YYOLOOP OF THE PECTOBAL BEOJON IN PIBHEB, 196

8. Further Observations on the Myology of the Pectoral

Region in Fishes. By E. W. SEAKN, B.Sc., Ph.D.,
F.Z.S., Biology Master at Onndle School.
(Text-figures 1-4.)
[Reeeired October a%,lB@: Red Febraary 19,1994.1

I. Introduction.
11. The Myology of the Peetmal Fkgion in Lepidorirem and Polyptsnu.
1. The L n t d Muscle.
8. The Pectoral oirdla
3. The Pectoral Piu.
111. The Cleithro-byoideus Muscle in the Teleoebi.
1V. The Dugre- of Homolopy Iwtweeu the bfuacles of the Pectoral Region in
various orderd of FiFishes oud in Urodele BntrucLiaus (with Key).

I. IXTBODDCTIDN.
The following observatione are supplemental to two previous
pnpera on the myology of fishes: the first, *‘Ont he Nature of
the Iaternl Muscle in Teleostei” ( P m . Zool. Soc. 1914), paved
the way for the second, “The Comparative Myology of the
Shoulder Girdle and Pectoral Fin of Fishes” (Trans. Roy. Soc.
Edinburgh, 1919). The notes and drawings for this latter work
mere made before the war, and at the date of their publication I
had neither the time nor the opportunity fo complete the
programme which I had undertaken. The Dipnoi were left
untouched, and though flgures of Polyptems were drawn they
were not described. Moreover, beyond a Key to t h e comparative
myology, no general conclusions were arrived at. The monograph,
however, incomplete a8 it admittedly was, led to interesting
correspondence with other workers both nt home and abroad.
Stimulated by this, and at the instance of my friend Prof.
D. M. 8. Watson, F.R.S.,F.Z.S., on the fir& available oppor-
tnnity I resumed my studies at the point where they had heen
relinquished. I am indebted t o Prof. Graham Ken-, F.R.Y.,
F.Z.S., for the specimens of Lepidosiren and Polypterm upon
which my observations are based.

OF TEE PECTORAL
II. THE MYOLOGY IN LEPIDOSIREN
REGION
AND P O L Y P T E l i W .

1. The Lateral IIInscle.

The composition of the lateral muscle of the other Osteichthes
conforms closely with that of the Teleostei, coi~sistingof four
series of hollow cones, two above aud two below the lateral
h e . I n the pectord region foLir areas can still be traced ;. of
13*
196 DB. E. W. 8HANN ON TEE lifYOuMY OF

these, the two dorsal are completely severed from the two ventral
by the passage of the horizontal septum which runs from beneath
the lateral line to the vertebra There are no “red fibres” in
connection with the lateral line.

LEPIDOSIBEN.
I n Lepidoeiren several peculiarities of the lateral muscle are
worthy of special consideration.
The general arrangement of the lateral muscle behind the
pectoral girdle closely resembles that of Polypteruu, except for
the greater thickngss of the ventral body-wall. This resemblance
may be’ due i n part to the fusiform shape of the body which is
shared by both form& . A t the same time it diffem widely from
the condition which has been described in a fusiform Teleestean,
namely, Conger. In these presumably primitive forms the
myomeres remain more nearly i n juxtaposition to the vertebm
which bear them; or, i n other words, they form very much
shallower cones than are found in the Teleost myomeres. The
dorsal posteriorly directad cones, large in Yolyptem~s,in Lepido-
eireii so far preponderate as entirely to exclude the smaller
anteriorly directed cones from appearing at the eurface. Owing
t o the great thickness of the abdominal wall the conical structure
of the ventral moiety is retained almost as far forward as the
pectoral girdle. A very thick fascia separates the dorsal from
tlie ventral moiety.
The superficial divisions of the lateral muscle of Lepidosiren are
illustrated in text-fig. 1. I n the side view (A) the 1atero-dorh;il
portion is seen as a massive muscle whose fibres slope rrttlier
steeply from below anteriorly to above posteriorly ; it bas a very
extended origin upon the whole side of the skull as far forward
as the eye. The ventral moiety anterior to the tip of the
pectoral fin is divided by distinct fascise into superticial areas as
well as into layers. The superficial areaa are numbered similarly
i n each of the drawings A, 13, and C ; of these areas 4 is con-
sidered to be the latero-vential portion, the remainder derivatives
of the mesio-ventral portion.
On makinga mid-ventral longitudinal incision (as in C,mn. to 6)
a distinct fascia is found completely separating a thin outer layer
from a thick inner layer. Following the incision por;teriorly the
fxscia is found to disappear in the 6th myomere behind the base
of the pectoral fin; here the fibres of 5 become confluent with
those of 4. Dissecting from the mid-ventral line to the riglit
( i .e. left of the animal) and then upwards the whole of 1, 1 a, 2,
and 5 come away re:dily from t h e underlying part, 3. B shows
tlie appearance of the left aspect after the superficial muscle
h y e r I n s been removed ; veritr:illy nre sern the cut edges of 1 u,
2: and 5. I have not encoiuitered this continuity of muscle from
the mesio-ventral portion t o t4e mandible in any other ~ I O U I of J
fishes.
 THE PECTORAL REOIONJN FIBHIB. 197

Part 3 is a massive muwle trtking origin on the hyoid and

passing back beneath the pechntl fin t o be inserted in prt 4;
the pectornl girdle is embedded in this m u d e (where it is shown
by transparency in B), but there is no fascia to delimit a

Text-figure 1.
+a.

.L.

Lateral Muscle of Lepidoriwn (X $1.

A. Left aspect, superficial. B. Left blapect, deeper layer.
C. Ventral aspect, suprficial.
Br.o.,blanchid aperture ; Hy., hgoid ; L.D., latero-dorsal pr3rtion ; L.Z., latern1
line; Xn., mandible; P.f.,pectoral tin; P.g., pectoral girdle; T.,t o n p .
1, la,2,3,4,6, parts of the vontlal moiety of the lateral muscle.

cleithro-hyoideus such as is found in the vast niaj’o~tyof

Osteichthes. The parts 3 of either side are separated by a thin
vertical septum near their origin, and a little further back
become widely separated by the pericardium, of which they form
198 DE. E. W. SEBhh” OH THE MYOLOQY OF

the lateral walls. Part 4 overlies part 3 to a certain extent, and

its free edge can be disected away from the portion of 3 which
underlies it M f a r as the wavy line shown in B ; at this level the
fibres of parts 3 and 4 become intermixed. By the 7th myomere
behind the base of the pectoral fin (one behind the tip of the fin)
all trace of areas and layers i s lost.
Parts 1, 1a, and, to a great extent, 5, serve to depress the
mandible. Part 3 serves to retract the hyoid and thus lower
the tongue. Part 2 serves as a constrictor of the pharynx ; its
deeper fibres, especially in the forward area, run obliquely to the
snperficial ones (as shown in C) ; thus itr power of enfolding the
pharyngeal cavity ia greatly increased.

PoLYmns.
Owing to the relative straightness of the myocommata in this
form there is little external evidence upon which to define a
mesio-dorsal from a latero-doiwl portion. The greater bulk of
the dorsal moiety runs forward beneath the pectoral girdle
to h k e origin on the occiput. In the pectoral region the grenter
part of the muscle, judging by the direction of its fibres, appeals
to belong to the latero-dorsal portion.
There is also below the lateral line very little distinction
betwcen the Iatero-ventral and the mesio-ventral portions.
Internally, in the dorsal moiety, the upper posteriorly directed
cones are so large as almost to obliterate the lower ones. Owing
to the position of the anus, the conical structure of the ventral
moiety is practically non-existent; it is composed almost through-
out its length of a single layer covering the abdominal cavity.

2. The Pectoral Glirdle.

LEPIWSIREN.
The great reduction of the skeletal element^ of the shoulder-
girdle exhibited by hpidosiren resulte in considerable departure
from the normal distribution of the shoulder-mnscles. No skeletal
element occurs above the mticnlar cartilage of the fin, the stid
cartilage being connected to the basi-occipital by a strong liga-
ment (such as occurs in many Teleosts). No part of the dorsal
moiety of the lateral muscle contributes to the fixation mus-
culature of the girdle. The ventral sympliysis, formed here by
the clavicles, is embedded in the deeper layer of the mesio-ventral
portion, as was noted above. I n consequence of this it makes no
appearance on the ventral surface, as is the mse in almost all
other fishes ; nnd, as a further consequence, there is no retractor
mesio-ventralis pectomlis inferior, nor m y muscular connection
between the g i d l e and the mandible. The general relations of
the girdle to the lateral muscle are shown in text-fig. 1, while
 THE PEi?”)RAL REGION IN FI8He8. 199

text-fig. 2 illustrates the diapoaitiona of t h e smaller mudes.

Hrimphry* has worked out hhe myology of Cerdodus, and tha
synonyms c i a below refer to his observations on that type.

Text-Egure 2.

Shoulder Muscles of Lcpidos(+en (X 1).

Leh asp&, after removal of constrictor museleg aud almost tlta eirtiw me&-
veutrnl portion of left side. A., articular cartilagr, !,&owing iuriertiolr Of
p.1.v.p.) ; Ey.e., ceratorllal (cut) ; Lig., ligament; U.,lataral liue; T., toiigue
For other letters see text.

(1) Tlu Pootm’or Xueclca.

(a) lhrsal abmnt.
(6) Retractor l&ero-ventralis pectomlk.
(c) Retractor mesio-ventralis pectoralis.
(1)) R&wctor Zatero-ventralis yectorm?is (r.I.-v.p.) ; synon. Ser-
ratux, Hurnplrry :-
The muscle is p r t of that numbered 4 in text-fig. I. It is a
thin sheet tmversed by myocommrrta, and is attached to the
psterior border of the articulni cartilage, as well (IS spreadi~lg,
to a slight extent, upon the adductor m u ~ c l ofs the fin. Only the
dorsal tibres of the Iatero-ventral portion (muscle 4)are inserted
on the girdle, nnd may therefore be mid to constitute the retractor;
the muscle sprsads ventrally to the finover muscle 3, but it is no
longer inserted on the girdle.
(c) Retrllclor ~sio-ventrahkpectardir (r.m.-v.p.) ; a p o n .
Pechrnlh ( m d Latissimus dorsi ?), Hrimpliry.
By reference to text-fig. 1. €3 it will be wmn th& the who16
girdle of either side is embedded in a mass of muscle numbered 3.
The mass ioderived from the m e ~ h - v e n t l dportion of the lateral
mu6cle, though not comprising the whole of that portion, and
those f i b m which a m attached the posterior border of the
girdle must function as retractors. At the same time there is no
fascia to ’clktingriinh n definite retractor muwle from those p r h
of the same maw which run forward, exterualiy to the girdle, to
take origin on the ceratohyal.

* “The Muscles of CcraWdrr,” Jonru. An&. and Phys. pi., 187%

200 DU. E. W. SHANK ON TEE XYOLOOY OF

The r.m.-v.p. tliiw conatifuted take8 origin fmm the wltole

posterioi. border of the girdle, underlying at its upper extremity
the r.l.-v.p. descrilnd nbove.
The homology of this muscle is somewhat obsciire, hiit, i t
appwently represents in pFrt the r.m.-v.p. snp. m d med. of other
fishes. The r.m.-v.p. iuf. is undoubtedly wanting in Lepidoaiveia.

( 2 ) The A~JerabrMuecles.
(a) Dorsal wmting.
(b) Protimtor Ititero-vent~nlispectoinlis.
(c) Muscles derived f i ~ m the nic&-wntF;il portion :
(i.) Levator pectoidis.
(ii .) Ul~itjirolhrnnchides.
(iii.) Claviculo-hyoideiis.
(h) ~ ~ o t r n c t olatero-uentrdi8
r peclorulis (p.l.-v.p.) ; Synon.
Tmpexiiis, Yuulphry :-
This is a. suid1 muscle which runs from its origin on the h i -
occipital, parallel to the l i g ~ m e nmentioned
t ;riiove, to its iiisertiori
on the anterior outer border of the articular cartilage. lty
teildiilous insertion is in pnrt continuous with that of the r.1.-v.p.
over the outer surface of the airticultw cartilage ; further, since
t h e fibres of the two muscles also take the same direction, there
C L I I be little doubt that they belorig to the mame portion of the
lateral musclg.
(c) Muscles derived from the mesio-ventml portion :
(i.) 1;evcstor pectoralia (1.p.) ; synon. Sterno-cleidomastoideus,
Humphry :-
This muscle is represented by R very slender pla.t,e of fihres
wliich forms the posterior rim of the brnnchial cha.mber. From
its origin on tlie bnsi-occipitd, imtiieili~tely henenth the pro-
tmctor, it runs almost vertically downwards to he iiiserted on the
iutier border of the cleithrurn. Some of the nnterior fibres,
bending sharply forward nmr their insertion, extend beneath the
origins of the cleithro-branchialus.
(ii.} Cleithro-branchiah ext:ternua (c1i.-hr.ext.) :-
A small, biit well-defined, strap of muscle taking origin from
the inner upper surface of the cleithriim and pawing forwrirdfi to
he inserted on the lower posterior rim of the second brancliirrl
cnrtil~ge. Beneath this lies n smaller, hiit nevertheless distinct,
muscle wliich runs to the lower posterior rim of the third
branchial cartilage; this constitutes tlie cleithro-branchidis
internus.
The cleithro-branchiales thus differ in their mode of imertion
from t h e condition to be ohserved in nll other g r o u p of t h e
Osteichthes. This may be due in p r t to the extreme reduction
of the brnnchial skeleton in general and of thc two posterior
arches in particular ; at the same time it suggests an arrangement
 TnE PN‘TORAL REGION IN FISTIES. 201

moro nearly akin t o t h e segmental plan of the generalised fish

type such RS occurs in existing Elasmobmnchs.
(iii.) Chvimdo-hyoidkm (cla.-hy.) ; synon. Coraco-hyoideus,
Humphry :-
Taking origin on the swollen ventral extremity of the c e m
tobyal, the whole of whose posterior circumference it grips, a
stout muscle riins directly backwards, expanding laterally as it
goes. This muscle is illustrated in text-fig. 1, where it is denomi-
nated 3, and in the context its insertion is fully discussed. The
innermost fibres run to the pectoral girdle where they are
inserted into its outer surface, both into the clavicle and the
cleithrum. “hese fibres constitute the claviculo-hyoideus, as
shown in text-tig. 2 (ch.Ry.); it is clear, however, that they are
i n no way delimited from the remaining fibres of muscle 3,
nlthough they are separated by a tolerably distinct fascia from
their fellows of the opposite side.
The clavico-hyoideus of Lep’dosiren i s not readily comparable
with t h a t of tlie Chrondrostei or with the cleithro-hyoideus of
other Oateichthes. I n few other fish known t o me does any
portion of the muscle which runs from the hyoid arch to t h e
pectoral girdle pnss that structure and become directly confluent
with the ventrsl hody-wall. The condition is certainly not to be
regarded as primitive, for i t finds no parnllel in t h e Chondrichthes
On the other hand, it mnnot entirely he nttributed to the reduc-
tion of the slioulder-girdle, t h e lower Iialf of which (especially
the ventral symphysis) is not nearly so reduced as is the case in
many other Osteichthes. W e have here, in fnct, an approach to
the condition in the Amphibin, whei-e muscles completely invest
t h e ventml aspect of the girrlle, so that none of its bkeletal parts
nppenr beneath the skin (with the exception of the newly
inteicdated sternal elements).

POLYPTERUS.
(1) Th Posterior Mwrcles.
((6) Retractor dorsnlis pectoralis.
( 6 ) Retmrtor latero-ventidis pectoralis.
(c) Retrnctor meaio-rexitnlis pectoralis :
(i.) superior.
(ii.) medins.
(iii.) inferior.
(a) Rctrector rloi*scclispectoralis (R.d .p.) :-
The p e n t rnam of the dors:tl moiety r u n s forward beneath the
girdle to be inserted on the skull, but n few of the superfieid
fibres rlre inserted into the inner aspect of the supraclavicle aid,
to a less extent, of the cleithrum. These fibres constitute a
retractor mnscle ; and since there is no distinct division between
the mesio-dorsal and latero-dorsal portions, it seems well t o
designate i t simply “ r e t r a c t ~ rdorsalis pectoralis.” It is illus-
202 DR. E. W. SBANK ON THE MYOLOQY OF

trated in text-fig. 3, R, but the main point of inwrtiou is higher

up than the terminrrtion of the index line.
(b) Retractor latero-ventraliopectoralis (r.1.-v.p) :-
A very few fibres from the latero-ventral portion succeed in
reaching the ghdle immediately below the lateral line, where
they are attached to the inner aspect of the lower postclavicle.
(c) Betractor ntesio-uei~rulis
pectorulio :
(i.) Superior (R.m.-v.p.sup.) :-
This muscle is clerrly shown in text-fig. 3 , B ; it is not attached,
ns might appear to be the case, t o the lower postclavicle, but rum
beneath that bone td inserted on the inner aspect'of the
cleithrum.
(ii.) Hedim :-
Does not appear in either of the illustrations, but it may be
defined 8~ that section of the mesio-ventral portion which occupies
the axil of the fin. It ends in the fascia which covers the
nddnctor muscles of the fin. It is not clearly defined, however,
either from the '' superior" or the " inferior " section.
(iii.) Inferior (R.m.-v.p.inf.) :-
As is seen in text-fig. 3,A, this Hi the only one of the series of
retractors which is not traversed by mywornmata right up to its
point of insertion. The muscle is thin and is attached throughout
its depth to the posterior rim of the cleithrum. The R.m.-v.p.inf.
of either side are separated from one another by a marked septum
in the mid-ventral line.

( 2 ) The Anterior Nwcks.

(a C a) Protractor pectoralis.
(i.) Anterior.
(ii.) Posterior.
(c) Muscles derived from the masio-ventral portion :
(i.) Levator pectoralis.
(ii.) Cleithro-branchiales.
(iii.) Cleithro-hyoideus.
(n & b) P~otruciol.pectoralis anterior (P.p.xnt.) :-
This is a short compact muscle running from the otic region of
the skull to the inner aspect of the supraclavicle ; it is seen in
text-fig. 3, B. Beneath it lies a smaller muscle which I propose
to a l l the protractor pectoralis posterior.
From the nature of their origins and insertions as well RS from
their functions, these two muscles appear to be homologous with
the sirnilas pnir which occur in all Osteichthes. At the snme
time it is worthy of note that they nppaar, from their position
high above the lateral line, to he derivativesof the dorsal moiety
(a)wther than of the latero-ventral portion ( b ) ; in the absence
 THE PECTORAL REaION IN FISHER. 203

Pectornl musculature of Polyptsrur ( X 1).

A. Ventrnl.nspset,after removal of constrictors. U. Left Intern1 anpet.
Br.mr., Urnncliio-msnnibulnna ; Cli., Cleithrum ; Hy.c., CeratollyaL,
M u d e a as iu text.
204 DR. 1. W. SIIANN ON THE HYOLOQY OF

of developmentnl evidence, however, i t does not appear to be

jwtifiable to alter the nomericlature on this account.
(c) Muscles derived from the mesio-ventral portion :
(i.) Levator pectovalis :-
A cylindrical muscle running from the basioccipital to the inner
upper aspect of the cleithrum. It does not appear in text-fig. 3, B,
being hidden by the suprabranchial constrictors ; it runs paralld
with the ligament (shown in the illustration as a white streak
immediately above the constrictors), and just ventral to it.
By reason of its direction it functions rather as an additional
protractor than as a levator, but, from its general topography,
there c m be little d?+t of its homology with the levator
pectoralis of Teloosteans.' -
(ii.) CZeiOro-branchiales(C1i.-br.) :-
I find three distinct parts of this muscle which, arising close
together on the inner aspect of the cleithrum, run as follows (see
text-fig. 3, B):-
(a) Externus, a,short broad strap riinning almost vertimlly to
the last ceratobranchial, just below the bend of that structure.
(b) 17~ternu.s~ a cyiindrical muscle taking origin inside the
above and running obliquely forwards to the ventral extremity
of the last ceratobranchial.
(c). Tertius, a smaller cylindrical muscle lying between the two
f osegoing.
I have lately had some correspondence with Prof. D. M. S.
Watson about the cleithro-branchiales of Polyptern. Prof.
Watson found that on one side of his specimen this muscle
formed a single continuous sheet, while on the other it was
divided into two distinct bundles. I n the two specimens of
P. senegalus dissected by me, and on both sides of them, I foiind
the same arraugement of the cleitbro-branchiales which I have
already illustrated and described. I therefore propose to retain
my original description, though it is quite possible that a certain
amount of variation occurs.
(iii.) Cleithro-hyoideus (c1o.-hy.) :-
A powerful muscle, devoid of myocommata, which, arising from
the anterior face of the lower third of the cleithrum, runs for-
w;wds to be inserted on the bme of the hyoid arch. It is epclosed
ventrally b the clavicle, but sends no fibres to that structure.
In the miLventrd line it is contiguous with its fellow of the
opposite side, from which, however, it is separated by a fascia
throughout its depth.

3. The Pectoral Fin.

LEPIDOSIRXN.
Owing to the extreme reduction of the pectoral fin in Lepiclo-
siren the muscles are excessively small and simple. The adductor
 TEE PECTORAL REQION IN FIBHE8. 205

on the inner surface and the abductor on the outer each consists
of R single compact bunch of fibres which, taking origin 011 the
artioulltr cartilage, is inserted on the bawl mrti'tctge of the fin.
I can find no trace of muscular tissue between the segments of
the fin-axis.
POLYPTERW8.
(1) Adductor.
(i.) Adductor mperf;ci.li. (Adds.) :-
A narrow strap-like muscle which takes origin on the anterior
border of the scspuls, and, in part also, of the comcoid. It runs
Text-figure 4.
Add.? Di1.p.
I
! 5 3 ,Pt. cla.

I
I
I \

Mbt. Ahd.s. &I.&

Iiiner a9prct of left peetors1 fin of Polypterns ( x 2).
Cli., Cleithrurn ; Cor., Coracoid ; dlpf., Metnpteryginm ; Pt.clu., I'ostclnviule (the
indpx liue runs to the upper) ; Se., Scapula. Muscles as i i i test.
' obliquely downwards to be inserted on the mehpterygi:rl ratlial,
giving off a few fibres to the inuer border of the inetqiterygiuur.
This muscle functions as R dilator rather than R S an nrlductor ;
but it evidently corresponds to the d d s . of Teleostea~ie,which,
in maiiy cases, is also very s m d l and only partially covers the
a~~a.p.
(ii.) Adductor profundtzs (Add.p.) :-
The largest muscle of the adductor series, arises st the junction
of the scapula and coracoid, and spreads fan-wise, gctthering fibres
from the fascia covering t.he Dil.p.ii., from the mesoptyrypi:il
cartilage, and from the metnpterygium, till i t covers the wliole
distal area of the inner surface of the fin. A t their extreniitly
tlie fibres pass into a tendinous sheet which is iuserted on tlie
b:ises of the lepidotrichia. Some of the deeper fibres of this
muscle run obliquely (about 60 degrees) to those =en oii the
surface; there is no fascia, however, between the two sets of
fibres.
(iii.) Dilator posterior (Di1.p.) :-
From the fact that this muscle takes origin behind the glenoid
border it evidently belongs to the adductor series. It agrees
206 DR. E. W. SHANN ON THE M Y O L W Y OF

i n position, moreover, with the dilator posterior which occurs in

many Teleosteans, but difFers from any such known to me in
that it is divided into two prts (dwignated i. and ii. in text-
%-4).
Part i. is a triangular muscle which takes origin oh the inner
aspect of the cleithrum immediately above the centre of the
scapula, whence its fibres run obliquely downwards to their
insertion on the upper border of the propterygium; a number
of the dorsalmost fibres, however, become collected into R
tendon which, after forming the upper rim of the fin muscu-
lature, becomes inserted on the propterygial radial. !l%kmuscle
is also partly visible from t h e outside of the fin, as appear8
in text-fig. 3, B. .
Part ii. has the shape o f a - flattened o n e which, taking origin
at its base on the upper posterior border of the scapula, diveb
beneath the adductor profundus to its insertion along the inner
border of the propterygium.
These muscleu serve to keep the upper supports of the fin b u t :
and so to dilate the fin, during the back stroke induced by the
adductor profundus.

( 2 ) Abductor.
(i.) Abdmtor super-cia& (Abd.sup.) :-
This and the other muscles of the abductor seiiea are illus-
trated in text-tig. 3. Abdsup. is a thin sheet of muscle having
an exteusive origin on the outer aspect of the cleithrum. Owing
to the fact that its fibres are inserted on the fascia which covers
the nbductor profundus, only a >mall portion of the muscle,
and this in the dorsal region, reaches the distal area of the
fin ; no part of it, however, comes in direct contact with the fin
skeleton.

(ii.) Abductor profundus (8bd.prof.) :-

Arising from the outer angle between the scapula and the
cleithrum this Lmge muscle spreads fan-wise, being augmented by
contributions from the outer aspects of the propterygium, of the
mesopterygial cnrtitage, and of the metapterygium. It ends in n
tendinous sheet which is inserted on the bases of the lepidotricliia.
As in the case of the adductor profundus, some of the deeper
fibres run obliquely to those seen on the surface ; again, however,
these is no fascia between the two sets of fibres.

(iii.) DiZutor antei*ior (Di1.a.) :-

This muscle takes the form of a slender cone which, arising a t
its basal end from the outer aspect of the cora.coid, runs along the
outer ventral border of the metfipterygium on which its fibres are
inserted. It forius the ventral border of the fin musculature, and
is p % l yvisible from the inner aspect (see text-fig. 4) its well as
from the outer.
 THE PECTORU REGION IN FIBHFS. 207

111. TIIECLEITHBO-HYOIDBUS MUSCLEIN THE TELEOBTEI.

CleiJmhyoidarLs. Bynon. 8terno-hyoid, Stannins, Vogt &
l'ung ; Retractor hyoidei, Owen ; Hyo-p&oralie, McMurrich ;
Hyo-clavicuhrb, Hamburger :-
A powerful muscle which forms the ventral W e r of the gill-
chamber. It has (L double origin on the- cleithrum; the larger
portion arises from .the outer aspect of t h e chithrum, usually i n
its lower third, while a smaller portion arises from the region of
the symphysis, where it comes in contact with its fellow of the
opposite fide. The insertion mcum on the b e of the hyoid arch.
The muacle is usually much reduced in biilk as it approaches
tile point oE insertion, but Zeus forms an exception.
I n a few of the more syecitrlised Teleostettns a part of t h e
cleithro-hyoideus runs from its insertion on the hyoid external
to the cleitliivm so as to become confluent with the muwle of the
ventral body-wall (i.e. Retractor mesio-ventralis pectoralis in-
ferior). This recalls the condition which we have observed in
Lepidosiren ; but in no Teleostean, does the unattached outer
portion of the muscle so far preponderate over the cleithro-
hyoideus proper as iY the wse in the Dipnoon. Speciahation i n
this respect hue apparently taken place along vaxious lines, and
those which have come to my notice in the courw of these investi-
gations will now be considered.
I n most cases t h e ventral symphpsis of the two cleithra is
vieible on removing the skin. In T2Gglu gurnardwr a cylindrical
muscle runs on either aide of the ventral middle line from t h e
urohgsl to the symphysisof the cleithw. A t the same time,
another portion of the cleithro-hyoideus runs from the outer
posterior border of the brucihyrtl to t h e inner aspect of the
cleithriim at the level of the origin of the aleithro-branchialis
internus. Between these t a o portions a thifd is inserted on t h e
hsihyal and runs clear of the girdle, between the pectoral and
pelvic fins (where i t narrows considerably), to become coefluent
with t h e ventral body-wall. All three portions are separated
from one another by fnscis, and none show any trace of myo-
commata. A precisely Bimilar condition wm found in Peri-
OJddUdWlUS.
In Bhnnius pholk the general disposition of the cleithro-
hyoideus is similar to that i n Trigla, but here the ventral portion
of the cleithro-hyoideus proper is much more in evidence than
the upper outer portion. I n Motella mwtelct, on the other hand,
conditions are reversed. The sympliysis of the c1eithi.a is scarcely
visible, the ventral portion of t h e cleithro-hyoidens proper is
reduced to a mere thread, and the outstanding featuw on the
ventral aspect is the large free portion. This upper portion
differs from that in the foregoing types in that it arises on t h e
level of the cleithro-brauchialis exterrius and is iuserted on the
upper aspects of the uro- and basihyal; there is, moreover, no
fascia behveen it and the free portion.
208 DR. E. W. IIHAPI" ON THE MYOLOUT OF

In Pholie gunnellue, the symphysis is wmpletely h!&den by a

muscle which runs direct from the urohyal to the iudimenbry
pelvic girdle. A large portion of the ventral muscle runs forward
above the foregoing to be inserted on the posterior aspect of the
basihyal ; this represents the free portion of the cleithro-byoideus,
but differs from that of the above types in that it is traversed by
niyocommata. Above this again, but not separated from it by a
definite fascia, is the cleithm-hyoideus proper.

Iv. TEEDEQREES
OF HOMOIAX+Y
BETWEEN TEE MUSCLESOF THE
PECTORAL
RWIONi n various Orders of Fishes, and in
Urodele Batrachians.
(References as in op. cit., and not as in this communication.)
Attention waa d m t n in my previous pnyer (we Introduction)
to the chaotic condition of the myologicnl nomenclature of the
pectoral region in Fishes, prior to my work. After perusal of the
text, and particularly of the Keys to the muwulature in Ela!nio-
bwnchii and Teleostei, my observation mill be fount1 to be justified.
In adopting a uniform system of nomenclnture i t w a s ~iecessnry
to select one of two mnin lines of procedure: ( a ) to retain tlie
existing names for muscles (giving precetlence, in cases whew
two or more names had been applied t o the same muscle, either
to priority or to suitability), or (b) to iiitroduce new names.
After due consideration I decided to follow the latter course.
This haa led, in some cases, to the coinage of such cumbemonie
terminology aa appears to demand an exp1an:ttion ; the explana-
tion, moreover, involves the discussion of a wider subject, namely,
the morphological value of the new nomenclature.
As a result of the investigations on the laternl muscie which
were set forth in my previous papers, it was shown that a veiy
considerable degree of similarity in the general build of the body
exists in all the main Orders of living fishes. I n the embryonic
state the similarity is very much more marked than in the adult ;
but, even in the adult, at least behiud the pectoral girdle, the lateral
muscle retains a decided constancy of structure. The origixi:il
nietameric segmentation of the body muscle is maintained ; and
although the individual myomeres are bent into c o n i d form they
are still separated (right up to the pectoral girdle or, at all events,
close up to i t ) by myocommatrt. The division of the lateral
muscle by a horizontal septum benearth the lateral line, giving
rise to a dorsal and a ventral moiety, is R constant morphological
f;ictor. Of more doubtful value, however, is the further sub-
division of each moiety iuto a latewl and a niesial portion. It
has beeu shown, for instance, that though the conical con-
formation of the lateral and mesial portions itl fairly constant
throughout the Chondrichthes on the one hand, and the
Osteiclithes on the other, yet between these t w o Snb-CIRsses
certJiu discrepancies exist. The, diviaion of the moieties
 THE PEC'TORAL R E G 1 0 9 IN L'IRHES. w
into lateid and niesiltl portions is at best not very clearly
defined, especially in the forward :wea with which me are
chiefly concerned, nnd oue cimiiot affirm that i t litis any t,riie
morphological valne. At the smue time, its use for descrip-
tive purposes is beyond doubt, for it divides the lateral muscle
behind the girdle into groups whose probable liomolom it mould
be dificult to deny (see Key a t end of section). Though the term
'6 latero-dorsd " is merely descriptive of relative position, for
example, the term " Retrnctor latero-dorsalis pectoralis " is
believed to imply a Iiomologous muscle in the various Orders
in which it, occurs. The fact that the majority of the muscles
attached to the pectoral girdle of fishes are merely fixation
muscles, and not, concerned with its actual movement, gives them
a greater constancy of arrangement than is found in the higher
Vertebratn. This constancy of arrangement first suggested to
me the nniform system of nomenclature which I wns led
to adopt.
The special mosculn.t,ureof the pectoial girdle and fin in any
given fish develops as n series of outgrowths from a definite
number of myomeres, thongli, as Goodrich (38) has shown, the
number of myomerer; which comnience the contribution in the
embryonic stage may be conr;iderably in advance of those which
a,ctually form the miiaciilature of the arlult (a certain definite
proportion of the oi.igina1 muscle-buds heing discn.rder1 in the
process of construction). The fact that muscles nre developed
from different embryonic myomeres in different groups of fishes
does not necessarily imply that they are not homologous. A s an
extreme exnmple of the argument one may cite the n~uscles
of the pelvic fin in Teleostei; the homology of these niuscles
throughout the Order has never been questioned, and yet they
may be developed from inyomeres many places removetl (c. 9. in the
Salmon and the Cod). Granted that liomologous muscles are not
necessarily derived from numerically homologous myomeres, i t
follows eqiially that we cannot look for them to be innervated by
iiumerically homologous nerves. Thus, for the purposes of a
general inyological classification one is confined, to a large extent,
to considerations of disposition and arrangement of parts. But
there are degrees of homology, and the closer the relationship i t
isdesired to show between the myology of any two types, t h e
closer must he the attention paid to myomeric development and
to innervation, both embrjonic and adult.
The homologies suggested by similar names in my classification
are of the broad general type which I have specified ahove. As
a rough gauge of the degree of homology which is implied,
1 should suggest that i t is comparable to that which exists
hetween the pectoral niyology of the Batrachie and the Reptilia,
hut it is not so close as that which exists between any two Orders
of those Classes respectively. I regam1 the hiatus between living
Fishes and Amphibians in respect of their pectoral musculn.ture
P R O C . ZOOL.Soc.-1924, No. XIV. 14
210 DR. E. W. IIHAPI" ON THE MYOLOUT OF

as greater than that which exists between the latter and any other
Class of Tetrapods. For this rewon I have avoided the use of
terms which are nserl in connection with Bntrachiaii inyology.
A t the same time, I liave drawn attention to possible homologies
which may exist between the inuscles of the pertoral girdle in
Fishes and certain of those exhibited by Urorlele Ratrachims.
Bearing in mind the foregoing proviso, we will proceed briefly
to summarise the evidence collected on the various muscles of the
pectoral region (cf. Key to the Comparative Myology of the
Pectoral Region, at end of section),

(1) The Posterior Muscles.

(a) Retrwctor lutero-ckPrsnlis pectoralis (R.1A.p.) :-
Usually small and only slightly diferentinted. I n Bntoids,
however, it is well developed and devoid of myocommnta. As a
rille i t is certain thRt the inesio-dorsal portion makes no con-
tribution townrds its formation ; the Holocepliali, however, form
a n exception to this stntement, nod for this reason the niiiscle
lins been nained in that Order simply Retractor clor.srtlispectoralis
(R.tl.p.). I n the Holocephali, moreover, this and the succeeding
iniiscles of the posterior series are more highly differentiated
tiinn in any other Order of living fishes.
It is possible that this mriscle, after being rotated into a more
vertical position, became the foreriinneiu of t h e Lztissimns clorsi
of Urodeles. The origin from the dorsal element of the shonldcr-
girdle and the insertion on the clorsal portion of the body-
mnscnlattture seem to snggeht something deeper than a merely
mecl~anic~l resemblmce between these two muscles.
(b) Ilstwctor latero-ventralis pectoralis (R. 1.-v.p.) :-
Usually small and slightly differentiated. Rettei developed in
the Chondrichthes than in the Oateichthcs. Attains its highest
development in the Holocephnli, where it is divided into two
distinct sheets (externus and internus).
This musc!e has been thought by Hnmphry and others to be
homologous with the SerratLis of Urodeles ; a snggestion which,
from the relative positions and proportionate sizes of the two
muscles in qnestion, seems quite possible.
( c ) Retractor mesio-ae?ttralispectoralis (1t.m.-v.p.).
(i.) S'wpei-ior :-
Usually rt large musvle which may or may not cover the
foregoing. Except in the Holocephali, i t is traversed by
myocommata close np to its origin on the girdle.
This niiiscle, especially RS represented by the Osteiclithes,
forms the lateral body-wall, and, almost undoubtedly, gave rise
to the Obliquus of Urodeles. It is not actually divided into two
superimposed layers, but, owing to retaining a trace of its conical
formation, the deeper fibres ran obliquely to the sriperficinl.
 THE PEC'TORAL R E G 1 0 9 IN L'IRHES. 211

(ii.) Medius :-
Only attains considerable development in the Chondrichthes,
while in all Orders of the Oateichthes it is greatly reclnced, and
in the Chondrostei absent.
The R.m.v.p.med. is not represented in living members of the
Rstrachia. Jts presence depends upon the occurence of a very
wide glenoid border, such as occurs i n primitive fishes, but is
gri~duallylost in the higher forms, while in living Batrachians i t
is reduced to a ball-and-socket joint.
(iii.) I22ferior :-
Is almost invariably large and well defined, arid frequently
loses its inyocomma.ta before reaching the girdle. The curious
exoeption presented hy Lepidosiren is discussed in section 11. of
this paper.
This muscle forms the ventral body-wdl, and Owen regarded
its resemblance to the Rectus abdominis of the Ratrachin as one
of the surest liomologies between the inyology of that Clnss and
of the Fishes.

(2) The dwterior itlziscles.

(a) Protrccctor lcctero-dorsalis pectorulis (P.l.d p . ) :-
Is undeniably represented only in Rhino and the Rays. I n
the Holocephali n very definite protractor occurs in the region
of the dorsal moiety; hiit, as i t appears to he derived from t h e
mesio-dorsal portion (in pnrt) as well as from the latero-dorsal, it
has heen named Protractor clorsalis pectoralis (P.d.p.), CJ the
corresponding retractor in the Holocephali.
It beeins unlikely that a muscle wliicli only appears in certain
specialised Chonricl~tl~oirl~ should have a counterpart in the
Batracliia. Kevertheless, the muscle dehignated " 22 " by Owen
(73,fig. 140) corresponds as closely in position with the P.1.4.p.
as does that clesipated '' 11 " (=latissimus dorsi) with the
R.1.4.p.
(h) Protractor kctero-ventrdis pectoralis (P.l.-v.p.) :-
The muscle is clearly defined throughout the Elasmohranchii,
hut is wanting in the Holocephali.
In the Osteichthes t h e muscle, which I have there named
Protractor pectoralis (P.P.), is divided with marked eonstnncy
throughout the Orders into two closely apposed portions (anterior
and posterior). A question a t once arises as to whether these
two portions are not homologous respectively with the Protractors
Iatero-dorsalis and latero-ventralis pectoralis of Elasniobranchs.
This supposition, if correct, would simplify the classification ; but
the evidence is against its acceptance. In the Dipnoi and
Chondrostei both protractors undouhtedly belong to the ventral
moiety of the lateral muscle. I n the Polypterini and in some
14*
212 DB. E. IT'. SHANN OX THE MYOLOQY OF

Telestoei, on the other Iiand, both protractors possibly belong to

the dorsal moiety?judging by their position high up on the body.
In the absence of developmental evidence I have not altered
the nomenclature on this accoiiiit, but I have compromised to the
extent of using the term Protractor pectoralis (omitting the
committal 6' Iatero-ventrtilis ").
The Protractor pectoralis of Osteichthes is very suggestive of
the Trrtpezins (=Protractor scnpulae, Owen) of Urodela, wliich
is also divided into two parts, although these parts in the
UrocleIe nre niore widely separat,ed and more mobile.
(c) Muscles derived from the rnesio-ventral portion.
(i.) Levtor pectorulis (L.P.) :-
Whether this muscle is actually derived from the Iatero-ventra1
or from the mosio-ventral portion would he difficult to afiriii,
aiitl for that reason i t has not been coupled with tile term '& niesio-
ventral '? in the nomenclature. I n character and level of insertion
i t seems to correspond in ninny mays with the Retractor niesio-
ventralis pectoralis superior (though always smaller in actual
size) ; i t lins therefore been rlnssi tied with the inesio-ventral
clerivntives. The iiiain point is that in all Orders, except the
(jhondrostei, a well-defiiied muscle is foniitl t o riin from t,lie
skiill to the pectoral girdle at n level decidedly lower than
t,lie prot,ractor ninscles described above. I n Osteichthes the
nlnscle is ncconipn.iiied by a. tendon, with which it may o r 1na.y
not be in actual contwt. The relations of this tendon to the
llones of the axinl skeleton n.nd of the pectoral girdle have been
cliscnssed under the heading " Teleostei."
It is possihle that the Levator ppctoralis of Fishes is the
forerunner of the Levator scapulz of Urotleles, but the latter
(lifers in its origin, which is from the cervical vertebm instead
of from the skull.
(ii.) C o r u c o - ~ ~ ~ ~ ~c o~~t ~uzZ~i~s~ z((3.-a.c.)
c ? c i s :- .
r\ specialiserl part of the ventral musculatnre only found in
Elasmobranchii.
(iii.) Co,.uco-bl.nitchicLles, etc. :-
I n all fishes a muscular connection exists between the pectoral
girdle and the branchid apparatus. There is, Iiolvever, a very
tlistinct difference in n.rrangement betweeii the Chonrlrichthes
and the Osteichthes in thiq respect.
I n the Chonrlriclithes me find a more or less distinct mnscle
bundle ronning from the coracoitl to the bxse of ench of the
bi-anchial arches. These Inindles are known as the cornco-bran-
chiales (C.br. 1-5).
I n the majority of Osteiclithes mnscle buntlles iwi from the
cleithrum to the posterior branchid n.rch only. These are known
as the Cleithro-branchirlles (CLbr.). I n Chondrostei there is
b u t one such bundle. In Teleostei there are two, extenins and
 TI1E PECTOHAL REGIOS IS FISIIES. 213
illternus (the iiisertion of tlie latter is interim to tliat of tlie
foriiier, and its origin is 1iigher.up on tlie cleitlirum). I n Poly-
pteriiii a third cleitliro-briliichialis occiirs between tlie externus
:dthe internas.
In Lepiclosire?~we find n somewlint iiiteriiiediate coiidition
I,etweeii the Clioiidriclitlies :uid the reuiniiiiiig Osteichthes. There
are two cleithro-br;tncliiales, aiid these are inserted on the second
and third branrhial cartilages respect,ively,
This element in the pectoral rnnscnlature of Fishes lins no
counterpart in the adult Urodele.

(iv.) (!oraco-Agoitlezis, etc. :-

In all fishes a pair of contiguous uiuscles, oiie oii eitlier side of
the median plane, runs from the pectoral girdle to the base of tlie
liyoid arch. The muscles are nilrued according to tlio skeletal
element of the girdle from which they arise : thus, in the Clkon-
drichthes we have a Uoraco-hyoideuti (C.-hy.), in Dipiioi and
Uliondrostei a Claviculo-liyoideus (Ula.-liy.), aiid in Polypterini
aiid Teleostei n Cleitliro-hyoideiis (CL-liy.).
Tile coraco-hyoideus of Elasiiiobraiicliii tliKers sliglitly from
tliat of IIolocepliali in that it arises froin the coraco-arcual
septuin instead of from the coracoid cii,ltilage itself.
Tliere caii be littlejcloubt that tlie Sterrio-lipid of Urodeles lias
bee11 derived froin tliis eleiiient of tlie pectoral niusculatui~eof
Pislies, tliougli, as one iiriglit wpect, its relative size is iiiucli
retlucecl.
It is iiiterestiiig in tliis connection to find R “ Puboliyoideus”
(Owen) in the Salaiiiander. This muscle, iiisertecl on the hyoitl
and ruuning back iii the ventral body-wall till it reaches tlie
pelvic girdle, vividly recalls the curious coiidition observed iu
connection with the claviculo-Iiyoideus of L q d o s i ~ e ia~i d the
cleithro-liyoideus of c w b i n Teleosteaiis where :L ‘‘ free portion ”
of the muscle runs back, exteriial to tlie pectoral girdle, to
bec’onie confluent with tlie ventral body-wall.
(C-lim.) :-
(v.) Coi.ccco-Z~goincciicEibi~laris
A very specialised modification of the ventral iuusculature
found only in lfuicc of the types examineci, aiid probably confined
to the Batoids.
(vi.) ~‘orc~co-nia~~tlibu~ffii~is
(C.-iu.) :-
A muscle liigbly developed iii all C‘hoiidriclitlies. It is appa-
reiitly waiitiiig 111 the Osteiclithes with tlie possible exception of
CemtotZzcs, for which Humphry (52) hns described a ‘‘ Genio-
Coracoid.”
( 3 ) The Fiti Vzisclea.
(b) Addtwtov.
‘!Ae inner or upper iiinscuIatnre of the fin wliich serves to draw
i t ill towards the side of the fish. With tlre exception of Lepido-
214 DR. E. IV. S H A K S ON Tlik: MYOLOGT OF

aiven (ill which the fin is so far degenerate as liardly to be

conipnrahle witli t h a t of :my other of the, types selected), the
:i.tltluc!toriiiiiscle is divisible irtto two superirriposetl layeis, super-
ticidis ( Atld.sop.) mid profuiidus (AdcLprof.). The two layers
:we not very clearly defined from one another except in the
Polypterini and Teleostei.
I n I’olppterua and in certain Teleostearis there is a third
deiiieiit of tlie add iictor series, which lilts been described :IS
I )ilirtor posterior (Di1.p.).
(I)) xlhtltrctov.
TIie outer or lower iiiirscuIatrire of the fin whiclr serves to draw
iL forwards away froin&lie body. I t is divided i n a inanner pre-
cisely siiiiilar to tlie adtldctor. Thus we have an abductor super-
ficialis (Ald.sup ) niitl :tii abductor profuiidus (Abdprof.) ; in
Lepirlosiwir tliere is but oiie layer, while in l‘olyptertccr i d ill
t,liose Teleosteans wlricli possess a clil;itor posterior we have, iti
:iililitioii, ii dihtor :Interior (Di1.a.).
In rert;iiu ‘releostwiis whose pectoral fiiis Ii:ive become modified
for progression oil ;L solid srirfn.ce remarkable specialisation of the
tiorillally siiiiple iiirisculature occurs. The muscles of the pectoral
h i of ~ ! o p k i u were s described iii some detail in this connection.
Such cases, however, are rare, ant1 do riot detract from tlie valiie
of it geiieral noinenclatnre which is a.pplicable to the vast
iiiiijority of fish types.
There is n.t present no evidence upon whicli to base even the
viigiiest con1parison betweeii the myology of the fin of any know11
tisli : i i ~ t ltli:rt of the lirnb of a.ny known Batrachian. Both start
as a series of ventral bi1ds from certain definite inyomeres, hit
tlieiiceformnrd each follows its owii course in building the adult
struct,tire, and ontogeiiy otfers as yet no clue to the phylogeny of
the Tetrapod limb.
 TJlE I’KCTOIIAL IlEXIUN IS E’ISIILS. 215

I-

64
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