NOTES ON THE LIMB MYOLOGY OF PROCYON

CANCRIIVORUS AND OF THE URSIDE. By

BERTRAMC. A. WINDLE, M.A., M.D. (Dubl.), Professor of
Anatorny in the Queen'8 College, Birmingham.
THE following notes on the myology of the limbs of Procyon
were made in the course of a series of dissections of the muscular
system of the Carnivora. As I have been able to compare my
observations on this animal with the accounts given by other
writers1 of the myology of the forms generally regarded as
standing nearest to it, the subject seemed to be sufficiently
complete to warrant a short note of the group.
I have abstained from any lengthy comments or references,
partly from a desire to make this paper as brief as possible, and
partly because I hope to publish in time a more detailed
account of the musculature of the Carnivora, in which this note
will be embodied.
Muscles of Back and Shoulder.-The dorsi-humeral panni-
culus is very extensive. It commences posteriorly on either
side by a slip attached to the fascia covering the root of the tail,
covers the latissimus dorsi, passes round its outer border, and is
there continuous with the abdominal panniculus, forming with it
the brachio-lateral subcutaneous muscle of some authors The
combined sheet is inserted into the humerus under the deepest
stratum of the pectoral muscle. A somewhat similar arrange-
ment is found in Ursus. Trapezius consists of two portions-
the first or occipito-nuchal arising from the occipital bone and
upper part of the ligamentum nuchae, in close relation to the
lower part; passing backwards it is joined by the levator
claviculae arising from the mastoid process. Below the point
of junction is a transverse tendinous intersection, underneath
which, but lying in the fascia and unconnected with any muscle,
1 (1) "Short Notes on the Myology of the American Black Bear (Urnus Ameri-
can)," by Professor J. Shepherd, Jour. of Anat. and Phys., vol. xviii p. 103;
(2) " On the Myology of the Limbs of the Kinkajou (Cercoleptes caudivolvus)," by
J. Beswick-Perrin, Proc. Zool. Soc., 1871, p. 547; (3) "Notes on the Myology
of the Coati-mondi (Nasua narica aud N. fumca)," by H. W. Mackintosh, B.A.,
Proc. Roy. Irish Acad., vol. ii. ser. ii., No. 1, p. 48.
VOL XXIII. (N.S. VOL III.) F
82 PROFESSOR BERTRAM C. A. WINDLE.

is a minute spicule of bone, the clavicle. Below this, the muscle

which corresponds here to the clavicular part of the deltoid is
attached to the humerus, in common with part of the pectoral,
at the deltoid tubercle. The lower portion of trapezius, which
is connected with the remainder of the lig. nuche and some of
the dorsal vertebrae, is inserted as usual and forms one single
sheet. This portion at its outer border is joined by the levator
scapulae, which arises from the anterior surface of the transverse
process of the atlas, passes under the first part of the trapezius
and separates it from the second. In Ursus the anterior part
of the trapezius receives a slip from sterno-mastoid, levator
claviculae joins it on its deep surface at the tendinous intersec-
tion, and levator scapulae is quite distinct from the second part
of trapezius at their insertion. In Cercoleptes the levator
claviculhe arises from the digastric groove, and is inserted into
the rudimentary clavicle and into the anterior part of trapezius.
The lower trapezii of opposite sides are connected with one
another by an aponeurosis but have no bony attachment. In
Nasua the trapezius consists of three separate portions- scapu-
laris superior and inferior and clavicularis, which in N. fusca
is joined by cleido-mastoid and is attached to clavicle, brachilais
anticus and humerus. Latissimus dorsi consists of two parts,
an anterior which is much the larger, gives off dorsi-epitroch-
lear and is inserted as usual, and a posterior connected with the
formeronly at its origin, which joins the dorsi-humeral muscle
and is inserted with it. In Ursus there is not this division, but
the muscle gives off dorsi-epitrochlearis and a slip to the dorsi-
humeral. In Cercoleptes one part is joined with teres major
and inserted as usual, a second joined by part of dorsi-humeralis
is connected with the pectorals, and from the former comes
dorsi-epitrochlearis, which receives slips from dorsi-humeralis
and teres major. The connection between dorsi-epitrochlearis
and teres major is present in Nasua narica, wanting in N. fusca.
The pectoral muscles consist of (1) a portion arising from the
manubrium and part of gladiolus, which expands as it passes
outwards, and is inserted in close connection with the superficial
part of the deltoid; (2) a deeper narrow portion, arising from
the manubrium and first costal cartilage and inserted into
the capsule of the shoulder-joint; and (3) a portion arising from
 THE LIMB MYOLOGY OF PROCYON CANCRIVORUS. 83
the whole of the sternum below 1, and from the costal carti-
lages and inserted into the shaft of the humerus below 2. The
abdomino-humeral panniculus, which arises from the whole of
the anterior part of the abdomen, lies close below this last part
but unconnected with it, and is inserted with dorsi humeralis and
the smaller part of latissimus dorsi beneath it. A Shepherd
describes in Ursus a muscle similar to 2 as P. minor, but it
does not correspond with the muscle known by that name in
human anatomy, but with one occasionally present in man, and
called pectoralis minimum; 1 and 3 are joined inseparably at
their origin in Ursus but soon divide. In Cercoleptes 2 is
wanting, 1 arises from anterior half of sternum, and 2-7 ribs,
and 3 beneath it from 2-7 ribs, passing to the great tuberosity
of the humerus, the capsule of the shoulder-joint, and the
coracoid process. A lower portion arises from 7-10 ribs meso-
and xiphi- sternum, and is inserted with dorsi humeralis. Nasua
has 2, "presterno-humeralis," 1 arises from the whole of the
sternum, and 3 deeper from side of sternum opposite first six
ribs. The lower abdominal part, P. quartus, arises in this animal,
from the cartilages of the lower ribs and the fascia over them.
Deltoid consists of the superficial part already described, and a
deep, quite separate, acromio-humeral. This portion joins the
superficial in Ursus. The occipital and other rhomboids form
a single muscle, as is the case in Cercoleptes and Nasua.
Shepherd describes R. major and minor as in man, but
MacalisterI says that Ursus has an occipital rhomboid also.
Levator anguli scapulae and serratus magnus f6irm a single
muscular sheet in all.
Muscles of the Am-m.-There is only a coraco-brachialis brevis
which commences by a very long tendon, and biceps has no
short bead. In Nasua the condition of these muscles is similar.
Ursus and Cercoleptes have coraco-brachialis brevis and longus
and a two-headed biceps. Supra- and infra- spinatus, teres
minor, and subscapularis have no special points of interest.
Teres minor is unsegmented from infraspinatus in Ursus and
Cercoleptes. Teres major gives a slip to dorsi-epitrochlearis in
Ursus and Nasua.
Triceps consists of longus arising in the usual situation,
1 Morphology of Vertebrates, p. 262.
84 PROFESSOR BERTRAM C. A. WINDLE.

externus arising from the neck of the humerus, internus arising

from below the insertion of coraco-brachialis for a short distance,
and anconeus arising beneath these but separate from them,
and also having an attachment to the external condyle. In
Ursus, longus arises from the whole of the axillary border of the
scapula, and anconeus from the back of the external condyle, a
fourth head of triceps arising just above it and joining the
common tendon. Anconeus is unsegmented from internus in
Cercoleptes. In Nanu nariax the anconei are united to biceps;
in N. f"sca anconeus internus arises from the supra-condyloid
process.
Mwscles ofthe FoYre-Arn and'Hand.-Pronator radii teres has
a single origin and normal insertion. In Ursus and Nasua narica
this last emends to the end of the radius. Palmaris longus
commences as a single muscle but soon divides; the larger part
going over the annular ligament becomes expanded, and ends in
slips for the three central digits; the smaller part is inserted into
the pisiform. In Ursus this muscle is absent. Flexor carpi
ulnaris is double, one portion coming from the common tendon,
the second from the inner side of the olecranon, and both being
inserted into the pisiform. There is a similar arrangement in
Ursus. In Cercoleptes Mr Perrin describes a muscle as "palmaris
longus internus," which arises from the fascia covering F. carpi
ulnaris. This is doubtless a second part of the last-mentioned
muscle; it is supplied by the ulnar nerve, as he points out. Flexor
sublimis digitorum consists of two parts--() smaller, arises from
the common tendon, crosses the deeper part, and joins that part
of F. profundus which goes to polled; (2) arises beneath the
first from the common tendon, gives off from its deep sur-
face a tendon which joins F. profundus in the palm, and then
breaks up into slender perforated tendons for the three central
digits. In Ursus the muscle divides into three slips:-(1) gives
perforated tendons to index, medium, and annularis; (2) gives
perforated tendons to minimus, and a slip to pisiform; (3)
passes to pollex. In Cercoleptes the muscle divides into two
lateral portions, which join profundus, and a mesial which splits
into four tendons,-one deep to profundus, three superficial
perforated for the three central digits. In Nasua, four per-
forated tendons are given to the four inner digits. In connec-
 THE LIMB MYOLOGY OF PROCYON CANCRIVORUS. 85
tion with this muscle must be mentioned a small muscle arising
ln Procyon from the annular ligament and pisiform bone, and
inserted into the sheath of the flexor profundus, but not per-
forated by it, at the base of the proximal phalanx. A similar
muscle is described in Cercoleptes, but is perforated by the deep
flexor. In these four forms we have inr the relations of the
superficial flexor of minimus a very interesting series of stages
of regression, as I read the facts. In Nasua the perforated
tendon comes as usual from sublimis; in Ursus it does so also,
but gives a slip to pisiform as well. In Cercoleptes the proximal
portion of the muscle has disappeared, but the distal arising from
the pisiform is still perforated. Finally, in Procyon it no longer
has any perforation, but is only connected with the sheath of
the tendon of the deeper muscle.
Flexor profundus, besides the portions mentioned above, re-
ceives a perfectly distinct adjunct, arising with but separate
from sublimis; this lies between the two muscles joining the
deeper in the palm of the hand. The great mass of the deep
muscle arises in two portions, which subsequently unite from
the front of the radius of the ulna. Ursus also has a portion
arising from the internal condyle. Pronator quadratus covers
two-thirds of the length of the bones of the fore-arm in Procyon
and Cercoleptes; in the others it is disposed as in man.
The superficial extensor of the digits arises with the first part
of the deep, and passes to the four inner digits. The deep ex-
tensor consists of two parts; (1) arises with the superficial, and
lies on its ulnar side for a short distance, it then passes beneath
it and ends in tendons for minimus, annularis, and medius;
(2) arises from the proximal part of the radial border of the
ulnar, and ends in two tendons, one of which goes to index, the
second representing extensor secundi pollicis. Extensor ossis
metacarpi pollicis is a very strong muscle. In the other animals,
as to the deep extensor, in Ursus 1 is much larger than the
superficial muscle, and 2 goes only to polled, index having no
deep extensor. Na&usa narica has separate extensores minimi,
annularis et medii, and indicis et pollicis. In Nasnua fuWca
medius gets a slip from both parts of the deep extensor.
Supinator brevis has no ulnar attachment, but arises from the
orbicular ligament of the radius. In Ursus it arises also from
86 PROFESSOR BERTRAM C. A. WINDLE.

the external condyle, but not from the ulna. In the hand
Procyon has superficial adductors for pollex, index, and
minimus, as have the other animals. Pollex has abductor,
opponens, and flexor brevis; minimus has an abductor, and
flexor brevis. The three central digits have each a pair of
interossei.
Mwsles of the Thigh. Gluteus maximus is thin but fairly
extensive, arising by a thin, strong aponeurosis from the greater
part of the iliac crest, and being inserted as usual. Gluteus
medius is very large and strong, and arises from the upper two-
thirds of the outer surface of the ilium. It is more or less
continuous with pyriformis in Cercoleptes. Gluteus minimus
consists of two imperfectly separated portions; the anterior,
which I look upon as gluteus quartus, arises from the anterior
edge of the ilium below sartorius and tensor vaginae femoris,
the posterior and smaller part taking origin from the lower part
of the ilium below medius; the latter is partly inserted with
the former and partly beneath it to the great trochanter.
Quartus and minimum are fused in Nasua narica; there is no
mention of the former in the other forms. Biceps femoris
arises altogether from the tuber ischii, and soon expands into a
large sheet of muscle, which is attached (1) to the whole of the
fascia of the thigh; (2) to the outer side of the patella; (3) to
the front of the tibia by an aponeurosis more tendinous in appear-
ance than the rest of its connections; and (4) to the fascia of
the leg nearly as far down as the ankle. In Ursus it is also
inserted by a round tendon into the os calcis, and there is an
accessories. In Cercoleptes, biceps has a caudal head. In
Nasua narica, accessories arises with agitator caudea, and is in-
serted with biceps; in Nasuqa fwsca it arises with the caudal
head of semitendinosus, and is inserted into the middle of the
fibula
Semitendinosus arises by two heads-one from the tuber ischii,
the other from the upper caudal vertebra; it is a small muscle,
and is inserted beneath the gracilis. Ursus has only the ischial
head, Nasua has both; in Cercoleptes the muscle is large and
bicipital, agitator caudae arising with its caudal head, and is
fused with it as far as the junction with the ischial portion;
Nasua has a separate agitator caudse; but this muscle was not
 THE LIMB MYOLOGY OF PROCYON CANCRIVORUS. 87
present in Procyon,' unless represented by the caudal head of
semitendinosus.
Semimembranosus arises from the tuberosity and adjacent part
of the ramus of the ischium, and is very closely connected with,
though separate from, the adductor sheet of muscle; it is in-
serted (1) into the inner side of the tibia, beneath the internal
lateral ligament, and (2) into the inner condyle of the femur.
In Ursus it is described as consisting of two parts, one inserted
into the tibia, the other and larger joining the adductor.
Sartorius is a large muscle, arising from below the anterior
superior spine of the ilium, and being chiefly inserted by a
strong aponeurosis into the anterior and inner parts of the
patella. It also sends down a tongue-like process of muscular
fibres on the inner side of the patella, the tendon of which is
attached to the tibia above that of the gracilis. Tensor vagina
femoris is a strong and separate muscle attached as usual.
Gracilis forms a thin but extensive sheet arising from the rami
of the pubes and ischium, and thus curving round the other
muscles above; it is inserted by a flat tendon into the middle
third of the anterior surface of the tibia. Pectineus arises from
the very prominent ileo-pectineal ridge, and is inserted as usual.
The remaining adductors are two in number, (1) a small muscle,
arising partly beneath the pectineus and partly internal to it
from the front of the pubes as far as the symphisis, it is in-
serted behind the pectineus; (2) an exceedingly large sheet of
muscle, arising under cover of the gracilis from nearly the whole
of the descending ramus of the pubes and the anterior part of
its body, and from the ramus of the ischium, and inserted
into the posterior part of the femur from a short distance below
the lesser trochanter to the lower end, and into the internal
condyle. Besides this last-mentioned muscle, Shepherd describes
a small adductor " a narrow ribbon-shaped muscle, which arises
from the tuberosity of the ischium in common with the semi-
membranosus, and is inserted into the upper part of the internal
condyle of the femur and adductor tubercle; a little before its
insertion it joins the greater portion." I am inclined to look
1 Macalister, Morph. of Vert. Animals, p. 263, says that the Procyonide have a
separate agitator caudle muscle, but it did not exist as an isolated muscle on
either side in my specimen.
88 PROFESSOR BERTRAM C. A. WINDLE.
upon this muscle as more properly belonging to the semimem-
branosus than to the adductor, though, of course, the connection
between these two muscles is very intimate. Cercoleptes and
Nasua have each three adductores. Quadratus femoris is a
large and quite separate muscle.
There is a strong psoas parvus in all the forms, and ilio-psoas
presents no special points of interest. The same remark applies
to the two obturator muscles; the gemelli are very large and
distinct. Pyriformis is also well developed and quite distinct,
some of its fibres arising outside the pelvis from the transverse
processes of the caudal vertebrae. There is a single-headed
rectus, and the remaining parts of the quadriceps are as usual.
Mwseles of the Leg and Foot.-The two heads of gastro-
cnemius are separable in the greater part of the leg, there is no
separate plantaris, and soleus has only a fibular origin. The
combined tendon is partly inserted into the lower portion of the
posterior part of the os calcis, a bursa intervening between the
tendon and the bone. The superficial part of the tendon, how-
ever, which is here separated from the deep by a second bursa,
passes over the os calcis, forming a kind of hood-shaped aponeu-
rosis, from which in the sole of the foot arises the short flexor
of the toes. Ursus is described by Shepherd as having a three-
headed gastrocnemius, the outer of which he looks upon as
representing plantaris; all three are attached only to the os
calcis. Cercoleptes has a large plantaris, the tendon of which
passes into the sole of the foot, forms the plantar fascia, and is
closely associated with the flexor brevis, a similar condition
existing in Nasua. Popliteus is a large, strong muscle.
Flexor tibialis arises from the upper third of the tibia, from
the head of the.fibula, and from the fascia overlying tibialis
posticus. Flexor fibularis, which is much larger, arises from the
fibula and adjacent fascia. The tendons of these muscles unite
in the sole, and give slips to the four outer toes. Flexor acces-
sorius arises from the inner side of the os calcis, turns round this
bone into the sole of the foot, crosses the combined tendons
of the long flexors with which it is slightly connected, and
passes joined by a slip from the combined tendon to the hallux.
From its side and from the combined flexor tendon spring two
small muscular bellies, which end in slender tendons passing
 THE LIMB MYOLOGY OF PROCYON CANCRIVORUS. 89
severally to those of the flexor brevis which go to the 2nd and 3rd
digits. Flexor brevis arises, as has been stated, from that part
of the tendo Achillis which passes over the heel, and gives per-
forated tendons to the four outer toes. As regards the flexor
group in the other forms, accessories in Ursus passes directly
into the combined tendon; in Cercoleptes, brevis supplies tendons
only to 2, 3, and 4 digits, each of these tendons receiving a fleshy
slip from accessories, the remainder of which joins the combined
tendon. Tibialis posticus has a small muscular belly and a
long tendon inserted as usual. Extensor longus digitorum
gives tendons only to the three outer toes; extensor brevis,
which is a well-developed muscle, gives tendons to the four
inner. Extensor longus hallucis is smalL Peroneus longus is,
as usual; the tendon of brevis passes behind that of longus, runs
along the side of the foot, and is inserted into the base of the
middle phalanx of the fifth toe. There is no separate peroneus
quinti. This last muscle is present in all the other forms.
Tibialis anticus is a large muscle.
Hallux has an' adductor, abductor, and flexor brevis. The
first muscle is described by Perrin in Cercoleptes as " obliquus
tarsi" Minimus has adductor, flexor brevis, and an abductor
inserted into the spur at the base of the metatarsal bone. Each
of the other digits has a pair of interossei.