased power and mobility in protraction of
servations correlate with those of Yalden the humerus; such capability is probably uti-
('66b, '721, which indicate wide ranges of lized in climbing and in defensive posturing.
radio-ulnar deviation and axial rotation in t h e The morphology of t h e forearm suggests
carpus. t h a t T a m a n d u a is capable of powerful prona-
The grasping ability of t h e hand is increased tion and supination of t h e radius. Most of t h e
by the development of t h e palmar pad (against elbow flexors produce simultaneous supina-
which t h e claws may be opposed) and by t h e tion; together these two movements would
wide range of flexion of t h e metacarpo-pha- impart a scooping action to t h e hand as i t is
langeal joint. It may be predicted t h a t such brought toward t h e body.
grasping ability would be extensively utilized The axial rotators of t h e humerus are well
both in climbing and in tearing functions. developed in Tamandua. Anteaters walk with
The triceps musculature in T a m a n d u a has their elbows somewhat abducted, and humeral
been rearranged such t h a t t h e medial head axial rotation therefore plays a n increased
assists flexor digitorum profundus in flexion role in protraction and retraction of t h e distal
of t h e distal inter-phalangeal joints; t h e fact portions of t h e limb during locomotion. It is
that its tendon passes through or close to t h e predicted that rotation of t h e humerus is also
axis of t h e elbow joint permits simultaneous important in producing side-to-side deviations
flexion of t h a t joint and the claws. The em- of t h e manus during tearing functions, but t h e
phasis upon flexion of t h e distal inter-pha- mechanism for transmitting large torques
langeal joint as t h e primary mechanism through t h e elbow is not well understood.
 FORELIMB ANATOMY IN TAMANDUA
Tamandua exhibits increased capabilities
for powerful anterior and posterior rotation of
the scapula, primarily due to increased tan-
gential orientation of rhomboideus, serratus
magnus, and levator scapulae with respect to
the border of the scapula. Scapular rotation is
probably used primarily to supplement pro-
traction and retraction a t the gleno-humeral
joint.
ACKNOWLEDGMENTS
I wish to thank the following people for
reviewing preliminary drafts of the manu-
script: H. R. Barghusen, S. W. Herring, R. E.
Lombard, and L. B. Radinsky. Their comments
and suggestions were most helpful. The assist-
ance of J. Hives with some of the illustrations
is greatly appreciated. I also wish to thank the
following institutions and their curators for
the loan of specimens in their collections:
Field Museum of Natural History, Chicago;
Museum of Vertebrate Zoology, Berkeley; Na-
tional Museum of Natural History, Smithson-
ian Institution, Washington; and the Lincoln
Park Zoo, Chicago. D. A. Meritt of the Lincoln
Park Zoo was also particularly helpful in
arranging for behavioral observations of cap-
tive animals. Most of the work reported herein
was done in partial fulfillment of the require-
ments for a Ph.D. in the Department of Anat-
omy, University of Chicago; the remainder
was done during a postdoctoral appointment
in t h e Department of Anatomy, University of
Illinois a t the Medical Center, Chicago. This
research was supported by institutional funds
from the University of Chicago and by a
fellowship from the Field Museum Center for
Graduate Studies in Systematic Zoology and
Paleontology.
365
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 PLATE 1

EXPLANATION O F FIGURES

14 Articular surfaces of head of metacarpal and base of proximal phalanx in digit 111
of Tamandua, stereo pair.

15 Articular surfaces of head of proximal phalanx and base of middle phalanx in digit
111 of Tamandua, stereo pair.

16 Articular surfaces of head of middle phalanx and base of distal phalanx in digit 111
of Tamandua, stereo pair.

366
FORELIMB ANATOMY IN TAMANDCIA PLATE 1
Bruce K. Taylor

14

15

16

367