S6M 1035

Zoology Series

VOLUME 61

NZ
a
NATU RAL

HISTORY
MUSEUM

NUMBER 2 30 NOVEMBER 1995

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© The Natural History Museum, 1995

Zoology Series
ISSN 0968-0470 Vol. 61, No. 2, pp. 91-138

The Natural History Museum
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London SW7 5BD Issued 30 November 1995

Typeset by Ann Buchan (Typesetters), Middlesex
Printed in Great Britain at The Alden Press, Oxford

Bull. nat. Hist. Mus. Lond. (Zool.) 61(2): 91-101

Issued 30 November 1995

Preliminary studies on a mandibulohyoid

‘ligament’ and other intrabuccal conne
tissue linkages in cirrhitid, latrid and
cheilodactylid fishes (Perciformes :

Cirrhitoidei)

| PRESENTED
| GENERAL LIBRARY

~es

PETER HUMPHRY GREENWOOD‘,

Visiting Research Fellow, The Natural History Museum, Cromwell Road, London SW7 5BD and
Honorary Research Fellow, J.L.B. Smith Institute of Ichthyology, Private Bag 1015, Grahamstown,

6140 South Africa
CONTENTS
IOC Neti OMe n bois ope anbinanecact eisai seed act ae bae's vanjag. int «wo oegemaNNer bigetci Seas Pamala sss adeisnsasebeeingatun sass opbeelenigde ous 91
MIatenic AGN ICE OGSI oiiecsancma idan ape clo aianuicins pciathaniisn: «sn Wanpdsciemmaracainn tcloal-iitiueeebaisinaeissat= <piasinger aa deananeltaianad 92
Ligamentous and other connective tissue linkages between the mandible, the palatoquadrate, the hyoid arch
ANG the ODELCHiaE SCLIES II (ALEC CIUUMITOIG FANTIISS oo. cis reas ap aiineinieisceiaiaasicinanl sjnuinine Sete Mansdes hae ease sane ee a dere. aes 93
Whe tanagly Cue Dae yo oe cscs ssi nciasienha steerer eee Melita ds a cas oa catemeNalastN cieemnernNUM NEN daetwesaNieNUeeN Ty seis rsewe sracntipes 93
SU CM ANETLLV MIE ALENCL AC RUPEES. Re eS T ONCE Rts cin ncn Oy OPS Ma Peis oo ve w rice seni n bigs nse Se pagslasldivecainneeie' Gass ive acpiadsisamsicniticnis singcaens 93
SE EULAIILY © MELO AGI CA CMS mI arias fais caieasoasnae sen ih seri iden ds doeee dasines steals vokeaidas(e nisin mains ex ap esa man siceacolces(oe 97
Discussion and Conclusions .................0s0cee00 RNP a cnc SR RA te a acne ne naerias toate sue wonton tenatn hen nortnceac nee Nts: 98
PROMOS PEMEMIGKI deka PeRIa Ie do sconce conch Mahnancece¥Gitenvestepesthrtdedsedhcccdavacteatecds-sseeeteree ieteceecs. 100
IUCECHEHEES © occadeucsscstecads smcbreceascnceiatectacser asd secaaiicr sels diac seschie cs houtestiarastsesnecactesstsncaesessa: «divs aeeMemtines onc 101
Synopsis. In certain taxa of at least three groups of percomorph fishes belonging to the cirrhitoid families

Cirrhitidae and Latridae, there is a connective tissue linkage between the mandible and the hyoid arch, suggestive of
the mandibulohyoid ligament described in certain sub-percomorph groups. This ligament is generally thought to be a
feature of lower teleost fishes, although a mandibulohyoid connection has also been identified in a few more derived
taxa. The mandibulohyoid connection in the Cirrhitidae examined would appear to originate from the tendinous
aponeurosis associated with the Aw division of the adductor mandibulae muscles, but its derivation in the latrid
species Acantholatris monodactylus remains undetermined. The Aw aponeurosis in A. monodactylus, as well as in
the latrid Mendosoma, and in two genera of Cheilodactylidae (viz. Chirodactylus and Cheilodactylus) ramifies
extensively over the paladoquadrate arch and part of the opercular series. This system, together with various
intrabuccal ligaments is described from representatives of the three cirrhitoid families studied.

It is concluded that, contrary to several earlier ideas, a mandibulohyoid linkage is of taxonomically and
phylogenetically widespread occurrence in teleosts but that it might be derived from different connective tissue
sources. The value of this connective tissue complex in phylogenetic studies has yet to be established, but it appears

to be of use in at least establishing intragroup relationships within the Cirrhitoidei.

INTRODUCTION

A recent anatomical study of certain cirrhitoid fishes (sensu
Greenwood, 1995) has revealed a number of markedly differ-
ent ligament and tendon systems which separately or con-
jointly link the mandible with the hyoid arch, the
palatoquadrate arch and the opercular series. Some of these
connections have a degree of complexity not previously
recorded among teleost fishes.

Of particular functional interest are the two types of direct

+ Dr Greenwood died 3 March 1995.

© The Natural History Museum, 1995

and ligament-like connections between the ceratohyal and
mandible in some cirrhitid species, and a third type found in
one of the latrid species examined (family placement after
Greenwood, 1995). These linkages invite comparison with
the so-called mandibulohyoid ligament generally thought to
be commoner in lower teleosts than in perciform taxa, or
even restricted to the former groups (see Verraes, 1977;
Lauder & Liem, 1980; Lauder, 1982; but see also Osse, 1969,
Springer et al, 1977, and Aerts et al., 1987 for certain
perciforms, and Anker, 1974, for gasterosteiforms). A man-
dibulohyoid ligament also occurs in the semionotiform lepi-
sosteids Lepisosteus and Atractosteus (Wiley, 1976).

92

The nature of the hyoid-mandibular connection in the
cirrhitoids, a derived percomorph group (Greenwood, 1995),
and that described in other teleosts and in the Lepisosteidae,
raises doubts about the strict homology of the connection in
these various taxa.

Functionally, it would seem that a mandibulohyoid connec-
tion is of importance both in adults (see Aerts et al., 1987)
and in early larval stages (Verraes, 1977); its phylogenetic
history in this context is discussed at length by Lauder (1982).

Regrettably, the feeding studies on cirrhitoids, from which
this anatomical study arose, are not sufficiently advanced or
refined to allow informed speculation on any correlation
between the morphology and the feeding habits of these
fishes. Furthermore, many other cirrhitoid taxa remain to be
studied before it will be possible to evaluate what significance
the different types of intragroup mandibulohyoid and other
linkages (or absence thereof) may have in unravelling the
taxonomy and phyletic relationships of the group. Neverthe-
less, there are indications from this study, and from informa-
tion in the literature on various forms of mandibulohyoid
connections, that further investigations may yield insight into
the biomechanics of feeding and into historical relationships.

MATERIALS AND METHODS

Material

Clupeidae: Etrumeus terres. RUSI 34140 (1 speci-
men)

Salmonidae: Oncorhynchus mykiss RUSI 36417 (3
specimens)

Characidae: Hydrocynus vittatus RUSI 19355 (1
specimen)

Cirrhitidae: Amblycirrhitus bimacula RUSI 77-20 (3
specimens)
Cirrhitichthys oxycephalus RUSI 40526
(3 specimens)
Cirrhitops fasciatus RUSI 2375 (1 speci-
men; ex Hawaii)
Cyprinocirrhites polyactis RUSI 12339
(3 specimens)
Paracirrhites arcatus RUSI 30975 (2
specimens)
Paracirrhites forsteri RUSI 39419 (3
specimens)

Latridae: Acantholatris monodactylus RUSI

33485 (2 specimens)
Mendosoma lineatum
RUSI 33613 (1 specimen)
RUSI 33626 (1 specimen)
RUSI 26176 (1 specimen)

Cheilodactylidae: Cheilodactylus fasciatus DIFS unreg. (2

specimens)
Cheilodactylus pixi DIFS unreg. (3
specimens)
Chirodactylus brachydactylus DIFS

unreg. (3 specimens)

DIFS: Department of Ichthyology and Fisheries Science,
Rhodes University, Grahamstown. RUSI: J.L.B. Smith Insti-
tute of Ichthyology, Grahamstown.

P.H. GREENWOOD
Method

The entire opercular series, palatopterygoid arch and hyoid
arch of one side, together with the mandible, premaxilla and
maxilla of that side, were dissected away from the head.
Muscles, tendons and ligaments were examined on the dis-
sected side, and checked on the contralateral aspect which
was left in situ.

All specimens had been fixed in formalin and preserved in
either ethyl or propyl alcohol.

In the absence of ontogenetical information on the devel-
opment of ligamentous and tendinous systems in these fishes,
ligaments and tendons in the various taxa are presumed to be
homologous if their places of origin and insertion are similar.

ABBREVIATIONS FOR FIGURES

Abbreviations for tendons and ligaments are given separately
with each figure.

Muscles:

Add. mand.

Iu&m: Adductor mandibulae A, upper and main divi-
sions respectively

Aw: Aw division of adductor mandibulae muscle

Avt: Tendinous aponeurosis of adductor mandibulae
Aw

Geh: Geniohyoideus

Im: Intermandibularis

St: Sternohyoideus

Skeletal elements:

Ang: Anguloarticular
Bb: 1st basibranchial
Ch: Ceratohyal

Dt: Dentary

Cts Ectopterygoid
Ent: Entopterygoid
Epi: Epihyal

B: Raised facet for articulation with epihyal
Ga: Gill arch

Hyom Hyomandibula
Hyp: Hypohyals

Ihyl: Interhyal

lop: Interoperculum
Max: Maxilla

Mt: Metapterygoid
Pal: Palatine

Pop: Preoperculum
O: Quadrate

R: Retroarticular
Sy: Symplectic

V: Vomer

MANDIBULOHYOID CONNECTION IN CIRRHITOID FISHES

LIGAMENTOUS AND OTHER CONNECTIVE
TISSUE LINKAGES BETWEEN THE
MANDIBLE, THE PALATOQUADRATE, THE
HYOID ARCH, AND THE OPERULAR SERIES
IN THREE CIRRHITOID FAMILIES

The family Cirrhitidae

On the basis of their mandibulohyoid connections, two
distinct groups can be recognised within the cirrhitid species
examined. A third group, represented by Amblycirrhitus
bimacula (Jenkins), has no macroscopically detectable man-
dibulohyoid linkage (see below).

Group I species (viz. Cyprinocirrhites polyactis [Bleeker],
Cirrhitichthys oxycephalus [Bleeker] and Cirrhitops fasciatus
[Bennett] have a stout, ligament-like connection between the
ceratohyal of each side and the coronoid process of the
corresponding dentary ramus (Fig. 1A). Group II species
(viz. Paracirrhites arcatus [Cuvier] and P. forsteri [Schneider]
also have a ligament-like band of tissue stemming from the
lateral aspect of each ceratohyal, but here it links each hyoid
arch with, predominantly, the corresponding quadrate, on
which it inserts immediately above that bone’s process for
articulation with the anguloarticular. Part of this tissue,
however, is apparently continous with the tendinous insertion
of the Aw division of the adductor mandibulae muscle, (Fig.
1C). There is no macroscopically obvious and clearly defined
connective tissue linkage between the mandible and hyoid
arch in Amblycirrhitus bimacula (hereafter referred to as
Group III).

In all three groups the adductor mandibulae Aw division
originates on the quadrate through a posterior extension of
the muscle’s tendinous central aponeurosis, and thus is of the
basic perciform type as defined by Gosline (1986). The
extension is well-demarcated and moderately deep, and lies
across the quadrate-anguloarticular joint. Amblycirrhitus
bimacula (the single Group III species examined) is excep-
tional in this respect because the tendon lies very slightly
above the jaw articulation. Members of all three groups have
the fascia covering the Aw muscle extending posteriorly onto
the lower half of the quadrate, part of the preoperculum, and
the upper margin of the interoperculum as well.

Within the three species of Group I there are differences in
the association between the mandibulohyoid connection and
the tendon of the adductor mandibulae A, muscle inserting
on the maxilla. Cyprinocirrhites polyactis is unique in having
what appears to be a short branch of the mandibulohyoid
connection arising near the latter’s attachment to the coro-
noid process of the dentary and then joining the maxillary
tendon of the adductor mandibulae A, muscle (Fig. 1A). In
Cirrhitichthys oxycephalus and Cirrhitops fasciatus , the maxil-
lary tendon partially fuses with the mandibulohyoid connec-
tion at the point where the two cross over each other (the
latter lying medial to the maxillary tendon). From the point
of fusion a short section (interpreted as a continuation of the
maxillary tendon) runs into the tendinous central aponeurosis
of the adductor mandibulae Aw muscle (Fig. 1B).

There are also intergroup differences in other ligamentous
and tendinous linkages (Fig. 1). Species of Groups I and III
have a small upper, anterior division of the adductor man-
dibulae muscle A, inserting onto the maxilla only via the
ligamentum primordium. Group II species, in contrast, have

93

that division of the muscle inserting on the maxilla through
both the ligamentum primordium and the maxillary ligament
of adductor mandibulae A, muscle. In all three groups the
major (ie lower) division of the muscle is attached to the
ligamentum primordium and the maxillary ligament, the
latter inserting on the ventral aspect of the maxilla, and the
former on the bone’s dorsolateral aspect.

Other intergroup differences involve the epihyal-
interopercular and the interhyal-interopercular ligaments
(For comparison of these and other ligaments with the
situation in other cirrhitoid families, see pp. 94 and pp. 97-98
and Figs 2-4). Group II species have the latter ligament
partly associated with the epihyal as well as the interhyal, as
does the single Group III species dissected; in Group I taxa,
however, the ligament is confined to the interhyal. The
epihyal-interopercular ligament shows more marked inter-
group differences, especially when species of Group I are
compared with those of the other two groups, a difference
possibly associated with the manner in which the epihyal
contacts the interoperculum. In Group II taxa, the lateral
face of the epihyal head articulates with a well-defined,
prominently raised and posteriorly directed facet situated a
little below the dorsal margin of the interoperculum and
slightly behind the bone’s midpoint. The epihyal-
interopercular ligament in these fishes is short and stout,
originates on the lateral face of the epihyal near its dorsal tip,
and runs forward at approximately 45 to the sagital plane. It
inserts on the upper and anterior faces of the prominence
supporting the facet on the interoperculum against which the
epihyal articulates.A similar epihyal-interopercular ligament
occurs in the single Group III species examined, viz. Ambly-
cirrhitus bimacula. However, in this species, unlike those of
Group II, the interopercular facet is located on a relatively
lower base.

The epihyal-interopercular ligament is most distinctive in
Group II. In species of this group (unlike the other groups)
the epihyal articulates directly with the medial face of the
interoperculum and not with a facet carried on a distinct and
elevated base (albeit only slightly so in the single Group III
species examined). The ligament itself is a prominent feature
originating (as in other groups) on the dorsal tip of the
epihyal’s lateral face, from which it runs anteriorly onto the
dorsal margin of the interoperculum at a point near the
bone’s anterior tip, where it is narrowly separated from the
attachment point of the mandibulo-interopercular ligament
(cf. Acantholatris monodactylus Fig.3 & p. 94).

A short interhyal-metapterygoid ligament is present in all
three groups.

No interhyal-preopercular ligament is present in any exam-
ined species of the three groups (cf. the other cirrhitoids
described below).

A stout mandibulo-interopercular ligament (Fig. 1C; lig. 3)
is present in taxa of the three groups. It is confined to the
lateral face of both elements in all species except Cyprinocir-
rhites polyactis and Cirrhitichthys oxycephalus (both members
of Group I). In these two species it divides anteriorly to insert
on both the lateral and the medial aspects of the anguloarticu-
lar bone.

Also common to species of the three groups is a short and
deep, ventrally located ligament connecting the anguloarticu-
lar and dentary.

94
The family Latridae

There are clear-cut differences in certain aspects of the
ligamentous and other connective systems in the two latrid
species examined, namely the monotypic genus Mendosoma
lineatum (Gay) and the species Acantholatris monodactylus
(Carmichael) of that polytypic genus. Also, an upper division
of the adductor mandibulae muscle A, is absent in A.
monodactylus whereas in Mendosoma it is an elongate, rather
thin element which lies lateral to the major part of the muscle
and extends over the greater part of its length. The minor
division, unlike the major one, has no direct connection with
the ligamentum primordium and inserts on the maxilla,
together with the major division, via the maxillary tendon of
the A, muscle.

In Mendosoma the adductor mandibulae Aw division is a
very thin muscle, largely tendinous and with a single posterior
extension of its tendinous aponeurosis. This runs slightly
below the upper point of the articulation of the lower jaw
with the quadrate, to which bone it is attached a short
distance from the anterior border (Fig. 2; tendon 3). In other
words, it is of the basic percomorph type sensu Gosline
(1986), except that in Mendosoma it has one prolongation
extending along the symplectic, another running ventrally to
attach to the medial aspect of the preoperculum, a third,
directed dorsally to insert on the quadrate, and a fourth
directed obliquely backwards to attach to the ventral aspect
of the interoperculum medially and anteriorly.

In all essentials, the adductor mandibulae Aw tendon
system’s extension onto the interoperculum and preopercu-
lum in Mendosoma is very similar to that in Acantholatris,
with that in Mendosoma, as it were, foreshadowing the more
clearly differentiated condition in Acantholatris (cf. Figs 2 &
3).

There is no ligament-like connection between the mandible
and hyoid arch in Mendosoma (cf. Acantholatris; Fig. 3).

The epihyal-interopercular ligament is stout and short,
connecting the lateral aspect of the epihyal with the dorsal
margin of the interoperculum a short distance anterior to its
slightly raised facet for articulation with the epihyal. Unlike
the backward-facing facet in those cirrhitids in which it
occurs, that in M. lineatum faces forward (Fig. 2), as it does in
the other latrid examined (Acantholatris monodactylus; and
in the cheilodactylids dissected).

A discrete interhyal-interopercular ligament (present in
cirrhitids) is apparently lacking in M. lineatum (as it is also in
Acantholatris, and Cheilodactylus).

Like the two latter genera, but not in the cirrhitids exam-
ined, Mendosoma has a_ well-developed interhyal-
preopercular ligament and another, more dorsally placed
ligament between the interhyal and the metapterygoid (Fig.

P.H. GREENWOOD

2; lig. 7). This latter ligament I consider to be the homologue
of the interhyal-quadrate ligament in Cheilodactylus, and the
ligament in Acantholatris which runs from the interhyal to
both the quadrate and the entopterygoid (see p. 98 & p. 99
respectively).

Mendosoma has discrete lateral and medial divisions of the
mandibulo-interopercular ligament, with the medial division
terminating a short distance behind the anterior tip of the
interoperculum (Fig. 2; lig. 6), and the lateral division
extending much further posteriorly.

The anguloarticular-dentary ligament is short and stout,
markedly stouter than in any cirrhitid species examined, and
stouter than that in Acantholatris.

As compared with the ligamentous and other connective
tissue systems in Mendosoma lineatum, those in Acantholatris
monodactylus are considerably more complex, (as they are
when compared with the cirrhitid species studied). As was
noted earlier (p. 94), there is no obvious sub-division of the
adductor mandibulae A, muscle in A. monodactylus. How-
ever, anteriorly the upper third of the muscle, unlike the
other two-thirds, is free from the ligamentum primordium
and inserts on the maxilla only through the maxillary tendon,
to which the major part of the muscle is also attached.

Acantholatris monodactylus has a substantial Aw portion of
the adductor mandibulae muscle. From the muscle’s
mediolateral tendinous aponeurosis a stout and relatively
short branch (tendon 3 in Fig. 3) runs posteriorly to insert on
the anteromedial aspect of the preoperculum’s horizontal
limb.

A second stout and much longer tendon from the Aw
muscle (tendon 5 in Fig. 3) extends from the ventral margin
of the muscle above the anguloarticular bone, and runs
obliquely backwards to attach to the medial aspect of the
interoperculum a short distance from that bone’s anterior tip.
This tendon, unlike tendon 3, is not derived from the
aponeurosis of the adductor mandibulae Aw muscle but
originates directly from the muscle itself. Immediately after
its origin, tendon 5 is attached to the anterodorsal aspect of
the anguloarticular’s medial face. It then passes over that face
of the retroarticular, and attaches to the medial aspect of the
interoperculum a short distance from the bone’s anterior tip.
Since this tendon links the mandible with the interoperculum
it would appear to be the functional equivalent of the
mandibulo-interopercular ligament in the other species
described above. However, a true and very long mandibulo-
interopercular ligament is also present in A. monodactylus
(Fig. 3; lig. 6). Anteriorly it has an extensive attachment to
the lateral face of the anguloarticular and retroarticular
bones, as well as another on the posterior face of the
retroarticular. From here the ligament extends across to, and

Fig. 1 A: Cyprinocirrhites polyactic (Group I species) Medial aspect of the left lower jaw, cheek region and hyoid arch, viewed obliquely
from above, to show the mandibulohyoid connection (semi-schematic). The branchial skeleton is displaced to the right. About times natural
size. 1: Mandibulohyoid connection; 2a: tendon from lower part of adductor mandibulae A, muscle to maxilla; 2b: continuation of tendon
2a, joining tendinous aponeurosis of adductor mandibulae muscle Aw. Lig. prim: Ligamentum primordium.

B: Cirrhitops fasciatus (Group I species) Diagramatic representation of mandibulohyoid connection and related tendons; medial aspect of left
side to demonstrate the second form of tendinous relationships within species of Group I. Abbreviations as in Fig. 1A.

C: Paracirrhites forsteri (Group II species). Medial aspect of the right lower jaw, cheek region and hyoid arch, viewed somewhat dorsally; the
branchial skeleton and hyoid arch considerably displaced to the left and posteriorly in order to reveal the mandibulohyoid connection.
(Semi-schematic). About times natural size. 1: Posterior portion of mandibulohyoid connection, inserting partly on the quadrate, and partly
continuous with tendinous aponeurosis of the adductor mandibulae muscle Aw (Awt); 2b: ventral continuation of maxillary tendon of
adductor mandibulae muscle A,; 3: interopercular-mandibular ligament; 4: tendon of adductor mandibulae A, muscle; Lig. prim:

ligamentum primordium.

MANDIBULOHYOID CONNECTION IN CIRRHITOID FISHES
Lig. prim.

Add. mand. I u 2

Add. mand. I m ©
, = NGA

C Lig. prim.

- Max

95

96

P.H. GREENWOOD

Pal

Fig. 2. Mendosoma lineatum Medial aspect of left lower jaw, cheek region, and hyoid arch. Scale = 2mm. 2a: Maxillary tendon of adductor
mandibulae muscle A,; 2b: extension of tendon 2a, joining tendinous aponeurosis of adductor mandibulae muscle Aw; 2c: tendon of
adductor mandibulae muscle A,; 3a & b: extensions of adductor mandibulae muscle Aw’s tendinous aponeurosis; 6:
interopercular-mandibular ligament; 7: interhyal-metapterygoid ligament.

along, the dorsal and dorsolateral margins of the interopercu-
lum, ending at a point about midway between the bone’s
anterior tip and the face of the prominent, forward-facing
articulatory facet for the epihyal (cf pp. 94). Here it attaches
to a slight eminence on the dorsal margin of the interopercu-
lum. At first sight the ligament appears to be continuous with
the epihyal-interopercular ligament (Fig. 3; lig. 4) which also
inserts at that point. Careful dissection reveals, however, that
the two are separate entities (see also p. 93 and p. 94

respectively for the situation in cirrhitids and the latrid -

Mendosoma).

Apart from the more complex condition in cheilodactylids,
this double linkage of the mandible with the interoperculum,
one involving both tendons and ligaments, seemingly has not
been recorded in any other teleosts. However, it also occurs
in Mendosoma (see p. 94 and Fig. 2) where the lowermost
arm of the Aw aponeurosis is attached to the anteromedial
aspect of the interoperculum, and in Cheilodactylus (see
below, and tendon S in Fig. 4).

As in Mendosoma, the anguloarticular-dentary ligament in
A. monodactylus is short and stout.

An elongate and broad ligament (lig. 7 in Fig. 3) connects
the upper face of the interhyal with the quadrate and, mainly,
with the entopterygoid. This connection is similar to that in
Cheilodactylus (see Fig. 4, and p. 98), and, from its intercon-
nections would appear to be homologous with the ligament
joining the interhyal with the metapterygoid in Mendosoma
(Fig. 2; lig. 7) and the cirrhitid species examined.

The interhyal-interopercular ligament, present in all mem-
bers of the Cirrhitidae examined, is absent in the latrids and
cheilodactylids dissected. An interhyal-preopercular liga-
ment, present in the other cirrhitoids studied except the
cirrhitids, is also developed in Acantholatris. Here, although

very short, it is stout and has an extensive attachment area on
the interhyal and on the preoperculum, which it joins at the
point where the upper, vertical arm of that bone begins to
curve forward to form its horizontal arm.

A feature unique to Acantholatris monodactylus amongst
the cirrhitoid taxa examined is the presence of a well-defined
ligament connecting the hyoid arch and the dentary, a linkage
in no way associated with the adductor mandibulae Aw
muscle or its aponeurotic system (see Fig. 3). Posteriorly, this
ligament is attached to the summit of a distinct prominence
on the anterior face of the ceratohyal and situated immedi-
ately below the ceratohyal-epihyal suture. Anteriorly, the
ligament inserts on the dentary conjointly with the anterior
end of the anguloarticular-dentary ligament (see above and
Fig. 3; lig.1).

The family Cheilodactylidae

The account which follows is based on dissections of Cheilo-
dactylus fasciatus Lacepéde. Since the situation is virtually
identical in two other cheilodactylid species studied, Cheilo-
dactylus pixi (Smith) and Chirodactylus brachydactylus
(Cuvier), the term Cheilodactylus is used to cover all three
taxa. What interspecific differences do exist are noted on
page 98.

Of all the cirrhitoid species examined, the ligament and
tendon systems separately or conjointly linking the mandible,
the hyoid arch, the opercular series, and the palatoquadrate
arch in Cheilodactylus are by far the most complex. The
greatest similarities, however, are with those systems in the
latrid Acantholatris monodactylus (cf. Figs 3 & 4). In the
cheilodactylids examined, and like A. monodactylus, there is
no obvious subdivision of the adductor mandibulae A,

MANDIBULOHYOID CONNECTION IN CIRRHITOID FISHES

97

Fig. 3 Acantholatris monodactylus Medial aspect of left lower jaw, cheek region and hyoid arch. Scale = 2mm. 1: Anguloarticular-dentary
ligament; 2a: maxillary tendon of adductor mandibulae muscle A,; 2b: extension of above tendon joining tendinous aponeurosis of
adductor mandibulae muscle Aw; 3: tendon of Aw muscle to preoperculum; 4: epihyal-interopercular ligament; 5: tendon from Aw muscle
to interoperculum; 6: interopercular-mandibular ligament; 7: interhyal-quadrate-entopterygoid ligament. Dashed outline that of the

mandibulohyoid connection.

muscle, whose insertion on the maxilla is identical with that in
the latter taxon.

The Aw portion of the adductor mandibulae muscle in
Cheilodactylus is noticeably less extensive than in Acanthola-
tris, but its tendinous connections with the interoperculum
and the palatoquadrate arch are more complicated than in
that taxon. Also, in Cheilodactylus the ventral extension of
the adductor mandibulae A, maxillary tendon is noticeably
stouter than in Acantholatris (cf. Figs 3 & 4) but, unlike
Acantholatris, in Cheilodactylus it is derived from the medial
and not the lateral tendinous aponeurosis of the muscle’s Aw
division. A most obvious difference between the two taxa is
the absence of a ligament connecting the hyoid arch with the
mandible in Cheilodactylus (cf. Figs 3 & 4).

A somewhat tendinous section of the adductor mandibulae
Aw division (tendon 3 in Fig. 4) in Cheilodactylus runs
posteriorly, becoming completely tendinous as it crosses the
hind margin of the anguloarticular and its joint with the
quadrate. It inserts on the anterior tip of the preoperculum
just below that bone’s dorsal margin. At a point near the
centre of the anguloarticular this partly tendinous section of
the Aw division of the adductor mandibulae muscle gives off
a ventroposteriorly directed branch which soon becomes
completely tendinous. The anterior part of this tendon (Sa in
Fig. 4), immediately below its point of departure from tendon
3, is attached to the anguloarticular near its anterior margin.
It thus lies below the bone’s articulation with the quadrate.
The posteriad extension of tendon 5a runs backwards and
somewhat dorsally, seemingly joining the lateral face of a
broad, stout, dense, and obliquely orientated ligament-like
strap (5b in Fig. 4) extending from the midpoint of the
quadrate to the anteroventral surface of the interoperculum.
Together the two elements (ie 5a and 5b in Fig. 4) form a ‘Y’
shaped linkage between the anguloarticular, quadrate, and
interoperculum. Also, because the anterior arm of the ‘Y’ (ie

5a) is associated with an extension of the Aw muscle onto the
anguloarticular, the linkage involves that bone as well.

Without ontogenetic information it is difficult to decide
whether the element 5b of the ‘Y’ is, at it appears to be, a
branch of the tendon 5a (and thus is itself a tendon) or
whether it is strictly a ligament with which tendon 5a has
fused. That none of the other cirrhitoids examined has a
quadrato-interopercular ligament would add credence to 5b
being a true branch of Sa, and thus representing a consider-
able posterior extension of the Aw muscle’s tendon system.
Also, in the other cheilodactylids examined, the “Y’-shaped
connection gives no hint of it having originated from a tendon
and a ligament (see below). The potential complexity and
posterior extension of that system is clearly demonstrated in
another percomorph, the percid Gymnocephalus cernua (L.);
see Elshoud-Oldenhave & Osse (1976; fig. 4.1).

When comparisons are made with the latrid Acantholatris,
(see p. 94 and Fig. 3) it appears that the ‘Y’-shaped complex
in Cheilodactylus is, from its disposition and attachment
points, homologous with the simple tendon (5 in Fig. 3)
associated with the Aw portion of the adductor mandibulae
muscle in Acantholatris. Tendon 5 in that taxon is attached to
both the anguloarticular and the medial face of the interoper-
culum, and is separated by a short section of Aw from tendon
3 which inserts on the preoperculum (Fig. 3). In turn, and
also from its disposition and points of attachment, the latter
tendon would seem to be homologous with the longer tendon
3 in Cheilodactylus which also inserts on the preoperculum.
An early evolutionary stage in the development of this
complex in both Cheilodactylus and Acantholatris may be
represented by the tripartite posterior extension of the Aw
aponeurosis in Mendosoma, which also serves to link the Aw
muscle with the quadrate, preoperculum and interoperculum
(see p. 94 and Fig. 2).

The two other cheilodactylid species dissected, Chirodacty-

98

P.H. GREENWOOD

2a

\
A,

\
any
‘

st

Fig. 4 Cheilodactylus fasciatus Medial aspect of left lower jaw, cheek region and hyoid arch. Scale = 2mm. 1: Anguloarticular-dentary
ligament; 2a: maxillary tendon of adductormandibulae muscle A,; 2b: extension of above tendon joining tendinous aponeurosis of adductor
mandibulae muscle Aw; 3: tendon of adductor mandibulae muscle Aw to preoperculum; 4: epihyal-interopercular ligament; Sa: extension
of tendon 3; 5b: branch of tendon Sa, attaching to quadrate above and interoperculum below; 6: interopercular-mandibular ligament; 7:

interhyal-quadrate ligament; x: anguloarticular-quadrate ligament.

lus brachydactylus and Cheilodactylus_ pixi, have a
mandibular-preopercular-quadrate tendon system essentially
like that described above in Cheilodactylus fasciatus. In these
species the interopercular-quadrate branch (Fig. 4; 5b) does
not partly overlap that section of the complex (Fig. 4; Sa)
going to the anguloarticular. Instead, the two branches meet
in the same plane, with the result that the complex is clearly
single and ‘Y’-shaped. Since the specimens of Chirodactylus
brachydactylus (standard length 106 mm) and Cheilodactylus
pixi (S.L. 70-81 mm) are much smaller than the specimen of
Cheilodactylus fasciatus (S.L. 243 mm), the difference could
be related either to the larger size of the C. fasciatus specimen
or to individual variation.

The epihyal-interopercular ligament in Cheilodactylus is
short and broad (shorter even than that in the latrid Mendo-
soma; and unlike the long and anteriorly directed ligament in
the other latrid examined, Acantholatris, Fig. 4; lig. 4). As in
Acantholatris, but unlike Mendosoma, the interopercular
facet for the epihyal in Cheilodactylus is prominent and
well-developed (see Fig. 4). The interhyal-quadrate ligament
is long and flat (Fig. 4; lig. 7), again like that in Acantholatris,
but unlike its presumed homologue, the short and stout
interhyal-metapterygoid ligament in Mendosoma.

The interhyal-preopercular ligament in Cheilodactylus is
also short and stout. No discrete interhyal-interopercular

ligament is developed in the cheilodactylids, a characteristic
shared with the two latrid genera examined, but not with the
cirrhitid species studied.

A stout anguloarticular-dentary ligament is present, as it is
in the other cirrhitoids, but unlike those taxa Cheilodactylus
has a short and broad ligament (x in Fig. 4) connecting the
uppermost part of the anguloarticular’s posteromedial face to
the quadrate, where it is attached to the ventral rim of that
bone’s facet for articulation with the anguloarticular. This
small ligament, not found in any of the other cirrhitoids
examined, is almost entirely hidden by tendon Sa of the “Y’
shaped complex described above.

A very stout interopercular-mandibular ligament originates
laterally on the dorsal margin of the interoperculum near its
anterior tip, and inserts mostly on the lateral aspect of the
anguloarticular and retroarticular bones, but with a short
medial branch going to the posteromedial face of the retroar-
ticular (6 in Fig. 4).

DISCUSSION AND CONCLUSIONS

The taxonomically and phylogenetically widespread occur-

MANDIBULOHYOID CONNECTION IN CIRRHITOID FISHES

rence of a mandibulohyoid linkage in bony fishes (see Tcher-
navin, 1953, and references cited by Verraes, 1977, Springer
et al., 1877, Lauder & Liem, 1980, and above) certainly
seems to support the views of functional anatomists with
regard to its involvement in the mechanics of jaw opening. It
also refutes the apparently widespread view (see reviews in
Lauder & Liem, 1980; Lauder, 1982) that the linkage may be
a primitive character of neopterygian fishes, one lost in
higher teleosts (but see also Lauder & Liem, 1989, for later
views). However, although the mandibulohyoid connection
may be functionally homologous in both ‘higher’ and ‘lower’
bony fishes, there are indications that it may not be homolo-
gous in an ontogenetical and hence phylogenetic context (see
below). Nevertheless, the diversity of mandibulohyoid con-
nections already known in but a few teleost fishes strongly
suggests that the structural, functional and ontogenetic
aspects of this system need to be reevaluated.

Any attempt to establish or refute the homology of man-
dibulohyoid connections in cirrhitoid fishes with those in
other bony fish groups (see below) is hampered by a lack of
information on the ontogeny of the linkage in the various taxa
involved. Indeed, this problem also arises with the different
mandibulohyoid linkages found within the cirrhitoids them-
selves, namely those in the Cirrhitidae (p. 93) and that in the
latrid Acantholatris monodactylus (p. 97).

The cirrhitid linkage type in the Paracirrhites species
examined (p. 93) strongly suggests that the connection
between the mandible and the ceratohyal in these fishes is
derived from an extension of the central aponeurosis of the
adductor mandibulae muscle’s Aw portion onto the hyoid
arch (with, in addition, a partial insertion on the quadrate;
Fig. 1C and p. 93). In another cirrhitid group (viz. Cyp-
rinocirrhites polyactis, Cirrhitichthys oxycephalus and Cirrhi-
tops fasciatus) the connection also has a linkage with the
aponeurosis of adductor mandibulae Aw. Here it is effected,
somewhat indirectly, by a branch from the major mandibulo-
hyoid connection joining the maxillary tendon of adductor
mandibulae A, muscle, which tendon itself is derived from
the aponeurosis of the Aw portion of that muscle.This
association with the Aw aponeurosis in both cirrhitid groups
raises the possibility that ontogenetically, the mandibulohy-
oid linkage is through a tendon and not a ligament as it
appears to be in the salmonid Oncorhynchus mykiss (see
Verraes, 1977). It also raises the question whether or not the
so-called mandibulohyoid ligament (see below) in other
teleosts (and in the semionotiform Lepisosteidae; see below)
is truly a ligament. A similar problem arises with the third
type of mandibulohyoid connection found in cirrhitoids,
namely that in the latrid Acantholatris monodactylus. Here
the linkage is not associated with the Aw muscle, and has
both its origin and its insertion entirely on bone, thus
appearing to be a true ligament.

There is some indirect support for the idea that in members
of the Cirrhitidae the mandibulohyoid connection could be
derived ontogenetically from the adductor mandibulae
muscle bloc (sensu Edgeworth, 1935) of the early embryo.
This stems from the considerable posterior extension of the
adductor mandibulae Aw aponeurosis onto the bones of both
the palatoquadrate arch and the interoperculum in certain
other perciform fishes (see also discussions in Winterbottom,
1974; Elshoud-Oldenhave & Osse, 1976; Anker, 1978;Green-
wood, 1985) and, indeed in other cirrhitoids such as the
cheilodactylids.

An origin of the mandibulohyoid connection from the

99

adductor mandibulae Aw tendon system seems less likely in
the latrid Acantholatris monodactylus. Here the linkage
extends from the posterior tip of the dentary’s lower arm
(not, as in the cirrhitids, from its coronoid process or the
anguloarticular) to the upper part of the ceratohyal’s lateral
face (Fig. 3). At no point has this apparent ligament in
Acantholatris any association with the adductor Aw muscle or
any part of its tendon system. With regard to its attachment
to the lower aspect of the dentary, the connection is compa-
rable both with the loosely compacted and fibrous linkage
between the dentary and ceratohyal identified by Aerts et al.
(1987) in the cichlid Astatotilapia elegans, and with Osse’s
(1969) ligament XXIV in the percid Perca fluviatilis. In both
these species, however, the tissue has insertions on certain
branchiostegal rays as well as on the ceratohyal, and in
neither species does it have the ligament-like appearance of
the connection in Acantholatris monodactylus.

Aerts et al. (1987:97) describe in some detail the histology
of the hyoid-dentary connection in Astatotilapia elegans,
which seemingly is derived from the anterior, tendinous part
of the geniohyoideus muscle, with whose dorsolateral aspect
it is closely associated over much of its length. These authors
conclude (op. cit.: 99) that ‘In fact, the rostral part of the
interconnection can be interpreted as a parallel elastic com-
ponent of the protractor hyoidei’ (=geniohyoideus). The
posterior attachment of the connection is on the epi- and
ceratohyals dorsally, with, as noted above, a number of small
strands merging into the dermal layers of the skin-fold
between the hyoid and interoperculum. A mandibulohyoid
connection, superficially like that in A. elegans also occurs
(pers.obs.) in another haplochromine cichlid, Thoracochro-
mis buysi (Penrith); although its histology was not studied,
the linkage appears to originate from within the geniohyoid
muscle, and to attach to the hyoid arch at the epi-ceratohyal
suture.

At least with regard to its superficial features, Aerts et al.’s
description of the dentary-hyoid connection in Astatotilapia
elegans does not resemble the condition seen in Acantholatris
monodactylus. Here, the interconnecting tissue is clearly
separated from the geniohyoideus muscle over virtually its
entire length, and is much more compact and ligament-like.
However, posteriorly it does appear to fuse with the tendi-
nous insertion of the geniohyoideus at the point where both
elements attach to an elevation on the anterior margin of the
certohyal. The insertion of the geniohyoideus muscle then
extends down along the lateral face of the ceratohyal, but that
of the mandibulohyoid connection does not. Thus in adult
Acantholatris monodactylus the only suggestion of the con-
nection being derived from the geniohyoideus muscle is a
partially shared insertion with that muscle on the ceratohyal.
That suggestion is, unquestionably, far less convincing than
the evidence provided by the situation in Astatotilapia
elegans, but is one that could be clarified if studied ontoge-
netically in Acantholatris monodactylus.

A distinct mandibulohyoid ligament, superficially like that
in Acantholatris monodactylus, has been described by Wiley
(1976) in the semionotiform gars Lepisosteus and Atrac-
tosteus. The connection is labeled as a tendon in figure 9 of
Wiley’s paper, but is referred to, I believe correctly, as a
ligament in the accompanying text. The ligament in gars
differs from the ligament-like mandibulohyoid connection in
Acantholatris monodactylus in its points of attachment (epi-
hyal and retroarticular in the gars, ceratohyal and dentary in
A. monodactylus). Again, without ontogenetic information

100

from both taxa, nothing can be said about its possible
homology in the two species.

The concept that a mandibulohyoid connection (usually
referred to as a ligament) is essentially a feature of pre- and
lower teleost actinopterygians, has influenced theories relat-
ing to the evolution of feeding machanisms in teleosts. For
example, Lauder (1982: 279, also fig. 1) postulated that “The
first specialization involves a shift of insertion of the man-
dibulohyoid ligament to the interoperculum. The interoper-
culohyoid ligament characterizes the feeding mechanism of
eurypterygian fishes (=Aulopiformes + Myctophiformes +
Paracanthopterygii + Acanthopterygii; Rosen, 1973) and
effectively shifts the action of the hyoid and opercular cou-
pling onto the interoperculum. Only the interoperculoman-
dibular ligament transmits posterodorsal hyoid and opercular
movements to the mandible in the Eurypterygii, while other
teleosts retain the primitive two-coupling system of the
halecostomes’ (ie both a mandibulohyoid and an
interopercular-mandibular linkage). Verraes’ (1977) studies
on the development of Oncorhynchus mykiss show unequivo-
cally that in this teleost there is no ontogenetic shift of the
mandibulohyoid ligament’s mandibular insertion onto the
interoperculum. Indeed, the interopercular-mandibular liga-
ment develops independently (and later than the mandibulo-
hyoid ligament) with both connections persisting in adults
(Verraes, 1977; pers. obs.); neither is there any ontogenetic
evidence to show that the epi- (or inter-) hyal to interopercu-
lum ligament is the result of a preexisting mandibulohyoid
ligament shifting its mandibular insertion onto the interoper-
culum. Interestingly in that context, the latrid Acantholatris,
which has what appears to be a genuine mandibulohyoid
ligament (see p. 97) also has an epihyal-interopercular liga-
ment.

Thus, pace Lauder (1982), it would seem that cirrhitoids
(and other teleosts) with both a mandibulohyoid connection
and an interopercular-mandibular ligament have either
retained the primitive halecostome condition or, as seems
more likely, re-evolved it through some other form of con-
nective tissue linkage between the hyoid arch and the man-
dible.

Parenthetically, it may be noted that the importance of an
interopercular-mandibular linkage in the jaw-opening mecha-
nism of teleosts, stressed by Lauder op.cit. and other authors
(see for example Liem, 1978 & 1991; Aerts et al., 1987, and
references therein) is underlined, albeit indirectly, by the
condition in three of the cirrhitoid taxa examined. In Cheilo-
dactylus (Cheilodactylidae) and in Mendosoma and Acantho-
latris (Latridae) there is, in addition to the interopercular-
mandibular ligament a second such linkage effected through
an extension of the Aw muscle’s aponeurotic system onto the
interoperculum (see pp. 94 & 97 and Figs 2-4).

If, as suggested above, certain teleosts have re-evolved a
mandibulohyoid connection, it may have arisen in different
ways. This seems probable even within the cirrhitoids (viz.
cirrhitid and latrid types; see pp. 93 & 94), and in other
groups as well. In the ostariophysan Hydrocynus vittatus
(Characidae) for example, the mandibulohyoid connection
appears to be an extension of the epihyal-interopercular
ligament which, after its insertion on the dorsal margin of the
interoperculum, continues forward to bridge the small gap
between that bone and the retroarticular (pers. obs.). The
salmonid Oncorhynchus mykiss, by contrast, has no obvious
association of the mandibulohyoid connection with the
epihyal-interopercular ligament. Both are discrete entities

P.H. GREENWOOD

throughout their lengths despite having insertion points close
together on the epihyal (pers. obs.). The clupeid Etrumeus,
unlike the preceding examples, has no readily discernible
mandibulohyoid connection. However, the geniohyoideus
muscle has a thickened and tendinous dorsal margin which is
macroscopically continuous with the muscle from the latter’s
origin near the dentary symphysis to its insertion immediately
over the epi-ceratohyal suture (pers. obs). Superficially at
least, the situation in this clupeid shares certain similarities
with the mandibulohyoid link in the perciform cichlid Astato-
tilapia elegans (see Aerts et al., 1987, and p. 99 above). In the
clupeid, however, the differention of the linkage from the
associated muscle is at a somewhat lower level of develop-
ment than that in the cichlid.

Verraes (1977) highlighted the functional importance of
the mandibulohyoid connection in immediately post-hatching
stages of the salmonid Oncorhynchus mykiss. This apparently
ligamentous connection develops earlier than _ the
interopercular-mandibular ligament. Thus at this point in the
fish’s life-history it is an essential element in bringing about
jaw depression, and consequently it plays a major role in the
creation of the trans-buccal water current involved in respira-
tion and feeding (see also Lauder & Liem [1989] for a
discussion of this ligament in the feeding mechanism of
another salmonid, Salvelinus fontinalis). Recently, Aerts et
al., (1987), working with the cichlid Astatotilapia elegans,
postulated that a mandibulohyoid connection is also of crucial
importance in the feeding mechanism in adults of that spe-
cies.

Regretably, no experiental work has been carried out on
the feeding mechanisms of cirrhitoid fishes, nor is there
enough critical information on their feeding habits to deter-
mine what correlations may or may not exist between species
with or without a mandibulohyoid connection. It would be
interesting to know in what way the mandibulohyoid connec-
tion functions in cirrhitids such as Cyprinocirrhites polyactis.
Judging from preserved specimens it would seems to block
the sinking of the lower jaw when the hyoid is pulled
posteriorly by the contracting sternohyoideus muscle — a
somewhat anomalous situation, but possibly one that may be
associated with a specialized suction mode of feeding on small
crustacean zooplankters, apparently the principal food of this
species in South African waters.

As yet, the intrabuccal tendon and ligament systems are
known from too few cirrhitoid taxa to test its usefulness in the
intragroup taxonomy and phyletic relationships of those
fishes. However, the tendon system in the Cheilodactylidae
examined, when compared with that in the latrid Acanthola-
tris monodactylus (cf. Figs 4 & 3) supports the latter taxon’s
removal (see Greenwood, 1995) from the genus Cheilodacty-
lus and the family Cheilodactylidae in which it had been
placed previously. Those differences also provide an addi-
tional character complex for distinguishing the Latridae from
the Cheilodactylidae.

ACKNOWLEDGEMENTS. My thanks go to Professor Tom Hecht and Dr
Colin Buxton of Rhodes University’s Department of Ichthyology and
Fisheries Science, as well as to their students, for giving me access to
that Department’s collections of preserved material, and for collect-
ing other specimens when needed. To Dr Phil Heemstra of this
Institute, my thanks for information on, and discussions about,
cirrhitoid fishes. For her patience, forbearance and skill, it is a
pleasure to thank Huibré Tomlinson who once again has turned my

MANDIBULOHYOID CONNECTION IN CIRRHITOID FISHES

untidy manuscript into a legible typescript. Anthea Ribbink, who
also displayed those attributes when producing the anatomical fig-
ures, deserves my deepest gratitude for her essential contributions to
this paper. By no means least of all, I am indebted to Drs Mark
Westneat and Rick Winterbottom for their very constructive com-
ments on an earlier draft of the paper.

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Winterbottom, R. 1974. A descriptive synonymy of the striated muscles of the
Teleostei. Procedings of the Academy of Natural Sciences of Philadelphia
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Bull. nat. Hist. Mus. Lond. (Zool.) 61(2): 103-109

Issued 30 November 1995

A new species of Crocidura (Insectivora:
Soricidae) recovered from owl pellets in

Thailand

PAULINA D. JENKINS

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD

ANGELA L. SMITH
University of Cambridge, Cambridge

Synopsis. A new species of Crocidura (white-toothed shrew) is described from owl pellets from Loei Province,
Thailand. The craniodental morphology is compared with that of similar sized species of Crocidura recorded from

Thailand.

INTRODUCTION

A recent survey of bat roosts and owl pellets in Thailand by
one of us (ALS) and Mark F Robinson has increased knowl-
edge of the small mammal fauna of the area. Contained in the
owl pellets were skulls of 38 species of small mammals,
including an unknown species of Crocidura. This undescribed
species has sufficiently distinctive cranial and dental charac-
ters to warrant its description on the basis of these features
alone, although the external characters remain unknown.

MATERIALS AND METHODS

Regurgitated pellets were collected from roosting sites of
barn owls (Tyto alba [Scopoli, 1769]) at several localities in
Thailand. The pellets were dissected and the contents, usu-
ally incomplete crania and mandibles, were identified as far
as possible in the field. Voucher specimens were sent to The
Natural History Museum for confirmation of identification.
Included among these specimens was a series of Crocidura
which was proving difficult to identify and was thought to
include two species, C. attenuata Milne-Edwards, 1872 and C.
fuliginosa (Blyth, 1855).

Measurements, in millimetres, were taken using dial cali-
pers or a micrometer eyepiece and measuring stage on a
microscope. Cranial and dental nomenclature follows that of
Meester (1963), Mills (1966), Swindler (1976) and Butler &
Greenwood (1979). Abbreviations for the dental nomencla-
ture are given in the text.

RESULTS

Crocidura hilliana sp. nov.

HOLOTYPE. BM(NH)1994.90, collector’s number 467. Cra-
nium with damaged braincase, left mandibular ramus, com-

© The Natural History Museum, 1995

plete maxillary and mandibular dentition. Extracted from an
owl pellet from a roost at Wat Tham Maho Lan, Ban Nong
Hin, 48 km south of Loei, Loei Province, northeastern
Thailand, 17°06'N 101°53’E, altitude 575m.

PARATYPES. Eighteen specimens of crania with mandibles
and thirteen specimens of crania only, all from owl pellets at
the same locality as the holotype. Three specimens of crania
with mandibles and five specimens of crania only from Wat
Tham Pha Phu, 7 km north of Loei, Loei Province, 17°34’N
101°42’E, altitude 542m.

Diagnosis

Zygomatic process of maxilla broad and angular, interorbital
region narrow; coronoid process broad and deep. Upper and
lower first incisors robust, first upper unicuspid large and
broad relative to the other unicuspids, talonid of the third
lower molar (M;) reduced to a single cusp.

Description

Overlapping in cranial size with medium to large specimens
of Crocidura attenuata and smaller specimens of C. fuliginosa
but differing from both species in proportions (see Table 1
and Figs 1-6). Cranium and mandible robust; cranium angu-
lar in appearance, especially in dorsal view. The rostrum is
moderately deep and obliquely sloping anteriorly; the maxil-
lary region is broad, the zygomatic process of the maxilla is
broad and angular; the interorbital region is long and narrow,
its width increasing only slightly from anterior to posterior;
the zygomatic plate is positioned above the first upper molar
(M') and the anterior of the second upper molar (M7), its
posterior face is semi-circular and deeper than the anterior
face; the braincase is angular, with pronounced angular
superior articular facets in dorsal view, a squamosal crest is
present and lambdoid crests are well developed, meeting at
an acute angle at the midline. The horizontal ramus of the
mandible is moderately robust; the coronoid process is broad
and deep (see Fig. 4); the ascending ramus is long and low;
the condyle is nearly as broad or broader than deep (ratio of

104

P.D. JENKINS AND A.L. SMITH

Fig. 1
BM(NH)1933.4.1.183.

condyle width to height 91.2-113.3), the postero-internal
ramal fossa has a broad base and is approximately as broad as
deep; the mental foramen is positioned below the anterior
part of the first lower molar (M,).

The dentition is illustrated in Figs 2-6. The first upper
incisor (I') is robust, slightly proodont with well developed
posterolingual and posterobuccal cingula; the upper unicus-
pids are overlapping and crowded; the first upper unicuspid
(Un') is large and broad in comparison with the other
unicuspids, its breadth is equal to or greater than the distance
between the two first unicuspids and this tooth is more than
two thirds the height of I' and P*; the second and third
unicuspids (Un* and Un*) are subequal in size; the upper
premolar (P*) has a moderately small parastyle and robust
metacone; the first and second upper molars (M' and M7?)
show no significant distinguishing features; the third upper
molar (M?°) is short and slender with a slightly compressed
lingual basin. The first lower incisor (I,) is robust, long, deep
and curved, and the anterolingual ridge extends for circa
three quarters of the length of the tooth, diverging from the
ventral border, the posterior border of I, lies below the
middle of the lower premolar (P,); two thirds of the second
lower incisor (I,) are in contact with I, and one third of the
tooth is overlapped by P,; the postentoconid ledge is very
narrow in the first lower molar (M,) and yet more reduced in
the second lower molar (M,); the talonid of the third lower

Dorsal view of cranium from left to right of C. attenuata BM(NH)1911.9.8.26, C. hilliana BM(NH)1994.113 and C. fuliginosa

molar (M,) is reduced to a single cusp.

Etymology

This species is named in honour of John Edwards Hill, who
taught one of the authors (PDJ) the basics of mammalogy and
who also provided invaluable help in the identification of
some of the skull fragments of bats found during the survey.

Comparison with other species

Five species of Crocidura have been recorded from Thailand
(Lekagul & McNeely, 1977, Davison, 1984): C. fuliginosa
(including C. dracula Thomas, 1912 listed as a separate
species by Lekagul and McNeely), C. attenuata, C. pullata
vorax Allen, 1923 (listed as C. russula vorax), C. horsfieldii
indochinensis Robinson & Kloss, 1922 and C. monticola
Peters, 1870. Crocidura hilliana is separated from most
specimens of C. fuliginosa dracula by its smaller size (see
Table 1), while it is considerably larger than C. p. vorax
(condylobasal length <17.5), C. horsfieldii (condyloincisive
length <17.9, data taken from Heaney & Timm (1983) for
specimens from Vietnam) or C. monticola (condylobasal
length <17.4).

Crocidura hilliana falls at the middle to upper part of the
cranial size range of C. attenuata and the lower part of the size

NEW SPECIES OF CROCIDURA

Fig. 2 Ventral view of cranium from left to right of C. attenuata BM(NH)1911.9.8.26, C. hilliana BM(NH)1994.113 and C. fuliginosa

BM(NH)1933.4.1.183.

range of C. fuliginosa (see Table 1). It is readily distinguished
from both species by its robust, angular cranium, in which the
maxillary region is broad, the interorbital region narrow and
the anterior part of the braincase markedly angular (see Figs
1-3). In contrast, both C. attenuata and C. fuliginosa have a
proportionally narrower maxillary region, broader interor-
bital region increasing noticeably from anterior to posterior
and a more rounded braincase that is evidently broader than
the maxillary region. Lambdoid crests are more or less well
developed in both C. hilliana and C. fuliginosa, but they meet
at an acute angle at the midline in C. hilliana and a shallower
angle in C. fuliginosa; lambdoid crests are less developed in
C. attenuata and meet at a shallow angle. Squamosal crests
are absent or ill defined in C. attenuata, poorly to moderately
defined in C. fuliginosa but well-marked in C. hilliana. The
mandible of C. hilliana is considerably more robust than that
of either of the other two species (see Fig. 4). The horizontal
ramus of the mandible of C. attenuata is more slender than
that of C. hilliana, with a sinuous ventral border; the coro-
noid process is considerably narrower and shallower; the
ascending ramus is higher and the condyle is higher than
broad (ratio of condyle width to height 75.0-93.3). The
horizontal ramus of the mandible of C. fuliginosa is longer yet
shallower than that of C. hilliana, with a narrower, less robust
coronoid process and, as in C. attenuata the condyle is higher
than broad (ratio of condyle width to height 77.8—93.3). The

mental foramen lies below the posterior part of P, in C.
attenuata and C. fuliginosa but below the anterior of M, in C.
hilliana. Dentally the most obvious differences between C.
hilliana and the other two species is the comparatively large
anterior dentition (I', Un' and I,) relative to the rest of the
teeth, in combination with the narrow M® and the reduced M,
of C. hilliana, differing considerably from the condition in
either C. attenuata or C. fuliginosa (see Table | and Figs 2-6).

In detail the dentition of C. attenuata differs in the follow-
ing aspects from that of C. hilliana: 1' is slender and orth-
odont, the posterolingual cingulum is narrow, Un!' is
moderate in size and the distance between the two first upper
unicuspids is greater than the breadth of Un', Un? is smaller
than Un’ and Un’, and the unicuspids overlap only slightly so
that the rostrum is moderately long in appearance; the
parastyle of P* is moderately well developed. M? is variable in
different populations of C. attenuata; it is medium sized in
Indian and Burmese populations and thus readily distin-
guished from C. hilliana, and although only narrower on
average in the Chinese populations of C. attenuata, neverthe-
less, the lingual basin is less compressed than in C. hilliana.
The first lower incisor of C. attenuata is moderately slender,
straighter and more procumbent than that of C. hilliana; the
anterolingual ridge extends for two thirds the length of the
tooth and is subparallel to the ventral border of the tooth; the
posterior border of I, lies below the posterior part of [,; less

106

Table 1 A comparison of species of Crocidura occuring in Thailand and nearby countries.

P.D. JENKINS AND A.L. SMITH

C. hilliana C. attenuata C. fuliginosa

Thailand China India Thailand Vietnam China
Condylobasal length 21.0-23.5 19.8-20.7 19.7-21.6 22.0, 22.8 21.3-23.4 21.6-22.7
mean 22.20 20.20 20.23 22.58 22.20
SD 0.68 0.38 0.54 0.51 0.47
n 16 8 10 Z 15 4
Upper toothrow length 8.8-10.2 8.7-9.4 8.7-9.8 10.1-10.8 9.8-10.8 9.7-10.7
mean 9.45 9.00 9.12 10.42 10.25 10.17
SD 0.35 0.23 0.29 0.29 0.27 0.35
n 37 12 17 5 24 11
Maxillary breadth at level of M* 6.0-7.2 5.7-6.4 5.8-6.5 6.6-7.0 6.7-7.3 6.7-7.2
mean 6.57 6.14 6.08 G72 6.96 6.91
SD 0.30 0.21 0.19 0.16 0.15 0.17
n 38 12 17 5 23 11
Interorbital breadth 3.8-4.6 4.2-4.8 4.2-4.7 4.4-4.7 4.7-5.3 4.7-S.2
mean 4.27 4.46 4.39 4.55 4.93 4.93
SD 0.21 0.19 0.12 0.13 0.13 0.19
n 38 10 15 + 22 8
Braincase breadth 8.9-10.0 8.5-9.5 8.7-9.8 9.9-10.1 9.8-10.7 9.9-10.6
mean 9.56 9.09 9.08 10.00 10.24 10.14
SD 0.36 0.34 0.33 0.12 0.24 0.26
n 16 9 11 3 18 5
Mandible length excluding I, 10.7-12.7 10.0-11.5 10.1-11.2 11.8-12.4 11.7-13.1 11.5—12.7
mean ESS 10.71 10.62 12.09 12.36 12.06
SD 0.55 0.46 0.42 0.24 0.35 0.41
n 21 13 iy 8 26 11
Mandible height 5.2-6.3 4.4-5.1 4.5-5.2 5.2-5.8 5.45.9 5.2-6.0
mean 5.78 4.81 4.65 5.54 5.61 5.62
SD 0.28 0.22 0.21 0.18 0.16 0.25
n 74) 13 17 9 26 g)
Interorbital breadth: maxillary breadth 60.5-70.5 68.8-77.7 68.3-77.6 65.7-69.7 67.1-75.4 68.9-73.2
mean 65.00 72.36 72.40 67.43 70.86 71.09
SD 2.44 2.98 2.48 1.66 2.26 1255
n 38 10 15 4 22 8
Length of M?: upper toothrow length 5.2-7.0 6.4-6.9 7.1-8.0 6.8-7.9 6.6-8.0 6.9-8.0
mean 6.12 6.67 Uc) 7.48 7.28 7.34
SD 0.51 0.16 0.28 0.42 0.42 0.35
n 34 12 ily 5 24 10

than half of I, is in contact with I, and I, is one quarter
overlapped by P,; a postentoconid ledge is present in M, and
M.; the talonid of M; is relatively complete and an entoconid,
entoconid ridge and talonid basin are present.

Crocidura fuliginosa differs from C. hilliana in having a
moderately slender, orth-opisthodont I’ with a smaller
although well developed posterolingual cingulum; Un! is
moderate in size (c half the height of I’ and P*); in contrast to
the condition in C. attenuata, Un? is only slightly smaller than
Un’ and Un’; the lingual region of P* is characteristic in
shape; the mesostyle of M? is divided into two stylar cusps
(see Ruedi, in press) unlike either of the other species; M? is
medium in size and the lingual basin is not compressed. The
mandibular dentition is similar to that of C. attenuata. In
particular it is readily distinguished from C. hilliana by the
less robust, straighter, more procumbent first lower incisor;
slightly over half of I, is in contact with I,; the talonid of M, is
not reduced and an entoconid, entoconid ridge and talonid
basin are present.

DISCUSSION

It is known from the study of barn owl pellets in the British

Isles and Africa (Glue, 1967; Andrews 1990) that prey
skeletal elements are subject to little breakage or digestion,
contrary to the case for pellets of some other avian predators.
Certainly there is a degree of damage to all of the crania in
the current sample, none of which are intact. Crania and
associated mandibles occur in 48%; a few specimens are
nearly complete showing only minimal damage to the brain-
case, although the braincase is broken or absent in most
specimens. The toothrows are complete in 87% of specimens,
although the teeth may be loose in their sockets, with tooth
loss occuring generally at the terminal molar or unicuspid
loci. There is little evidence of digestive erosion of crania or
teeth. It has therefore proved possible to take most of the
standard cranial measurements on sufficient of the recovered
crania and mandibles to obtain significant data on size
variation. Similarly, the dentition is well preserved so that
diagnostic characters are readily observed and allowing the
samples to be aged. Shrews of the genus Crocidura show very
rapid dental maturation as nestlings, teeth are fully erupted
shortly after leaving the nest. The dental ages appearing in
these samples include fully erupted dentitions with no sign of
tooth wear, probably representing juvenile or subadult speci-
mens; dentitions showing slight to moderate wear, represent-
ing adults; dentitions showing extreme wear, representing old
adults.

NEW SPECIES OF CROCIDURA

107

Fig. 3. Lateral view of cranium from top of C. attenuata BM(NH)1911.9.8.26, C. hilliana BM(NH)1994.113 and C. fuliginosa

BM(NH)1933.4.1.183.

There have been few systematic collections of the small
mammal fauna in Thailand, which in consequence remains
comparatively little known; in particular the shrews are
poorly documented. Crocidura fuliginosa was recorded from
peninsular Thailand by Bonhote (1903), Kloss (1917) [as C.
aagaardii|, Robinson & Kloss (1923) and Hill (1960) [prob-
ably referring to the same specimen as Robinson & Kloss
(1923)], and from Koh Samui off the east coast of peninsular
Thailand by Robinson & Kloss (1914) [as C. negligens]. The
inclusion in this taxon of two chromosomally distinct but
morphologically cryptic species in Malaysia was discovered
recently by Ruedi ef al. (1990). Ruedi (in press) has
attempted to correlate morphological features with these
chromosomal forms, in order to assign specific names to
them, reserving the name C. fuliginosa for those specimens
with chromosomes 2n = 40, Fundamental Number 56 and
ascribing the other species, with polymorphic chromosomes
of 2n = 38-40, to C. malayana Robinson & Kloss, 1911.
Regrettably, examination of Malaysian specimens in the
collection of the Natural History Museum fails to confirm the
supposedly clearcut morphological distinction, with some
specimens exhibiting a mixture of the characters listed by
Ruedi, so negating the use of these morphological criteria.
Crocidura fuliginosa is a widely distributed species, occuring
from Burma in the west to China in the east and southwards
to Indonesia, including a number of named forms, whose

taxonomic status has been the subject of considerable discus-
sion (Medway, 1965, 1977; Jenkins, 1976, 1982; Heaney &
Timm, 1983; Corbet & Hill, 1992). The presence of cryptic
species in Malaysia, emphasises the lack of understanding of
the status of C. fuliginosa, suggesting that it requires further
revision and might be more appropriately considered as a
species complex. There are few records of this species from
regions other than peninsular Thailand, apart from that of
Lekagul & McNeely (1977) from Chiengmai, or Chiang Mai,
northwest Thailand (as C. fuliginosa and C. dracula). Fur-
thermore, there are no specimens of C. fuliginosa from
Thailand, other than peninsular Thailand, in the collection of
The Natural History Museum, while in the collection of the
American Museum of Natural History there are single speci-
mens from Nakhon Nayok, Khao Yai National Park and
Nakhon Ratchasima, central Thailand, plus an unconfirmed
specimen from Umphang, western Thailand. In the current
survey, C. fuliginosa was identified from prey remains of the
carnivorous bat, Megaderma lyra E. Geoffroy, 1810 collected
at Thung Yai—Huai Kha Khaeng Wildlife Sanctuary, western
Thailand; however there are only a few fragmentary speci-
mens, dubiously attributed to this species, among the owl
pellets from Loei Province. There are similarly few records of
C. attenuata from Thailand; Lekagul & McNeely (1977) listed
this species from Nakhon Phanom and Udon in the northeast,
and Chiang Mai, northwest Thailand. There was no evidence

108

TUT

Fig. 4 Lateral view of mandible from top of C. attenuata
BM(NH)1911.9.8.26, C. hilliana BM(NH)1994.90 and C.
fuliginosa BM(NH)1933.4.1.183.

of C. attenuata either amongst the owl pellet remains from
Loei Province or from remains found at M. lyra roosts in
Thung Yai—Huai Kha Khaeng. It is therefore uncertain if C.
hilliana is sympatric with either C. fuliginosa or C. attenuata.

Crocidura hilliana does, however, occur sympatrically with
a smaller species of Crocidura which proved difficult to
determine from the fragmentary skulls in the owl pellets.
Allen & Coolidge (1940) collected C. vorax (currently
grouped with C. pullata Miller, 1911 from the Himalayas, see
Hutterer, 1993) from northwestern Thailand, while a speci-
men from Lat Bua Kao, mainland Thailand, attributed to C.
fuliginosa by Kloss (1919) is also an example of C. p. vorax.
Several skulls attributable to this species were found in the
owl pellets from Loei, while a good series was recovered from
the M. lyra prey remains from Thung Yai—Huai Kha Khaeng,
where an additional skull was found in the faeces of a large
carnivore. The only other species of Crocidura listed by
Lekagul & McNeely (1977) from mainland Thailand was C.
horsfieldii indochinensis from Chiang Mai and Khao Yai
National Park . Most recently, Davison (1984), recorded C.
monticola from penisular Thailand. Neither of the last two
species mentioned above were identified from either area,
although pellets from Loei Province contained another shrew
Suncus etruscus (Savi, 1822), plus a variety of rodent and bat
species.

Since there has been so little systematic collection in
Thailand, it is impossible to make categoric statements about
the new species, however it seems likely that it is relatively
localised in its distribution. Even in areas where collecting

P.D. JENKINS AND A.L. SMITH

Fig. 5 Lateral view of left anterior dentition. Left: upper toothrow
(I' to P*); right: lower toothrow (I, to P,). Top: C. attenuata
BM(NH)1911.9.8.26; middle: C. hilliana BM(NH)1994.119 upper
toothrow, BM(NH)1994.118 lower toothrow; bottom: C.
fuliginosa BM(NH)1933.4.1.178. Scale 1 mm.

Fig. 6 Occlusal view of left upper toothrow from left to right of C.
attenuata BM(NH)1911.9.8.26, C. hilliana BM(NH)1994.121 and
C. fuliginosa BM(NH)1933.4.1.178. Scale 1 mm.

NEW SPECIES OF CROCIDURA

efforts have been more stringent, shrews are frequently
difficult to trap, perhaps giving a false impression of their
rarity as faunal components. The discovery of this new
species of shrew, apparently present as a sufficiently large
population to form an important and regular part of the diet
of the resident owls, is therefore not so surprising as it might
first appear. Because of the nature of the specimens, even
less information than usual is known about the ecology of the
new species, although some implications may be drawn from
knowledge of the ecology and behaviour of the owls. The
barn owl roosting sites of both collecting localities are caves
in limestone outcrops in or near temple grounds, surrounded
by bamboo and deciduous trees. Individual roost sites at Wat
Tham Maho Lan are generally within 0.5 km of cultivated
maize fields, while those at Wat Tham Pha Phu are within 1
km of rice and cassava fields. The home range of barn owls in
the British Isles and Africa is generally 1-2.5 km, rarely up to
3 km (Bunn et al. 1982; Andrews, 1990). Because of this small
hunting range, it may be inferred that this habitat which
extends for some distance around the roosting site is also the
habitat for the shrews on which they prey. Barn owls are
nocturnal and crepuscular in their hunting behaviour, the
implication being that the shrews are active for at least a
proportion of the same activity period.

ACKNOWLEDGEMENTS. We would like to thank the monks and nuns at
Wat Tham Pha Phu and Wat Tham Maho Lan for their generous
hospitality and their help in locating owl roosts. Mr Jarujin Nabhitab-
hata, Mr Preecha Leucha, Mr Surachit Wargsothorn and Ms Sunee
of the Ecological Research Department, Thailand Institute of Scien-
tific and Technological Research, Bangkok, provided much help and
advice, and kindly allowed access to their reference collection of
mammal specimens. We are very grateful also to the staff at Wildlife
Fund Thailand, in particular Mr Surapon Duangkhae, Mr Siripong
Thonongto and Mr Patric Corrigan, for their help and support. We
thank Dr Robert Mather (WWF) and his wife Noi, who provided
logistical support and Dr Mark Robinson who collaborated on the
survey, for his help and encouragement throughout the project. We
are indebted to Dr Robert Prys-Jones, Bird Group, The Natural
History Museum and Dr Rainer Hutterer, Zoologisches Fors-
chungsinstitut und Museum Alexander Koenig, Bonn, Germany for
their constructive reviews of the manuscript.

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Issued 30 November 1995

Redescription of Sudanonautes floweri (De
Man, 1901) (Brachyura: Potamoidea:
Potamonautidae) from Nigeria and Central

Africa

NEIL CUMBERLIDGE
Department of Biology, Northern Michigan University, Marquette, Michigan 49855, USA
CONTENTS
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Synopsis. The African fresh-water crab Sudanonautes floweri (De Man, 1901) is redescribed from the male syntype
from Sudan (designated here the lectotype) and a large series of other specimens. The species is recognised by a
combination of characters of the carapace, chelipeds, mandibles, and gonopods. Sudanonautes floweri is compared
to related species occurring in Nigeria and Central Africa. The species is found in guinea and woodland savanna
from northern Nigeria to southern Sudan, in tropical rain forest from south-east Nigeria to northern Angola
(including Bioko), and along the Zaire river and its tributaries. Sudanonautes floweri is one of the second
intermediate hosts of the human lung fluke (Paragonimus) in Africa.

INTRODUCTION

Recent major works on the taxonomy of the fresh-water
crabs of Africa (Bott, 1955, 1959, 1964; Monod, 1977, 1980)
recognise three species of Sudanonautes Bott, 1955 — S.
aubryi (H. Milne Edwards, 1853), S. africanus (A. Milne
Edwards, 1869), and S. pelii (Herklots, 1861). Since that time
a number of other species in this genus have been added
(Cumberlidge, 1991, 1993a, b). The subject of the present
work, S. floweri (De Man, 1901), was formerly considered by
both Bott (1955) and Monod (1977, 1980) to be a subspecies
of §. aubryi. Sudanonautes floweri is judged here to be a good
species, and is redescribed from a male syntype from Sudan.

Gonopod 1 of S. floweri is distinct (Fig. 2 d-f), and when
considered in conjunction with other characters of the cara-
pace and sternum (Fig. 1 a-c) and mandibles (Fig. 2 a-c), can
be used to identify the species unequivocally. This is impor-
tant, since S. floweri is one of the four species of Sudanon-
autes that serve as the second intermediate host of the human
lung fluke (Paragonimus) in Nigeria and Central Africa
(Voelker, et al., 1975; Voelker & Sachs, 1977; Nozais, et al.,
1980). However, the ambiguous descriptions of S. floweri and

© The Natural History Museum, 1995

S. aubryi in the literature (A. Milne Edwards, 1853; De Man,
1901; Bott, 1955; Monod, 1977, 1980) have led to the
misidentification of specimens of S. floweri as S. aubryi by
parasitologists (Voelker, et al., 1975, fig. 6; Voelker & Sachs,
1977, fig. 4).

The right mandible and the right first and second gonopods
of the type of S floweri were removed to illustrate these
structures from different angles and under magnification (Fig.
2 a-i). Specimens of S. floweri from Nigeria collected by the
author were either dug from their burrows at the sides of
streams, or were trapped in fishing nets set overnight in
ponds. One specimen (NMU 9.1V.1983) was caught by hand
under rocks in a dried river bed, immediately following the
temporary damming of the river by villagers. Four measure-
ments, carapace length, carapace width, carapace height, and
front width, were recorded from each specimen using digital
callipers. Carapace proportions were calculated according to
carapace length. These data were pooled and used for
descriptions of growth (Fig. 3 a,b). Statistical comparisons
between species were made between sexually mature adults
only (Table 1). The distribution of S. floweri described here is
based on the direct examination of a large number of
specimens from 73 different localities in 9 countries. Litera-

112

ture records are generally not reliable, and have not been
included.

The following abbreviations are used: AMNH, American
Museum of Natural History, New York, NY, USA; FMC,
Field Museum, Chicago, IL, USA; MCZ, Museum of Com-
parative Zoology, Harvard, MA, USA; MNHN, Muséum
National dHistoire Naturelle, Paris; NHM, The Natural
History Museum, London, UK; NNH, Nationaal Natuurhis-
torisches Museum, Leiden, The Netherlands; NMU, North-
ern Michigan University, Marquette, MI, USA; RCM, Royal
Congo Museum, Tervuren, Belgium; SMF, Senckenberg
Museum, Frankfurt am M., Germany; USNM, The United
States National Museum of National History, Smithsonian
Institution, Washington, DC, USA; ZIM, Zoological Insti-
tute and Museum, Hamburg, Germany; ZMB, Museum fiir
Naturkunde der Humboldt-Universitat, Berlin, Germany;
CW = carapace width at widest point; CL = carapace length,
measured along median line; CH = cephalothorax height,
maximum height of cephalothorax; FW = front width, width
of front measured along anterior margin; m = male; f =
female; coll. = collected by.

SYSTEMATIC ACCOUNT

Sudanonautes floweri (De Man, 1901)
(Figs 1 a-i, 2 a-j, 3.a,b, Table 1)

Potamon (Potamonautes) floweri, De Man, 1901:94-98,
100-101, pl. X (fig. 1-7); Rathbun, 1904, pl. XVII (figs 2,
6); Rathbun, 1905:193-195; Rathbun, 1921:406—-410, fig. 6,
pl. XX (fig. 2); Parisi, 1925:99.

Potamon (Potamonautes) aubryi; Balss, 1914, p. 405 (except
ZIM K13557 from Mukonje farm, Cameroon, not Pota-
mon aubryi H. Milne Edwards, 1853).

Potamonautes floweri; Balss, 1936:171, fig. 6.

Potamon floweri; Flower, 1931:734; Chace, 1942:211;
Capart, 1954:834, fig. 21.
Sudanonautes (Convexonautes) aubryi floweri; Bott,

1955:304-306, fig. 65, 100, a-b, pl. XXVIII (fig 2 a-d);
Monod, 1977:1218; Monod, 1980:384—385.

DiAGNOsIs. Mandibular palp 2-segmented; terminal seg-
ment single, undivided, with small hard, hair-fringed flap at
junction between segments (Fig. 2 a-c). Terminal segment of
gonopod | with raised lobe on cephalic part, separated from
caudal part by a conspicuous longitudinal groove; subtermi-
nal segment of gonopod 1 distinctly broadened on outer
margin (Fig. 2 d-f). Conspicuous raised ridges on sternum at
points where chelipeds articulate (Fig. 1 c). Carapace greatly
arched (CH/CL = 0.61, Fig. 1 b), very wide (CW/CL = 1.51,
Fig. 1 a). Vertical suture separating sub-branchial and subor-
bital regions meeting anterolateral margin at base of interme-
diate tooth (Fig. 1 b).

DISTRIBUTION. Nigeria, Cameroon, Bioko (= Fernando
Po), Central African Republic, Sudan, Zaire, Congo, Gabon,
Cabinda, Angola. It is likely that S. floweri is also present in
Equatorial Guinea. Rathbun (1921) and Balss (1936) pro-
vided details of the distribution of the species in Zaire.
Monod (1980) reported S. floweri from the basins of the Nile,
Zaire, Chari, and Lake Chad. The present work adds several
new localities in Nigeria, Bioko, and northern Angola.

N. CUMBERLIDGE

MATERIAL

LECTOTYPE. NHM reg. 1901.8.26.2, lm (CW 48.5, CL 30.5,
CH 17.8, FW 11.7 mm), from Bahr el Gebel, Sudan, coll.
Capt. S. S. Flower, 26.viii.1901. This specimen is here
designated the lectotype of S. floweri. De Man did not specify
types, so the material he examined was syntypic.

OTHERS. The catalogue number of material held at NHM
and NMU begins with the date (year, month, day) of
collection or acquisition. NIGERIA. NHM 1895.5.5.1+4,
Asaba, 150 miles up the Niger, coll. N. H. Crosse. NHM
1905.6.5.98-100, Sapele, junction of Jameson and Aethiop
rivers, coll. Dr. Ansoroye. NHM 1910.4.30.19-22, Oban
southern Nigeria, coll. P. A. Talbot. NHM 1938.7.1, Obubra,
southern Nigeria, coll. I. Sanderson. RCM 52.889, Jos, 1967,
coll. E. B. Guong. NMU 8-12.V.1975, Rosse, at Iguoriokhi,
Bendel State, 1f, 8-12.v.1975, coll. Bruce Powell. NMU
24.1V.1980, first or second roadside culvert, Calabar, 1f (CW
45.5 mm) dug from burrow, coll. J. C. Reid. NMU
30.IV.1982, Ogoja, Cross River State, lm, CW 50 mm, dug
from hole at edge of swamp at Ogoja, rain forest/ woodland
savanna, coll. B. D. Barrett. NMU 4.1.1983, Kaduna river
(year-round flow), Kaduna State, 4m, coll. Fatima Abdulka-
dir. NMU 1.]]1.1983, dug from holes, Kaduna, Kaduna
State, coll. Fatima Abdulkadir. NMU 4.1V.1983, foot of
Obudu plateau, Cross River State, 1m, fast white water, big
rocks, small rocks, sand gravel bottom, caught by villagers,
who dammed stream, dried river bed, caught crabs under
rocks, (with S. africanus, S. granulatus), coll. N. Cumber-
lidge. NMU 12.XII.1983, Yankari Game Reserve, Bauchi
State, Hippo Pool, dug from holes, 1m, 1f, coll. N. Cumber-
lidge. NMU 30.I1V.1984, pond near tributary of river Niger
(20 km east of river), Otta, Benue State, 1f, (CW 54 mm),
coll. John Iyage. NMU 12.VI.1984, dug from holes in banks
of river Samu, tributary of Niger, Pasakwauri, near Kagoro,
Kaduna State, 1m, If, coll. N. Cumberlidge. ZIM K3484,
Benin, 1m, 2f, xii.1909, coll. C. Manger. ZIM K30252, Njaba
creek, 15.11.1973, coll. J. Voelker. ZIM K30314, Cross river,
near Arochukwu, 6.iv.1974, coll. J. Voelker. CAMEROON.
NHM 1938.7.1.9-13, Mamfe, coll. I. Sanderson. NHM
2. VIII.1968, Kindongo, south Bakundu, west Cameroon, in
hole on forest floor about 100 yds from nearest (non-
permanent) water, coll. T. S. Jones. RCM 54.190, Kombe-
tiko, 5 km from Batouri, river Tanadi, 3 specimens, 2.11.1976,
coll. F. Puylaert. RCM 53.389, Olounou, 15-30 specimens,
15-17.vii.1971, coll. F. Puylaert. RCM 54.198, Bissiri May-
erey, 20.i1.1976, coll. F. Puylaert. SMF 2098, Bibundi,
20.viii.1948, coll. Justus Weil. SMF 2868, Bibundi, coll.
Justus Weil. SMF 1787, Victoria, 1907, O. Valley. NMU
24.X.1970, near Mamfe, crossing road by Baduma village,
Kumba-Mamfe road, If, coll. R. H. L. Disney. ZIM K3526,
Im, 1f, 24.xii.1911, coll. Dr. E. Fickendey. ZIM K25447,
Duala, 1m, 4.x.1912, C. Manger. ZIM K30397, Kembong,
near Mamfe, 26.iv.1975, coll. J. Voelker. ZMB 5552, Djeer-
fluss, 1m, coll. Schweinfurth ZMB 7789, Benue,
4-9 viii. 1889, coll. Staudinger. ZMB 8234, Barombi Lake, If,
coll. Zeuner. ZMB 10023, 1f, coll. Preuss. ZMB 10216,
Johann AlbrechtshOhe (modern name unknown, 4°40'N,
9°20’E), 1f, coll. Conradt. ZMB 13718, Victoria, lf, coll.
Deutsche Tiefsee Expedition. ZMB 14342, Douala, 2f,
5.xi.1910, coll. Shaeffer. ZMB 16440, Barombi Station, 1m,
1891, coll. Preuss. ZMB 16947, Douala, 1m, coll. Thorbeke.

REDESCRIPTION OF AFRICAN FRESH-WATER CRAB

ZMB 20161, Buea, 6f, 16.xi.1892, coll. Preuss. ZMB 20195,
Buea, 1m, 1f, coll. Preuss. ZMB 20199, Victoria, 4m, coll.
Preuss. ZMB 21300, river Sanaga, Douala grassland district,
1200 m, 1m, 11.i.1917, coll. Elbert. ZMB 21308, Douala, 1f,
coll. Thorbeke. CENTRAL AFRICAN REPUBLIC. RCM
55.399, Giako river, Bougua, 26.11.1982, coll. L. de Vos & J.
Kempeneeus. RCM 53.086, near Bangui, 22.xii.1967.
SUDAN. NHM 1912.12.31.52, Nyonki Nile, 2030 feet, 1f,
hatchlings, 28.iv.1912, coll. Sir F. T. Jackson. NHM
1912.12.31.53, Gondokoro, 1800 feet, 12.iv.1912, coll. Sir F.
T. Jackson. NHM 1913.9.10.1-3, Lado Nipo, 15 miles north
of Kojokaji, coll. S. S. Flower, zoological survey of Egypt.
NHM 1913.9.10.9-10, new cut to Zeraf, north of Shamfe,
coll. G. W. Graham. NHM 1918.12.13.1-3, Mongalla, coll.
S. S. Flower, Zoological Survey of Egypt. NHM
1922.11.22.7-11, Mongalla, Kanisa, vi.1914, coll. S. S.
Flower, Zoological Survey of Egypt. FMC, 400 miles west of
Juba, 7m, 18f, 22.xii.1884. ZAIRE. RCM 1666, Buta, 1934,
coll. F. Hutsebout. RCM 1.661—1.665, Bambesa, 1.viii.1924,
coll. J. Brejko. RCM 47.495, Epulu, ix.1956, coll. Dr. M.
Poll. RCM 46.159-46.160, Ngense, 1955. RCM
46.161-46.162, Ngense, 1955. MNHN BP5049, river Dougou,
affluent of Uele, 1m, coll. L. Didier, Mission du Bourg
Bozas, 1903. MCZ 10612, Faradje, 1m 1f, 21—23.ix.1915.
SMF 2405, Luki, coll. E. Dartevelle. SMF 2398, Ganda
Sundi, coll. E. Dartevelle. SMF 2385, Faradje, upper Uele,
v.1925, coll. Dr. Schoudeten (exchange, RCM 1083, 1079).
SMF 2383, Bambesa, coll. Krydag. SMF 1782, Duma, coll.
Telinbotz. All of the following AMNH material coll. H.
Lang, J. Chapin, AMNH Congo Expedition. AMNH 3338,
Faradje, 5m, 2f. AMNH 3339, Faradje. AMNH 3355,
Faradje, 3m. AMNH 3357, Faradje, 3m, If, x.1912. AMNH
3358, affluents of Nepoko river, near Gamangui (Ituri For-
est), 3m, 1f. AMNH 3359, Banana, 3m. AMNH 3359, Poko,
1m, 4f, x—xii.1913. AMNH 3377, affluents of Nepoko river,
near Gamangui (Ituri Forest), 3m, 1f. AMNH 3406, south of
Poko, x—xii.1913. AMNH 3409, affluents of Nepoko river,
near Gamangui (Ituri Forest), 1m. AMNH 3422, Van Kerck-
hoverville, 2m, 1f, iv.1912. AMNH 3448, Faradje, 1f, 1911.
AMNH 3453, Poko, Im, 4f, viii. 1909. AMNH 3458, north of
Ganza, If (ovig), 16.xii.1909. AMNH 3462, affluents of the
Tshope river, near Stanleyville. AMNH 3465, Yakukuku,
lm; Garamba, If, xi.1911. BIOKO. NHM 1905.7.19.12, coll.
Fernando Po Exploration Committee. ZMB 20164, 1m, If,
vii.1900, coll. Conradt. GABON. NHM 1908.6.2.22, Lam-
barene, Ogoué river, coll. M. Ansorge. NHM
1908.6.2.23-24, Abanga river, Ogoué river. NHM
1908.6.2.25, Fang forest, Ogoué river, caught on a mountain-
top during heavy tropical rain, 29.iv.1907. NHM
1908.6.2.25a, Masoma river, Ogoué river. AMNH 3367,
Libreville, 5m, 5f, ii.1916, coll. H. Lang, J. Chapin. AMNH
3369, 3m, 2f, 1916, coll. H. Lang, J. Chapin. FMC, Gabon or
Middle Congo, French Equatorial Africa, 1951-1952, coll. H.
A. Beatty. CABINDA. MNHN BPS5048 (1m, CW 54.7, CL
36.0 mm), BP5047 (1f, CW 56.6, CL 39.5 mm) Landana,
Cote de Loango, 4.1x.1898, coll. M. Petit. ANGOLA. NHM
1912.4.2.1-3, Luali river.

DESCRIPTION OF MALE LECTOTYPE

CARAPACE (Fig. 1 a,b). Ovoid, extremely wide, widest in

113

anterior third (CW/CL = 1.51), extremely high, with maxi-
mum height in anterior region (CH/CL = 0.61). Anterior
margin of front straight, curving under, front relatively
narrow, about one-quarter carapace width (FW/CW = 0.25).
Surface of carapace smooth with no deep grooves. Postfron-
tal crest consisting of fused epigastric, postorbital crests,
lateral ends with slight crenulations; mid-groove broad, shal-
low. Postfrontal crest contrasting colour to carapace, located
very close to, almost touching, postorbital margin; laterally,
postfrontal crest meeting, or nearly meeting, anterolateral
margin of carapace at, or near, epibranchial tooth. Exo-
orbital tooth blunt, low, intermediate tooth smaller than
exo-orbital tooth, epibranchial tooth small, low, a granule.
Anterolateral margin of carapace raised and granulated,
bigger granules at epibranchial corner, smaller granules
behind, continuous with posterolateral margin, or curving
slightly inward in hepatic region. Posterior margin about
two-thirds as wide as carapace width.

Face of of carapace with 2 sutures, 1 longitudinal, 1
vertical, dividing face and sides into 3 parts (Fig. 1 b).
Longitudinal suture dividing suborbital, subhepatic regions
from pterygostomial region, beginning under inferior medial
margin of orbit, and curving backward across side. Short,
curving, vertical suture dividing suborbital region from sub-
hepatic region (Fig. 1 b); suture beginning beneath interme-
diate tooth, curving down to meet longitudinal suture,
marked by row of small rounded granules. Third maxillipeds
(Fig. 1 d) filling entire oral field, except for transversely oval
efferent respiratory openings at superior lateral corners; long
flagellum on exopod of third maxilliped; ishium of third
maxilliped smooth, with faint vertical groove; merus with
flanged edges. Mandibular palp 2-segmented, terminal seg-
ment single, undivided, small hard, hair-fringed flap at junc-
tion between segments (Fig. 2 a-c).

PEREIOPODS (Fig. 1 f-i). Chelipeds of lectotype unequal,
right longer, higher than left. Dactylus of right cheliped not
arched, fingers enclosing long interspace when closed, palm
of propodus swollen. Fingers of right cheliped with 4 larger
teeth on lower digit and 4 larger teeth on upper digit,
interspersed with a series of smaller pointed teeth along their
lengths. Inferior margins of merus with rows of small teeth,
cluster of granules surrounding larger tooth at distal end.
Carpus of cheliped with 2 large pointed teeth on inner
margin, second smaller than first. Left cheliped similar to
right, but smaller in all respects. Walking legs (pereiopods
2-5) slender (Fig. 2 j), third pair longest, fourth pair shortest.
Posterior margin of propodus of walking legs serrated, dactyli
of walking legs tapering to point, each bearing rows of
downward-pointing sharp bristles; dactylus of fourth pair
shortest (Fig. 2 }).

UNDERSIDE. First transverse groove on sternum (between
sternal segments 2 and 3) complete; second groove (between
sternal segments 3 and 4) consisting of 2 small notches at sides
of sternum; sternum with conspicuous raised ridges at points
where chelipeds insert (Fig. 1 c). Segments 1-6 of abdomen
four sided, last segment triangular, sides indented, rounded
at distal margin (Fig. 1 e); segment 3 broadest, segments 3-7
tapering inwards (Fig. 1 e).

Terminal segment of gonopod 1 long (2/3 as long as subtermi-
nal segment), first half straight continuation of subterminal
segment, second half curving outward, tapering to pointed tip;
terminal segment with raised lobe on the cephalic part, separated

114 N. CUMBERLIDGE

Fig. 1 Sudanonautes floweri, lectotype, adult male from Bahr el Gebel, Sudan (CW 48 mm), NHM reg 1901.8.26.2. (a), whole animal,
dorsal aspect; (b), carapace, frontal aspect, (c) sternum; (d) left third maxilliped; (e), abdomen; (f), right cheliped, frontal view; (g), left
cheliped, frontal view; (h) carpus, and merus of right cheliped, superior view; (i) carpus, and merus of right cheliped, inferior view. Scale
bar equals 15 mm (h, i), 10 mm (c, f, g), 7.5 mm (a, b, e), and 3.75 mm (d).

from the caudal part by a distinct longitudinal groove visible from tapering diagonally to point at junction with terminal segment,
caudal and superior views (Fig. 2 d,f), not visible from cephalic forming roof of chamber for gonopod 2; subterminal segment
view (Fig. 2 e). Subterminal segment of gonopod 1 broadened beneath flap forming lower floor of chamber for gonopod 2 (Fig.
conspicuously on outer margin, fringed with bristles (Fig. 2 d,e), 2 d). Gonopod 2 (Fig. 2 g-i) shorter than gonopod 1 (reaching
with raised flap extending halfway across segment in distal part, only to the junction between last 2 segments of gonopod 1).

REDESCRIPTION OF AFRICAN FRESH-WATER CRAB

115

Fig. 2. Sudanonautes floweri, lectotype, adult male from Bahr el Gebel, Sudan (CW 48 mm), NHM reg 1901.8.26.2. (a), right mandible
anterior view; (b), right mandible superior view; (c), right mandible posterior view; (d), left gonopod 1, caudal view; (e), right gonopod 1,
caudal view; (f), right gonopod 1, superior view; (g), right gonopod 2, cephalic view; (h), right gonopod 2, caudal view; (i), right gonopod
2, caudal view, detail of terminal segment; (j) left pereiopod 2. Scale bar equals 10 mm (j), 1.5 mm (a-c, d-h), and 0.5 mm (i).

Terminal segment gonopod 2 cup-shaped, with pointed tip,
extremely short, only 1/15 as long as subterminal segment.
Subterminal segment gonopod 2 widest at base, then tapering
sharply inward, forming long, thin, pointed, upright process
supporting short terminal segment.

ADULT FEMALE. Right, left chelipeds same proportions as
male of same size, unequal in both length, height. Mature
female abdomen very wide reaching coxae of pereiopods 2-5.
Segments of female abdomen becoming gradually longer
distally, first, fifth becoming gradually wider, abdomen being

116

widest at groove separating fourth, fifth segments. Sixth
segment, telson together forming near semicircle.

GROWTH (Fig. 3 a,b, Table 1). Measurements and propor-
tions given in Table 1, Fig. 3 a,b. Sexual maturity judged by
development of female abdomen: abdomen of mature
females overlapping bases of coxae of walking legs, pleopods
broad, hair-fringed. Pubertal moult, from pubertal stage to
sexual maturity, occurring between CW 33-42 mm. Largest
known specimen, (male from Cameroon) CW 60.4, CL 39.9.
In Zaire, eggs produced in December; in Sudan, hatchlings
present in April. Dimensions of carapace varying with age
(Fig. 3 a). Relative proportions of carapace width (CW/CL)
and height (CH/CL) of juvenile and pubescent S. floweri not
significantly different (P >0.05) from adults (Fig. 3 b). Front
width becoming smaller with age: FW/CL of adult S. floweri
significantly more narrow (P <0.001) than that of juvenile
and pubescent animals (Fig. 3 b).

CoLour. (Living adults from Ogoja, Nigeria). Dorsal cara-
pace dark purplish brown, with a contrasting yellow-orange
postfrontal crest and yellow orbital border. Flanks light
brown, third maxillipeds pale brown with purple tinge, eye-
stalks white cream, cornea black, sternum and abdomen light
brown with purple tinge. Arthrodial membranes between
joints of chelipeds and pereiopods dark brown; dorsal surface

me
Ee
Be:
pd

Carapace Dimension (mm)

| ae : ae |
ea Sil a
ie a a

10 15 20 25 30 35 40 45
Carapace Length (mm)

N. CUMBERLIDGE

Table 1 Means (+ SE) of ratio of carapace width (CW), carapace
height (CH), and front width (FW), to body size (CL) of adult
Sudanonautes floweri compared to the adults of six closely related
species of Sudanonautes from Nigeria and Central Africa.

CW/CL CH/CL FW/CL
X24 SE X + SE X+SE
Sudanonautes floweri 1SZE= 0/01 _UGL= 010) 10B8e=
0.003
(n = 65)
Sudanonautes aubryi 1.377001 70527 = 10:01 0:38. 01002
(n = 63)
Sudanonautes africanus 1.38° + 0.01 0.43% + 0.003 0.36° + 0.004
(n = 26) (n = 14) (n = 15)
Sudanonautes granulatus 1.42*+ 0.01 0.51°+ 0.01 0.41% + 0.01
(n = 33)
Sudanonautes monodi 1.49? + 0.01 0.58" + 0.004 0.39 + 0.004
(n = 23)
Sudanonautes kagoroensis 1.52 + 0.02 0.50* + 0.01 0.39 + 0.004
(n = 9)
Sudanonautes orthostylis_ 1.45*+ 0.02 0.517 + 0.01 0.467 + 0.01
(n = 10)

Proportion significantly different from that of S. floweri: * = P
<0.001; ° = P <0.01; ° = P <0.05.

Carapace Proportion

0.0
10 15 20 25 30 35 40 45

Carapace Length (mm)

Fig. 3. Comparisons of 108 specimens of Sudanonautes floweri. a, dimensions of the carapace (CW, CH, FW) compared to body size (CL), r
values (all at df = 107) indicate a highly significant correlation (P <0.001) between size classes. b, relative proportions of carapace width
and height (CW/CL, CH/CL) compared to body size (CL), r values (both at df = 107) indicate no significant correlation (P >0.05) between
size classes; relative proportions of front width (FW/CL) compared to body size (CL), r value (at df = 107) indicates a highly significant

correlation (P <0.001) between size classes.

REDESCRIPTION OF AFRICAN FRESH-WATER CRAB

of chelipeds and pereiopods light brown, ventral surface light
brown. Specimens from the Ogoué river, in the Fang forest,
Gabon, with brown-pink carapace, shading into neutral
orange in middle; walking legs orange-vermillion.

VARIATION. The anterolateral margin is raised, marked by a
series of granules or small teeth in some specimens (from
Juba, Shambe, and Kojo-Kaji, Sudan; Ituri forest, Banana,
and Faradje, Zaire; and Ogoja, Kaduna, and Bendel State,
Nigeria). In other specimens (Poko, Zaire; Fernando Po, and
Luali, Angola) the anterolateral margin is completely
smooth. In specimens from Oban, Nigeria, the anterolateral
margin is smooth except for the epibranchial tooth (which is
the size of a large granule), followed by two smaller granules.
It is possible that the above variations of the anterolateral
margin are due to changes associated with growth. For
example, the adult male (CW 53.5 mm) from Juba, Sudan
(FMC) was the only one in which the anterolateral margin
was smooth out of 25 specimens of all sizes. This margin was
toothed or serrated in all the other specimens which mea-
sured CW 48 mm or less. A similar observation was made in
the series of specimens from Cameroon (RCM 53.389),
where the anterolateral margin of a large male (CW 60.4 mm)
was completely smooth, but that of smaller specimens was
granulated. Some specimens from Juba, Sudan, had serra-
tions on the dorsal surface of the dactylus of the cheliped
while other specimens from Juba, and from Nepoko, Zaire,
lacked these serrations.

ECOLOGICAL NOTES

Sudanonautes floweri is a common species of fresh-water crab
widely distributed in Nigeria and Central Africa. It is found in
the moister regions of the woodland and guinea savanna
zones from central Nigeria to southern Sudan. This species is
also found in the humid tropical rain forest habitats in
south-east Nigeria, south Cameroon, Bioko, Central African
Republic, Zaire, Congo, and Gabon. In Nigeria, S. floweri
occurs in the drainage basins of the lower Niger, Benue and
Cross rivers. Specimens collected from Yankari Game
Reserve, Bauchi State, Nigeria were dug from holes at the
base of tufts of tall grass clumps in a marsh at the confluence
of rivers Yashi and Gaji, an area heavily trampled by big
game, especially elephants. Many specimens of S. floweri
were caught on land during heavy tropical rain.

In Sudan, S. floweri lives both in the Yei river basin (a
tributary of the Nile), in the mountainous watershed between
the Nile and the Zaire rivers, and in the level papyrus swamps
(Flower, 1931). In Zaire, S. floweri has been reported from
the lower and middle reaches of the Zaire river, and in the
Ubangi and Uele rivers (Rathbun, 1921). The habitat of S.
floweri in Zaire has also been described by Rathbun (1921),
who summarised the field notes of Herbert Lang. S. floweri
was often found in heaps of rotting vegetation in water
courses, and Lang speculated that this habit may carry the
crabs downstream, explaining (at least in part) the wide
distribution of this species. Predators of S. floweri in the rain
forests of Zaire include crocodiles, monitor lizards (Varanus
niloticus), insectivorous otter shrews (Potamogale velox) and
several small carnivores, chiefly species of mongooses and the
African civet (Viverra civetta).

Sudanonautes floweri is common in shallow streams, rivers,

17

and ponds, and digs burrows near waterways. This species is
also found on land either next to water or some distance
away, since it is capable of breathing air, and functions well
for long periods out of water. The widened and highly arched
carapace, and the lack of teeth on the anterolateral margins
of the carapace of S. floweri are features often associated with
air-breathing and burrow-living. This body shape contrasts
with the more flattened, deep-grooved, and spiny carapace of
the more aquatic river-living species such S. faradjensis
(Rathbun, 1921).

TAXONOMIC REMARKS

The difficulties in distinguishing between S. aubryi and S.
floweri date back to the work of Rathbun (1904, 1905).
Although Rathbun (1905) described S. floweri and S. aubryi
as separate species, her description of P. (P.) aubryi was
based largely on specimens of S. floweri. Specimens from
Cabinda (MNHN BS5048) and Zaire (BP 5049) used by
Rathbun (1905) to describe S$. aubryi have been examined in
the present study and found to be S. floweri. This opinion is
supported by the photographs of the specimens from Zaire
and Gabon provided by Rathbun (1904: TVI, plate IX, figs 5,
8) which closely resemble S. floweri, and which are clearly
different from the photograph of the female type of S. aubryi
(Rathbun, 1904: TVI, plate IX, fig. 3). Unfortunately, Rath-
bun’s (1905) ideas were accepted by later workers with the
result that the descriptions of S. aubryi in Balss (1914, 1929),
Capart (1954), Bott (1955) and Monod (1977, 1980) all refer
to S. floweri rather than to S. aubryi sensu H. Milne Edwards
(1853).

COMPARISONS. Six species of Sudanonautes are sympatric
with S. floweri in Nigeria and Central Africa, viz. S. granula-
tus (Balss, 1929), S. kagoroensis Cumberlidge, 1991, S.
orthostylis Bott, 1955, S. monodi (Balss, 1929), S. aubryi, and
S. africanus. These taxa can be distinguished from S. floweri
as follows. The small hard flap on the mandibular palp at the
junction between the two segments (Fig. 2 a-c), and the
conspicuous raised ridges on the sternum at the points where
the chelipeds insert (Fig. 1 c), distinguish S. floweri from all
other species of Sudanonautes, which lack these features.

In addition, the raised lobe on the cephalic part of the
terminal segment of gonopod 1, separated from the caudal
part by a conspicuous longitudinal groove in S. floweri (Fig. 2
d,f) is also shared, in varying degrees, by S. monodi, S.
kagoroensis and S. granulatus. These three species can be
further distinguished from S. floweri by the following charac-
ters. The raised lobe on the cephalic part of the terminal
segment of gonopod 1 of S. monodi (Cumberlidge, 1991) is
considerably higher than that of S. floweri. In addition, the
carapace of S. monodi is significantly (P <0.001) flatter
(CH/CL S. monodi = 0.52, S. floweri = 0.61), and less wide
(CW/CL S. monodi = 1.37, S. floweri = 1.51) than that of S.
floweri (Table 1). Sudanonautes monodi has patches of
granules on the anterior corners of the carapace behind the
postfrontal crest, while S. floweri lacks these granules.
Finally, S$. monodi is found in dry sudan savanna from
Nigeria to Sudan, while S. floweri is absent from this region;
and S$. monodi is absent from woodland savanna and rain
forest where S. floweri is abundant.

Sudanonautes kagoroensis was described by Cumberlidge

118

(1991), and can be distinguished from S. floweri by examina-
tion of gonopod 1: the raised lobe on the cephalic part of the
terminal segment in S. kagoroensis is lower than that in S.
floweri, and the outer margin of the subterminal segment of
gonopod 1 is slim, while that of S. floweri is conspicuously
broadened (Fig. 2 d, e). Furthermore, the carapace of S.
kagoroensis is significantly (P <0.001) flatter (CH/CL =
0.44) than that of S. floweri (CH/CL = 0.61).

Sudanonautes granulatus was redescribed by Cumberlidge
(1993a) and can be distinguished from S. floweri as follows.
The carapace of S. granulatus is significantly (P <0.001)
flatter (CH/CL S. granulatus = 0.51, S. floweri = 0.61), and
less widened (CW/CL S. granulatus = 1.41, S. floweri = 1.51)
than that of S. floweri (Table 1). In addition, the dactylus of
the major cheliped of the adult male of S. granulatus is
dramatically arched, while that of S. floweri is only moder-
ately arched; the major cheliped of adult male S. granulatus is
as long as, or longer, than the carapace width (Cumberlidge,
1993a), whereas that of S. floweri is shorter (Fig 1 f) than the
carapace width (Fig. 1 a,b).

Three other species, S. aubryi, S. africanus, and S. ortho-
stylis, differ from S. floweri in that the terminal segments of
gonopod 1 of these species lack both a raised cephalic lobe,
and a distinct longitudinal groove in the caudal view. These
three taxa can be further distinguished from S. floweri as
follows. The carapace of S. aubryi is significantly (P <0.001)
flatter (CH/CL S. aubryi = 0.52, S. floweri = 0.61), and less
wide (CW/CL S. aubryi = 1.37, S. floweri = 1.51) than that
of S. floweri (Table 1). In addition, the carapace and post-
frontal crest of S. aubryi are a green-brown colour, whereas
these parts of S. floweri are uniformly red-brown with a
contrasting yellow postfrontal crest.

The terminal segment of gonopod 1 of S. africanus is thin
and needle-like, while that of S. floweri (Fig. 2 d) is wider and
has a distinct groove in the caudal view. The carapace of S.
africanus is significantly (P <0.001) flatter (CH/CL S. africa-
nus = 0.43, S. floweri = 0.61) and less wide (CW/CL S.
africanus = 1.38, S. floweri = 1.51) than that of S. floweri
(Table 1). The carapace of S. africanus has patches of raised
warts, while that of S. aubryi is completely smooth. Finally,
the pollex of the propodus of the major cheliped of S.
africanus has a large and conspicuously flattened tooth, which
is lacking in adult S. floweri.

Sudanonautes orthostylis was redescribed by Cumberlidge
(1993b), and can be distinguished from S. floweri as follows.
The terminal segment of gonopod 1 of S. orthostylis is
straight, lacks a visible groove, and curves outwards sharply
only at the tip, while that of S. floweri bears a longitudinal
groove and curves from the mid point (Fig. 2 d). The
carapace of S. orthostylis is significantly (P <0.001) flatter
(CH/CL S. orthostylis = 0.51, S. floweri = 0.61), and less
wide (CW/CL S. orthostylis = 1.44, S. floweri = 1.51) than
that of S. floweri (Table 1). The frontal margin of S.
orthostylis is significantly (P <0.001) wider than that of S.
floweri (FW/CL S. orthostylis = 0.46, S. floweri = 0.38, Table
1). The dactylus of the major cheliped of S. orthostylis is
broad and flat, while that of S. floweri is narrow. Finally, S.
orthostylis is a much smaller species, maturing at CW 22 mm,
compared to maturity between CW 33-42 mm in S. floweri.

ACKNOWLEDGEMENTS. I am very grateful to Mr. Paul Clark and Ms.
Miranda Lowe (NHM, London), Drs. D. Guinot and J. Forest
(MNHN, Paris), and Mr. Trefor Williams (University of Liverpool,

N. CUMBERLIDGE

UK) for loaning the specimens used in this work. The following
people are thanked for hosting visits: Dr. H. Feinberg, AMNH, New
York, USA; Ms. Ardis Baker Johnston, MCZ, Harvard, MA, USA;
Mr. Paul Clark, NHM, London, UK; Drs. L. Holthuis and C.
Fransen, NNH, Leiden, The Netherlands; Dr. R. Joqué, RCM,
Tervuren, Belgium; Dr. R. Manning, USNM, Washington DC.; Drs.
H.-G. Andres and G. Hartmann, ZIM, Hamburg, Germany; Dr. H.
Gruner, ZMB, Berlin, Germany; and the staff of the FMC, Chicago,
USA. I especially thank artist Mr. Jon C. Bedick of Northern
Michigan University, USA, for all of the illustrations used in this
paper. Part of this work was supported by a Faculty Grant from
NMU, Marquette, MI, USA.

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EEFERENCES

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Bull. nat. Hist. Mus. Lond. (Zool.) 61(2): 121-137

Association of epaxial musculature with
dorsal-fin pterygiophores in acanthomorph
fishes, and its phylogenetic significance

RANDALL D. MOOL
Milwaukee Public Museum, 800 West Wells St., Milwaukee, WI, U.S.A. 53233-1478

ANTHONY C. GILL
Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD

Synopsis. A survey of acanthomorphs reveals that epaxialis attachments to distal radials or the distal tips of
proximal-middle pterygiophores have a relatively restricted distribution. Four basic morphotypes are recognized: Type 0—
no distal insertions of epaxialis (lampridiforms, polymixiiforms, basal paracanthopterygians, zeiforms, beryciforms,
smegmamorphs, pleuronectiforms and many perciforms); Type 1 — partially separate epaxialis slip(s) inserting on to
dorsoposterior and dorsolateral processes of proximal-middle and/or distal radials (batrachoidids [Paracanthopterygiil,
scorpaeniforms, and among perciforms in blennioids, most cirrhitoids, apogonids, centrogeniids, latine centropomids,
grammatids, haemulids, percids, serranids, champsodontids and cheimarrhichthyids); Type 2 — insertions of epaxialis to
distal portions of pterygiophores without separate slips (possibly basal tetraodontiforms, various perciform taxa including
callionymoids, notothenioids, zoarcoids, and some cirrhitids, labrids, percoids and trachinoids); Type 3 — completely
separate slip of muscle dorsal to the main body of the epaxialis inserting on to anterior pterygiophore shaft with dorsal
insertions on to more posterior spine-bearing pterygiophores, and the first ray-bearing pterygiophore, then becoming
continuous with the supracarinalis posterior (percoid family Mullidae). Type 0 is considered to be plesiomorphic, and the
remaining morphologies apomorphic. Their phylogenetic significance is discussed in the context of other characters.
Among our conclusions, the Scorpaeniformes is awarded subordinal status within the Perciformes, and the centropomid

Issued 30 November 1995

Latinae is given full familial status.

INTRODUCTION

Within the last five years, there has been renewed interest in
higher relationships among acanthomorphs. The recent pub-
lication of the Symposium on Phylogeny of Percomorpha
(Johnson & Anderson, 1993) and other contributions
(Stiassny, 1990; Stiassny & Moore, 1992) have shifted the
focus somewhat from phylogenetic work on individual fami-
lies to broader studies involving interrelationships of subor-
ders and orders. Such studies are hampered by the difficulties
inherent in examining large numbers of taxa, determining
appropriate character complexes, and interpreting homolo-
gies among the variation within those complexes. In many
instances, characters are too complex or difficult to survey
resulting in an incomplete understanding of their distribution
within the included groups. During the course of investiga-
tions on the relationships among pseudochromoids (sensu
Mooi, 1990), we began surveying the relation of dorsal
epaxial myology to the dorsal-fin pterygiophores. Dorsal
epaxial myology appears to exhibit limited but sufficient
variation over a broad range of taxa and the character states
are simple enough to suggest it to be of high potential for
phylogenetic analysis of higher relationships among acantho-
morphs.

Epaxial muscles, the dorsal component of the body muscu-
lature, have received little attention from fish systematists.
Although some studies have used variation in the anterior
insertions of epaxial slips on to the head (e.g., Mooi, in press;

© The Natural History Museum, 1995

Stiassny, 1990), few workers using myological features have
surveyed this muscle group (Winterbottom, 1974a for a
review). Mok et al. (1990) were the first to report variation in
the relationship of the epaxial musculature with the dorsal-fin
pterygiophores. They found that in two percoid families, the
Grammatidae and Opistognathidae, the epaxial muscles
insert on to the distal portions of anterior dorsal-fin pterygio-
phores, and interpreted this as evidence for uniting the two
taxa as sister groups.

Our continuing studies on the phylogenetic positions of the
Grammatidae, Opistognathidae and other pseudochromoid
families have failed to provide corroborating evidence for a
sister-group relationship between the Grammatidae and
Opistognathidae. Moreover, a preliminary survey of epaxial
morphology in perciforms revealed that the reportedly
unique association of epaxial musculature with dorsalfin
pterygiophores described by Mok et al. (1990) is more widely
distributed (Gill & Mooi, 1993: 333). Here we present an
extensive survey of acanthomorph taxa, and show that,
despite having a wider distribution than indicated by Mok et
al., epaxial muscle/dorsal-fin pterygicphore associations nev-
ertheless appear to be relatively restricted within acantho-
morphs, and exhibit a number of recognizable morphologies.
We explore the possible phylogenetic significance of the
distribution of epaxial muscle insertions to dorsal-fin ptery-
giophores and their homology.

122

METHODS AND MATERIALS

Epaxial musculature/dorsal-fin pterygiophore associations
were studied in alcohol-stored specimens. An incision was
made through the skin along the length of the fish between
one third to one half the distance from the base of the dorsal
fin and the midlateral septum. The incision ran from the skull
to beneath the segmented-ray portion of the dorsal fin. The
skin was either removed or folded dorsally to expose the
underlying muscle. The inclinatores dorsales usually lifted up
with the skin, or were removed individually to permit exami-
nation of the epaxial muscles and the dorsal portions of the
pterygiophores. When appropriate, epaxial fibres were
traced anteriorly or posteriorly to ascertain their association
with the supracarinalis muscle system. The insertions of
epaxial fibres to pterygiophores were often re-examined on
cleared and stained specimens and dry skeletons in the
collections of the American Museum of Natural History,
Milwaukee Public Museum, National Museum of Natural
History, and The Natural History Museum. These specimens
are not listed in Table 1. Illustrations of muscles were made
with a camera lucida attached to a binocular dissecting
microscope.

Material dissected for myological observations is listed in
Table 1. All species examined during the study are repre-
sented in this list, although in many cases, multiple specimens
were examined, occasionally from different lots, and some-
times from museum collections other than those listed, par-
ticularly the Field Museum of Natural History and Royal
Ontario Museum. A complete list can be provided by the
authors. Institutional codes follow Leviton et al. (1985).

RESULTS

Many (if not most) fishes have some epaxial fibre insertion
near the proximal ends or near the middle of the dorsal-fin
pterygiophores, whereas some taxa have epaxial muscle
insertions on to the distal ends of the pterygiophores. We
recognize four morphotypes of epaxial musculature, Types 0
to 3. The consecutive numbering of the morphological types
is not meant to imply character transformations; the morpho-
types do not necessarily form a polarized transformation
series. The vast majority of acanthomorph fishes (including
putative basal taxa) exhibit an apparently primitive condition
of the epaxial muscles, Type 0, with no attachment to the
distal parts of the dorsal-fin pterygiophores, and with the
musculature usually lying well below the dorsal tips of the
pterygiophores (Fig. 1; Table 1).

Of those taxa that do exhibit insertions on to the distal
portions of the pterygiophores, epaxial fibres rarely insert on
to pterygiophores other than those bearing non-segmented
rays (spines), except where these ray elements are inter-
preted as secondarily derived from spines (e.g., pseudochro-
mids, zoarcoids, pleuronectiforms). In one scorpaeniform
and a perciform genus as discussed below, and probably the
paracanthopterygian Opsanus beta, there is insertion on
primary ray-bearing pterygiophores. Among the taxa with
dorsal insertions of epaxial fibres to spine-bearing pterygio-
phores, there are three recognizable morphologies. Although
these morphologies can be defined by specific taxa, their

R.D. MOOI AND A.C. GILL

apparent differences become somewhat subjective at the ends
of their respective morphological spectra.

Type 1 is characterized by a partially separate muscle mass
or series of slips of muscle fibres that insert on to the
dorsoposterior and dorsolateral processes of the proximal-
middle and/or distal radials of the pterygiophores. At least
some fibres originate from the main body of epaxial muscle,
but in extreme cases the dorsal muscle mass is detached
between successive myosepta, and anteriorly there can be an
elongate separate slip of muscle to an anterior pterygiophore
(Fig. 2). We observed this morphotype in a single paracan-
thopterygian species (Opsanus beta) (Fig. 3), blennioids,
most cirrhitoids, seven percoid families (Apogonidae, Cen-
trogeniidae, Centropomidae, Grammatidae, Haemulidae,
Percidae, and Serranidae) and two trachinoid families
(Champsodontidae and Cheimarrhichthyidae) among the sur-
veyed perciforms (Figs 1, 4-5, 12-17), and all but one
examined scorpaeniform (Figs 6-8) (Table 1).

Among examined scorpaeniforms with Type 1, Normanich-
thys crockeri exhibits a unique morphology (Fig. 8). The
epaxial muscles insert on to the lateral processes of the first
nine or ten pterygiophores as a separate mass of muscle.
Posterior to the first dorsal fin, epaxial fibres attach directly
to spineless (naked) pterygiophores and these fibres are not
arranged as a separate muscle mass. A separate muscle mass
is also present at the second dorsal fin, with insertions on to
those pterygiophores bearing segmented rays. This gradually
tapers out posteriorly and merges with the main body of
epaxial muscle. Other scorpaeniform and percoid taxa exhib-
iting Type 1 are quite consistent in their epaxial morphology;
even among unusual taxa such as Aploactis (a scorpaeni-
form), which has its dorsal fin placed far anteriorly over the
skull, a narrow tendon extends from the epaxial to insert on
to the third dorsal-fin pterygiophore. Differences arise in the
degree of muscle separation, size of the anterior slip, on to
which pterygiophores the muscle inserts, and on to which
radials of the pterygiophores the insertion occurs (cf. Figs
2-8).

Species with a Type 2 epaxial morphology lack the obvious
separation of the dorsal muscle bundle that inserts on to the
distal portions of the pterygiophores, and the anterior slip is
always absent. The insertions resemble sheets hanging on a
clothes-line, draping from one pterygiophore to the next (Fig.
9). In some taxa, the insertions are primarily via long
tendons, and the muscle fibres themselves are relatively
distant from the dorsal parts of the pterygiophores (Fig. 10).
In most elongate taxa, the muscles are much more dorsally
situated and the tendons are not as obvious. This morphology
is found in various perciform taxa, including some members
of the Cirrhitidae, Labridae, Percoidei, and Trachinoidel,
and all of the few examined members of the Callionymoidei,
Notothenioidei and Zoarcoidei (Table 1). The Tetraodonti-
formes have a modified condition of this basic morphology
which will be discussed below.

A Type 3 epaxial morphology was found only in the family
Mullidae (Fig. 11; Table 1). This type consists of a few
epaxial fibres inserting on to an anterior pterygiophore
relatively ventrally and on to a lateral wing along the main
shaft rather than on to a dorsal posterolateral process. A
completely separate slip of muscle sits dorsal to the epaxial
muscle and inserts on to the anterior pterygiophore and only
the posterior pterygiophores of the first dorsal fin. It extends
further posteriorly, inserting on to the first pterygiophore of
the second dorsal, and gradually narrows posteriorly, insert-

EPAXIAL MUSCLES AND ACANTHOMORPH RELATIONSHIPS

Table 1 List of taxa examined for epaxial muscle morphology. Morphological types: 0 — no association with distal tips of dorsal-fin
pterygiophores; 1 — partially separate muscle block or series of slips of muscle fibers that insert on to the dorsoposterior and dorsolateral
processes of the proximal-middle and/or distal radials of the dorsal-fin ptergygiophores; 2 — insertions to the distal portions of the
pterygiophores without an obvious separation from the main muscle body and with no separate anterior slip; 3 — completely separate slip of
muscle dorsal to the main body of the epaxialis inserting on to an anterior pterygiophore shaft with dorsal insertions on to more posterior
spine-bearing pterygiophores and the first pterygiophore bearing a segmented ray, then becoming continuous with the supracarinalis
posterior. Orders are listed phylogenetically following Johnson & Patterson (1993); suborders, families, and species are listed alphabetically
within orders. /ncertae sedis genera of Percoidei are listed alphabetically among families.

123

Taxon, Catalogue No., SL (mm)

LAMPRIDIFORMES
Veliferidae
Velifer hypselopterus, AMNH 49575, 118.0
POLYMIXIIFORMES
Polymixiidae
Polymixia lowei, AMNH 10116, 131.0
PARACANTHOPTERYGII
Aphredoderidae
Aphredoderus sayanus, AMNH 50907, 53.5
Batrachoididae
Opsanus beta, AMNH 52369, 115.0
Brotulidae
Dinematichthys sp., USNM 297347, 88.5
Gadidae
Urophycis floridanus, MPM 8409, 76.6
Lotidae
Lota lota, MPM 28380, 100.0
Percopsidae
Percopsis omiscomaycus, MPM 14060, 77.1
ZEIFORMES
Parazenidae
Parazen pacificus, AMNH 29459, 116.5
BERYCIFORMES
Holocentridae
Myripristis pralinus, USNM 285922, 113.5
Sargocentron vexillarus, MPM 30099, 56.7
Trachichthyidae
Hoplostethus mediterraneus, AMNH 49700, 117.0
SYNBRANCHIFORMES
Mastacembelidae
Caecomastacembelus congricus, AMNH 6157, 145.0
Mastacembelus armatus, FMNH 68484, 190.0
ELASSOMATIFORMES
Elassomatidae
Elassoma okefenokee, MPM 28810, 20.5
E. zonatum, MPM 14480, 28.5
GASTEROSTEIFORMES
Aulostomatidae
Aulostomus maculatus, MPM 25182, 174.2
Aulorhynchidae
Aulorhynchus flavidus, AMNH 58939, 123.0
Gasterosteidae
Culaea inconstans, MPM 26675, 50.2
Gasterosteus aculeatus, AMNH 37959, 54.0
Macrorhamphosidae
Macrorhamphosus scolopax, AMNH 84458, 85.5
MUGILIFORMES
Mugilidae
Agonostomus monticola, MPM 13806, 41.0
Mugil cephalus, USNM 152118, 93.2
M. curema, MPM 6817, 56.4
ATHERINIFORMES
Atherinidae
Atherinomorus stipes, MPM 30102, 53.4
Menidia beryllina, MPM 30404, 63.0
CYPRINODONTIFORMES
Cyprinodontidae
Cyprinodon variegatus, MPM 28940, 45.6
Fundulidae
Fundulus catenatus, MPM 15271, 70.8

Type

oo

==)

oo

ooo

Taxon, Catalogue No., SL (mm)

Poeciliidae
Poecilia mexicana, MPM 8283, 55.4
DACTYLOPTERIFORMES
Dactylopteridae
Dactylopterus volitans, USNM 307210, 59.5
PERCIFORMES
Acanthuroidei
Acanthuridae
Acanthurus triostegus, USNM 139750, 73.5
Ephippididae
Chaetodipterus zonatus, USNM 131415, 48.9
Scatophagidae
Scatophagus argus, BMNH 1976.4.13:2-7, 48.3
Anabantoidei
Anabantidae
Anabas testudineus, AMNH 13766, 65.0
Badidae
Badis badis, USNM 89076, 26.8
Belontiidae
Belontia signata, USNM uncat., 64.4
Macropodus opercularis, AMNH 10641, 38.7
Channidae
Channa arga, AMNH 79406, 121.0
C. obscurus, FANH 70260, 136.0
Nandidae
Monocirrhus polyacanthus, USNM uncat., 68.0
Nandus nebulosus, USNM 230323, 47.7
Polycentrus schomburgki, USNM 226071, 41.7
Pristolepidae
Pristolepis fasciata, USNM 305711, 75.7
Blennioidei
Blenniidae
Entomacrodus nigricans, MPM 18256, 55.4
Hypleurochilus aequipinnis, MPM 23034, 28.2
Ophioblennius atlanticus, MPM 24880, 52.4
Scartella cristata, MPM 18231, 62.0
Chaenopsidae
Acanthemblemaria greenfieldi, MPM 24876, 30.4
A. aspera, MPM 29983, 24.7
Emblemaria pandionis, BMNH 1938.2.2:2, 39.3
Stathmonotus gymnodermis, MPM 24881, 23.6
S. stahli, BMNH 1939.5.12:183-189, 18.8
Clinidae
Clinoporus biporosus, BMNH 1935.4.29:1-8, 89.5
Clinus cottoides, BMNH 1887.4.16:3-5, 93.0
Dactyloscopidae
Dactyloscopus tridigitatus, MPM 24981, 60.0
Gillellus uranidea, MPM 30131, 29.5
Labrisomidae
Labrisomus bucciferus, MPM 31163, 57.0
L. nuchipinnis, MPM 18253, 82.0
Malacoctenus gilli, MPM 24947, 49.1
M. versicolor, MPM 22469, 36.0
M. zonifer, BMNH 1861.8.13:33, 47.3
Paraclinus fasciatus, MPM 25004, 36.2
Starksia lepicoelia, MPM 29994, 23.5
Tripterygiidae
Enneanectes atrorus, MPM 30216, 21.0
E. boehlkei, MPM 11572, 18.2
E. pectoralis, MPM 22463, 26.5
Lepidoblennius haplodactylus, BMNH 1890.9.23, 63.6

Type

124

Taxon, Catalogue No., SL (mm)

Callionymoidei
Callionymidae
Synchiropus splendidus, MPM uncat., 59.2
Gobiesocidae
Gobiesox strumosus, AMNH 86887, 58.5
Carangoidei
Carangidae
Caranx latus, MPM 13771, 119.0
Oligoplites saurus, MPM 6364, 77.2
Selene vomer, MPM 2273, 75.1
Trachinotus rhodopus, MPM 6369, 107.0
Nematistiidae
Nematistius pectoralis, MPM 6367, 215.0
Cirrhitoidei
Aplodactylidae
Aplodactylus punctatus, USNM 227298, 58.0
Cheilodactylidae
Cheilodactylus variegatus, USNM 77574, 58.0
C. zonatus, USNM uncat., 73.5
Chironemidae
Chironemus marmoratus, ROM 40360, 125.4
Cirrhitidae
Amblycirrhitus bimacula, MPM 13509, 56.9
Cirrhitichthys oxycephalus, ROM 60291, 55.2
Cirrhitops hubbardi, ROM 59830, 64.5
Cirrhitus pinnulatus, ROM 47702, 101.0
Neocirrhitus armatus, ROM 59838, 44.1
Paracirrhites arcatus, MPM 13587, 66.7
Gobioidei
Butidae
Butis amboinensis, MPM uncat., 57.2
Eleotrididae
Eleotris pisonis, USNM 314448, 77.5
Gobtidae
Awaous taiasica, MPM 6811, 92.1
Bathygobius soporator, MPM 18232, 80.0
Odontobutidae
Micropercops sp., AMNH 10441, 44.4
Xenisthmidae
Xenisthmus balius, USNM 326758, 26.1
Labroidei
Cichlidae
Cichlasoma salvini, MPM 22851, 48.8
Etroplus suratensis, USNM 301169, 69.5
Embiotocidae
Rhacochilus argyrosomus, USNM 53969, 45.7
Labridae
Bodianus bilunulatus, MPM 13518, 76.3
B. diana, USNM 232355, 52.0
Cheilinus oxycephalus, USNM 262088, 62.1
Cheilio inermis, MPM 13369, 88.6
Choerodon graphicus, USNM 218548, 60.2
Coris variegata, USNM unceat., 86.0
Halichoeres bivitattus, MPM 8524, 73.6
Hemipteronotus martinicensis, USNM 37075, 85.0
Labroides dimidiatus, MPM uncat., 51.7
Sparisoma rubripinnis, MPM 30040, 62.6
Tautoga onitis, USNM 118352, 53.2
Thalassoma duperryi, MPM 13403, 77.7
T. lutescens, USNM 112696, 82.0
Pomacentridae
Abudefduf saxatilis, USNM 275040, 63.5
Amphiprion melanopus, USNM 309519, 68.0
Lepidozygus tapeinosoma, USNM 275893, 51.5
Notothenioidei
Nototheniidae
Notothenia sima, AMNH 5003, 82.5

Type

(SS) (=)

NNONNNNNOCNNCS (>)

Soe)

R.D. MOOI AND A.C. GILL

Taxon, Catalogue No., SL (mm)
Percoidei
Acropomatidae
Malakichthys griseus, USNM 184143, 60.2
Ambassidae
Ambassis sp., USNM 223376, 37.8
Chanda ranga, BMNH 1938.12.22:132-141, 40.0

Apogonidae

Apogon angustatus, USNM 261750, 57.0

Apogonichthys ocellatus, AMNH 33808, 43.0

Cheilodipterus macrodon, AMNH 33714, 68.0
Bathyclupeidae

Bathyclupea malayana,BMNH 1982.9.6:106-107, 117.0
Callanthiidae

Callanthias australis, AMS 1.18709-002, 88.0

C. platei, USNM 307594, 93.0
Caproidae

Antigonia eos, MPM 13598, 71.3

Capros aper, BMNH 1963.5.14:230-239, 43.5
Centrarchidae

Lepomis gibbosus, MPM 28675, 56.2

Micropterus salmoides, MPM 20246, 62.2
Centrogeniidae

Centrogenys vaigensis, USNM 245612, 70.0
Centropomidae

Centropomus armatus, USNM uncat., 108.7

C. ensiferus, ROM 61657, 47.7

C. pectinatus, ROM 61664, 61.0

C. undecimalis, ROM 40904, 118.5
Cepolidae

Cepola rubescens, BMNH 1970.4.18:3, 438.0

Owstonia totomiensis, BMNH 1986.10.6:61, 91.0
Chaetodontidae

Chaetodon multicinctus, MPM 13556, 89.7

C. miliaris, MPM 13466, 56.0
Datnioides quadrifasciatus, USNM 297256, 120.0
Dinolestidae

Dinolestes lewinii, USNM 59932, 138.5
Enoplosidae

Enoplosus armatus, USNM 48808, 77.5
Gerreidae

Eucinostomus gula, USNM 43216, 45.0
Glaucosomatidae

Glaucosoma scapulare, AMS 1.27325-002, 45.1
Grammatidae

Gramma linki, AMNH 35776, 36.3

G. loreto, MPM 15612, 50.4

Lipogramma anabantoides, AMNH 33061, 16.8

L. trilineata, FANH 95658, 24.2
Haemulidae

Anisotremus scapularis, USNM 127982, 55.8

Conodon nobilis, MPM 13778, 104.5

Haemulon aurolineatum, MPM 23228, 64.2
Hapalogenys sp., BMNH 1984.1.13:76-82, 55.0
Hemilutjanus microphthalmus, USNM 77623, 138.0
Kuhludae

Kuhlia rupestris, USNM 184110, 82.0
Kurtidae

Kurtus gulliveri, USNM 217310, 128.0
Kyphosidae

Girella tricuspidata, USNM 269547, 99.5

Sectator oxyurus, USNM 288880, 75.1
Lactariidae

Lactarius delicatulus, BMNH 1895.2.28:51, 87.0
Lateolabrax japonicus, USNM 64630, 87.0
Latidae

Lates albertianus, ROM 26537, 141.1

L. calcarifer, BMNH 1898.12.24:2, 113.5

L. mariae, ROM 28140, 125.2

L. niloticus, BMNH 1907.12.2:2959-2968, 48.5

Type

Taxon, Catalogue No., SL (mm)

Psammoperca waigiensis, BMNH 1933.3.11:312, 118.0
Lethrinidae

Lethrinus lentian, BMNH 1932.7.29:82-83, 70.8
Lobotidae

Lobotes surinamensis, USNM 156452, 46.6
Lutjanidae

Lutjanus griseus, MPM 8542, 48.1

L. kasmira, USNM 183109, 98.8
Malacanthidae

Caulolatilus affinis, USNM 211424, 104.5
Monodactylidae

Monodactylus argenteus, MPM 31026, 33.2
Moronidae

Dicentrarchus labrax, BMNH 1987.2.22:1—12, 42.5

Morone chrysops, MPM 4569, 78.3
Mullidae

Mulloidichthys martinicus, MPM 5321, 86.0

Parupeneus multifasciatus, MPM 13530, 79.0

Upeneus maculatus, USNM 126150, 76.0
Nemipteridae

Pentapodus porosus, BMNH 1984.8.20:27, 62.0
Notograptidae

Notograptus sp., USNM 173797, 174.0
Opistognathidae

Opistognathus maxillosus, MPM 30098, 98.3
Oplegnathidae

Oplegnathus fasciatus, BMNH 1905.6.6:154-161, 126.0
Ostracoberycidae

Ostracoberyx sp., USNM 307282, 83.0
Pempherididae

Parapriacanthus ransonneti, MPM 31028, 58.2

Pempheris schomburgki, FMNH 93774, 52.3
Percichthyidae

Percichthys altispinnis, USNM 163382, 70.1

Percilia gillissi, USNM 84341, 60.0
Percidae

Etheostoma nigrum, MPM 22420, 56.3

Perca flavescens, MPM 25409, 79.0

Percina maculata, MPM 20880, 76.7

Stizostedion canadense, MPM 6015, 86.7
Pholidichthyidae

Pholidichthys leucotaenia, USNM 289924, 77.0
Plesiopidae

Acanthoclinus fuscus, USNM uncat., 77.7

Assessor macneilli, USNM 295659, 40.5

Belonepterygion fasciolatum, USNM 273813, 34.0

Calloplesiops altivelis, USNM 261333, 60.4

Plesiops coeruleolineatus, USNM uncat., 45.5

Trachinops taeniatus, USNM 274580, 37.2
Polynemidae

Polydactylus approximans, USNM uncat., 60.2
Polyprionidae

Polyprion americanus, BMNH 1845.6.22:11, 190.0
Pomacanthidae

Centropyge bispinosus, USNM 259696, 61.5
Pomatomidae

Pomatomus saltatrix, BMNH 1967.2.1:80—-85, 74.7
Priacanthidae

Priacanthus hamrur, USNM 289285, 72.5
Pseudochromidae

Haliophis guttatus, USNM uncat., 137.5

Labracinus cyclophthalmus, USNM 309335, 85.8

Ogilbyina queenslandiae, USNM 290792, 59.3

O. salvati, USNM 278149, 50.3

Pseudochromis elongatus, USNM 290784, 35.6

P. fuscus, USNM 290345, 56.5

‘Pseudochromis’ diadema, USNM 290591, 32.9
Sciaenidae

Aplodinotus grunniens, MPM 16805, 66.4

EPAXIAL MUSCLES AND ACANTHOMORPH RELATIONSHIPS

Type

NNNNYNN WY

i=)

Taxon, Catalogue No., SL (mm)

Bairdiella chrysura, MPM 8954, 100.0

Cynoscion regalis, MPM 8969, 94.9

Equetus acuminatus, MPM 8522, 90.1

Leiostomus xanthurus, MPM 8934, 87.1

Menticirrhus littoralis, MPM 8443, 68.7

Micropogonias undulatus, USNM 142675, 67.5

Sciaenops ocellata, MPM 30424, 61.5

Stellifer lanceolatus, MPM 8936, 57.9
Serranidae

Alphestes afer, USNM 235696, 89.0

Anyperodon leucogrammus, USNM uncat, 103.5

Centropristis philadelphicus, USNM 142813, 75.8

Chelidoperca sp., USNM 322386, 80.0

Diplectrum macropoma, USNM 211397, 129.0

Epinephelus merra, USNM 309689, 75.5

Grammistes sexlineatus, USNM 166994, 62.0

Hypoplectrodes sp., USNM 198811, 67.5

H. maccullochi, USNM 42039, 102.1

Hypoplectrus puella, MPM 23461, 92.3

Liopropoma rubre, MPM 25083, 41.0

L. sp., USNM 322359, 76.5

Mycteroperca florida, USNM 176238, 59.7

Niphon spinosus, USNM 59739, 130.0

Paralabrax clathratus, USNM 54807, 53.0

Plectranthias nanus, USNM 288812, 24.9

Pseudanthias taeniatus, USNM 279782, 54.5

P. thompsoni, USNM uncat., 118.0

Pseudogramma sp., USNM 245340, 42.8

Serranus hepatus, USNM uncat., 73.0

S. tigrinus, MPM 30183, 58.3
Sillaginidae

Sillago cilliata, USNM 207647, 72.6
Sinipercidae

Coreoperca kawamebari, USNM 71331, 32.3

Siniperca chautsi, USNM 87082, 93.2
Sparidae

Diplodus bermudensis, MPM 18228, 76.5
Symphysanodontidae

Symphysanodon berryi, USNM 289922, 85.5
Synagrops bella, USNM 156955, 75.5
Terapontidae

Terapon jarbua, USNM uncat., 80.0
Toxotidae

Toxotes jaculator, USNM uncat., 45.0

Scombroidei

Scombridae

Scomber japonicus, AMNH 74945, 149.0
Sphyraenidae

Sphyraena barracuda, MPM 11496, 93.0
Trichiuridae

Trichiurus lepturus, MPM 8430, 316.0

Scorpaenoidei

Agonidae

Agonus decagonus, USNM 165146, 132.5
Anoplopomatidae

Anoplopoma fimbriata, USNM 208296, 123.0
Aploactinidae

Aploactis milesii, USNM 59980, 121.0
Bathylutichthyidae

Bathylutichthys taranetzi, BMNH 1994.7.22:1, 100.5
Caracanthidae

Caracanthus maculatus, USNM 140990, 34.5
Congiopodidae

Alertichthys blacki, USNM 318386, 80.0
Cottidae

Ascelichthys rhodorus, BMNH 1881.3.22:57-63, 50.0

Centrodermichthys analis, BMNH 1890.11.15:105, 56.7

Cottus bairdi, MPM 5878, 70.8

125

Type

NNONCNNN

—s— se ee ee ee ee ee es

126
Taxon, Catalogue No., SL (mm) Type
C. perplexus, USNM 258839, 51.5 1
Icelus hamatus, BMNH 1877.5.13:7-9, 60.3 1
Myoxocephalus scorpis, BMNH 1981.2.10:629, 50.5 1
Taurulus bubalis, BMNH 1981.2.20:776-794, 38.5 1
Cottocomephoridae
Cottocomephorus grewingkii, USNM 222075, 100.0 1
Cyclopteridae
Cyclopterus lumpus, USNM 197582, 83.8 1
Hoplichthyidae
Hoplichthys langsdorfi, USNM 309447, 123.0 1
Liparididae
Liparis agassizii, USNM 74697, 117.5 1
L. liparis, BMNH 1971.2.16:1757-1760, 99.5 1
Paraliparis hystrix, BMNH 1992.10.20:43-48, 87.8 1
Normanichthyidae
Normanichthys crockeri, USNM 176507, 56.7 1
Pataecidae
Aetapcus maculatus, BMNH uncat., 118.0 1
Pataecus fronto, BMNH 1914.8.20:282, 159.0 1
Platycephalidae
Thysanophrys japonica, USNM 70735, 119.0 1
Psychrolutidae
Cottunculus microps, BMNH 1981.3.16:550-553, 87.5 1
Psychrolues zebra, BMNH 1986.7.12:193, 41.5 1
Scorpaenidae
Pterois radiata, USNM 140491, 64.8 1
Scorpaena sonorae, USNM 59463, 67.6 1
Sebastes alutus, USNM 72461, 80.0 1
Triglidae
Bellator militaris, USNM 114793, 83.0 1
Stromateoidei
Stromateidae
Peprilus burti, MPM 8291, 92.4 0
Trachinoidei
Ammodytidae
Ammodytes americanus, AMNH 36780, 72.0 0
A. hexapterus, FMNH 80613, 106.0 0
A. lanceolatus, FMNH 34257, 177.0 0
A. personatus, USNM 104499, 86.0 0
Champsodontidae
Champsodon sp., USNM 150556, 64.2 1
Cheimarrhichthyidae
Cheimarrichthys fosteri, AMNH 98274, 71.0 1
Chiasmodontidae
Chiasmodon sp., USNM 186139, 110.0 0
Dysalotus alcocki, MCZ 60806, 112.0 0

ing on to no other pterygiophores, but becoming continuous
with the supracarinalis posterior. With such minimal fibre
sharing of this elongate separate slip with the epaxial, it
appears that the separate slip is likely to be a modified
supracarinalis posterior or supracarinalis medius. Although
no other taxon was found exhibiting this morphology, a few
taxa do have a tendon extending to the supracaranalis poste-
rior from the last fibres of the epaxial section that inserts on
to the pterygiophores. We observed this condition in the
cirrhitid Paracirrhites arcatus, some labrids (including Spari-
soma and Halichoeres), as well as some blennioids. This
tendon can be difficult to detect, and could be present in
other taxa, although no trace of this feature was found in
serranids or scorpaeniforms.

R.D. MOOI AND A.C. GILL

Taxon, Catalogue No., SL (mm) Type
Kali normani, USNM 207614, 159.6 0
Pseudoscopelus sp., ARC 8706465, 57.0 0

Creediidae
Crystallodytes cookei, FANH 63619, 41.0 2
Limnichthys fasciatus, AMNH 57282, 45.5 2
Percophididae
Bembrops anatirostris, AMNH 83323, 170.0 Z
B. gobioides, FANH 67070, 112.0 2
Pinguipedidae
Parapercis cephalopunctatus, FMNH 72471, 108.0 z
P. montillai, AMNH 50585, 94.0 2
Uranoscopidae
Kathetostoma albiguttata, FANH 45246, 99.0 2
Uranoscopus sp., USNM 113145, 80.0 2
Zoarcoidei
Anarhichadidae
Anarrhichthys ocellatus, USNM 57832, 585.0 2)
Bathymasteridae
Bathymaster signatus, USNM 24004, 130.0 2
Ronquilus jordani, MPM 394, 133.1 2
Stichaeidae
Anoplarchus purpurescens, MPM 366, 94.2 2
Zoarcidae
Lycodopsis pacifica, MPM 408, 117.3 2
PLEURONECTIFORMES
Achiridae
Achirus lineatus, MPM 13783, 95.0 0
Bothidae
Bothus lunatus, MPM 24885, 114.0 0
Cynoglossidae
Symphurus plagiusa, MPM 10525, 113.0 0
Paralichthyidae
Citharichthys spilopterus, MPM 8951, 103.0 0
Pleuronectinae
Pseudopleuronectes americanus, AMNH 33401, 119.5 0
Psettodidae
Psettodes erumei, BMNH 1904.5.25: 197-8, 83.4 0
Poecilopsettinae
Poecilopsetta hawaiiensis, MPM 13604, 106.3 0
Samarinae
Samariscus triocellata, MPM 13387, 67.0 0
TETRAODONTIFORMES
Balistidae
Rhinecanthus aculeatus, AMNH 50748, 52.5 2,
Monacanthidae
Pervagor spilosoma, MPM 13528, 78.4 2,
DISCUSSION

The insertion of epaxial muscle on to dorsal-fin pterygio-
phores is more widespread and exhibits more variation than
has been previously reported. The distribution of the various
recognized morphotypes suggests that it could have some
value for estimating phylogenetic relationships. The most
commonly encountered morphology among acanthomorphs,
that of no epaxial insertions to dorsal posterolateral processes
of dorsal-fin pterygiophores (Type 0), appears to be the
primitive condition, as it occurs in all basal acanthomorph
taxa (sensu Johnson & Patterson, 1993). Dorsal epaxial/
pterygiophore associations are absent from groups such as
lampridiforms, polymixiiforms, basal paracanthopterygians,
beryciforms, and smegmamorphs, as well as pleuronectiforms
(Table 1). Hence, Types 1-3 are apomorphic at some level.

EPAXIAL MUSCLES AND ACANTHOMORPH RELATIONSHIPS 127

SP

SCA
SN Vii Gp Ege key E
y Gh iA A A ZH YG AL
EPAX Yi MIRE ——<Zp :
EE___ =
'"4ZZ744 ED ‘D
CLA”

Fig. 1 Type 0 epaxial muscle as exemplified by Morone chrysops (MPM 4569, 78.8 mm SL). Inclinatores dorsales removed to expose medial
muscles. Note that there is no insertion of the epaxial musculature to the distal tips of the pterygiophores. Margins of muscle demarcated by
thicker lines; bone stippled. DD, depressores dorsales; DI, distal radial; ED, erectores dorsales; EPAX, epaxialis; PM, proximal-middle
radial; SCA, supracarinalis anterior; SCP, supracarinalis posterior; SN, supraneural; SP, spines; SR, segmented ray. Scale bar = 10 mm.

SP PM
oe DI EPAX ED

W/
ap (et eAI FLING Ai,
fa a SS aa Vy,
EB ——

SS
—_ Ss
4 SS

Fig. 2 Type 1 epaxial muscle as exemplified by Epinephelus merra (USNM 246689, 96.5 mm SL). /nclinatores dorsales removed to expose
medial muscles. Note the separate slip of epaxial muscle which inserts dorsally on to the second pterygiophore (directly behind the second
spine) and the additional insertions on to pterygiophores 3-8. Abbreviations and other methods of presentation as in Fig. 1. Scale bar = 5
mm.

Among these apomorphic morphologies, Type 1 is the
easiest to characterize and identify. It is found among a
restricted group of perciform families and is considered the
exclusive epaxial/pterygiophore association of the Scorpaeni-
formes (see below for discussion of Type 0 condition in
Bathylutichthys). A scorpaeniform sister group has remained
elusive and this has been a serious barrier to understanding
internal relationships of the Scorpaeniformes. The presence
of a derived Type 1 epaxial morphology in the Scorpaeni-
formes and a small subset of the Perciformes suggests that the
sister group of the Scorpaeniformes possibly lies within this
subset. Percoid taxa rarely have been considered candidates
for such status, although seven percoid families exhibit a
Type 1 morphology (Table 1; Figs 2, 4-5). Despite generally
being recognized as a heterogeneous and probably non-
monophyletic assemblage (e.g. Johnson, 1984), percoids
have been referred to as a single, identifiable taxonomic

SCA PM DI
SP1 pm SR1 dl: Wit IaeAaen~ hae —

SCA ZA. ED é ——
Leg Oe —_—$_———SSsSj=
Shi Oa be == —— SSS
——— SSSSn»7 ED ‘DD EPAX
SS Fig. 4 Type 1 epaxial musculature in three percoids: a, Apogon
EPAX : maculatus (MPM 24869, 64.6 mm SL), Apogonidae, with
insertions to the first through third pterygiophores; b,
Centrogenys vaigiensis (USNM 150792, 53.4 mm SL),
Fig. 3 Type 1 epaxial musculature in the batrachoidid Opsanus Centrogeniidae, with insertions to the first through seventh
beta (MPM 8919, 139.5 mm SL). Insertions to the 11th dorsal-fin pterygiophores; c, Perca flavescens (MPM 25409, 79.2 mm SL),
pterygiophore. SP1, first spine; SR1, first segmented ray; other Percidae, with insertions to the fourth through ninth
abbreviations and methods of presentation as in Fig. 1. Scale bar pterygiophores. Abbreviations and other methods of presentation

= 10 mm. as in Figs 1, 3. Scale bars = 5 mm.

128

R.D. MOOI AND A.C. GILL

EPAX PM+DI spy

Fig. 5 Type 1 epaxial morphology with extreme fibre separation from the main epaxial body of the epaxial muscle slip inserting on to
pterygiophores in Haemulon aurolineatum (MPM 23228, 64.2 mm SL). SP1, first dorsal-fin spine; SR1, first segmented dorsal-fin ray; other
abbreviation and methods of presentation as in Figs 1, 3. Scale bar = 5 mm.

group for so long that they have been reified; in practice,
most systematists regard the Percoidei as a bona fide taxon.
As a consequence ichthyologists have rarely examined taxa
from among the Percoidei as potential relatives of non-
percoid taxa (exceptions include Johnson, 1984, 1986, 1993;
Tyler et al. 1989), and few characters have been identified to
suggest a relationship among percoids and scorpaeniforms, at
least in part because few researchers have looked. These
same problems apply to the more inclusive Perciformes, for
which no satisfactory definition exists and membership is
often questionable; families considered perciforms are rarely
examined as either sister taxa or possible members of other
acanthomorph orders (although see Johnson & Patterson,
1993) because, in practice, the Perciformes is treated as a
monophyletic taxon.

Several additional characters suggest that a relationship
between scorpaeniforms and at least some of the ‘percoids’
with a Type 1 epaxial morphology is worthy of consideration.
For example, some larval serranids (particularly anthiines)
bear at least a superficial resemblance to larval scorpaeni-
forms, with suspensorial and cranial bones highly orna-
mented by spines and ridges (cf. Figs and descriptions in:
Baldwin, 1990; Johnson, 1984; Kendall, 1984; Washington et
al., 1984). Moreover, the general physiognomies of many
adult serranids bear striking resemblances to certain scor-
paeniforms. Although general similarities do not provide the
necessary evidence for relationship, they hint that there
might be more evidence than shared epaxial morphology; we
feel it is premature to dismiss these similarities as being due
to convergence before relationships are better understood.

The occurrence of Type 1 epaxial morphology in few
non-percoid perciform taxa (blennioids, some cirrhitoids and
some trachinoids) suggests that these should also be included
in a search for a scorpaeniform sister group, or considered for
inclusion among scorpaeniforms (Mooi & Johnson, in prep).
For example, blennioids also resemble scorpaeniforms in
having the supratemporal sensory canal enclosed by the
parietal (except in most tripterygiids where the cephalic
sensory canals are incompletely enclosed by bone; Springer,
1993:487 and pers. obs.). This condition is found in several
other perciform taxa, including at least some zoarcoids (sensu
Anderson, 1984; Travers, 1984b; all ‘zoarceoids’ according to
Gosline, 1968:46), some pseudochromids (Gill, in prep.), and
mastacembeloid synbranchiforms (Travers, 1984a), but these
taxa do not have a Type 1 epaxial morphology. Champsodon-
tids more closely resemble scorpaeniforms in having a serrate
ridge overlying the canal (Johnson, 1993:14; Mooi &
Johnson, in prep.), as well as Type 1 epaxials. Although
blennioid parietals lack the serrate ridge or spine over the
canal, the possibility of a blennioid/scorpaeniform relation-

ship deserves further study. Certain cottoids closely resemble
blennioids in dorsal gill arch morphology, notably in lacking
an interarcual cartilage, and in having only a single
infrapharyngobranchial (infrapharyngobranchial 3), which
articulates posteriorly with epibranchials 3 and 4 (e.g., com-
pare cottoids in Rosen & Patterson, 1990: figs 34A, C and
Yabe, 1985: figs 23, 24E with blennioids in Rosen & Patter-
son, 1990: figs 33A—B, 37, 38C—D and Springer, 1993: fig. 1).
Members of the cottoid family Liparididae further resemble
blennioids in lacking an uncinate process on epibranchial 1
(Kido, 1988: figs 12A—D).

Johnson & Patterson (1993: 591) found no evidence to
indicate a ‘pre-perciform’ position for scorpaeniforms, and
considered ranking them at the subordinal level within the
perciforms, ‘if only to stimulate the search for characters
justifying their individuality.’ We concur with Johnson &
Patterson’s proposal and award subordinal ranking for the
Scorpaeniformes, as the Scorpaenoidei, within the Perci-
formes. In addition to the justification provided by Johnson
& Patterson (1993), we believe this action will be a major step
forward in diagnosing a monophyletic Perciformes. There is
no contrary evidence for maintaining the two orders as
separate, and the epaxial morphology and other evidence
noted above suggests that the Perciformes is non-
monophyletic without the inclusion of the Scorpaeniformes.

The almost universal occurrence of Type 1 epaxial muscles
in the Scorpaenoidei has implications for its composition. It
casts doubt on the inclusion of the Dactylopteridae and
Bathylutichthyidae within the suborder, as neither family has
insertions of epaxial muscle to dorsal-fin pterygiophores
(Table 1). Johnson (1993: 7) also raised doubts about a
relationship between dactylopterids and scorpaenoids based
on the absence of a bone-enclosed supratemporal canal and

SCA SP DI ,PM

Fetal eh Ale
ra SSS

DD EPAX

Fig. 6 Type 1 epaxial musculature in a ‘primitive’ scorpaeniform
Anoplopoma fimbriata (USNM 208296, 122.2 mm SL). Note the
separate slip of epaxial muscle to the third dorsal-fin
pterygiophore, and other insertions of epaxial to as far posteriorly
as the ninth pterygiophore. Abbreviations and other methods of
presentation as in Fig. 1. Scale bar = 5 mm.

EPAXIAL MUSCLES AND ACANTHOMORPH RELATIONSHIPS

lack of parietal spines; Johnson & Patterson (1993: 579)
considered and rejected a relationship between dactylop-
terids and gasterosteiforms. The monotypic family Bathylu-
tichthyidae was recently erected by Balushkin &
Voskoboynikova (1990) and placed in the Scorpaeniformes
(our Scorpaenoidei) largely on the basis of trend characters
variably shared with some cottoid taxa. Although Bathylu-
tichthys could have secondarily lost Type 1 epaxial insertions,
its position in the Scorpaenoidei should be regarded as
provisional. The condition of the parietal and supratemporal
canal in Bathylutichthys could be informative, but requires
investigation.

Conversely, Mandrytsa (1991) has recently questioned the
inclusion of the Pataecidae in the Scorpaenoidei (his Scor-
paeniformes) based on a study of cephalic lateral-line struc-
ture. We have examined specimens of two of the three
pataecid genera (Aetapcus and Pataecus; Table 1) and found
that they have a typical scorpaenoid Type 1 arrangement of
their epaxial musculature, corroborating their current posi-
tion in the suborder. Ishida’s (1994) more detailed analysis of
various myological and osteological characters also conclu-
sively nests pataecids within the Scorpaenoidei (as the sister
group of the Aploactidae).

Winterbottom (1993) suggested a relationship of gobioids
with the scorpaenoid family Hoplichthyidae, but this is not
supported by our observations. Gobioids have no association
of epaxial muscle with distal portions of the dorsal-fin ptery-
giophores, whereas hoplichthyids exhibit a typical scor-
paenoid Type 1 pattern.

The shared Type 1 morphology in a subset of perciforms
(blennioids, some cirrhitoids, Apogonidae, some Centropo-
midae, Centrogeniidae, Champsodontidae, Cheimarrhich-
thyidae, Grammatidae, Haemulidae, Percidae, and
Serranidae) implies that closer relationships might exist
among these taxa than are presently recognized (cf. Figs 2,
4-5, 12-17). The enigmatic family Centrogeniidae is an
interesting example because its nomenclatural history reflects
the possible relationships suggested by epaxial morphology.
Centrogenys vaigiensis, the single included species, and/or its
junior synonyms, has variously been classified as a scorpaeni-
form (e.g., Day, 1875; Fowler & Bean, 1922), a serranid
(e.g., Jordan, 1923; Weber & de Beaufort, 1931; Paxton et
al., 1989), or has been suggested to bear a superficial
resemblance to cirrhitids (Gosline, 1966; Nelson, 1984).
Although Centrogenys does not fit comfortably into any of
these taxa as they are currently diagnosed, the similar Type 1
epaxial musculature suggests that a detailed anatomical com-
parison could provide considerable insight into their interre-
lationships.

In the Centropomidae, we found that extant members of

g 53
1k \ Zs

129

the subfamily Latinae (Lates, Psammoperca) have a modified
Type 1 epaxial morphology where the muscle insertions to
the pterygiophores are separate from the main epaxial body,
but are below the spine/pterygiophore articulation (Fig. 12);
this arrangement could also be described as a modified Type
0 morphology with a more dorsal position of the normally
proximal insertions. The Centropominae (Centropomus) dif-
fer in lacking such dorsal epaxial insertions to dorsal-fin
pterygiophores (Type 0) (Table 1). Greenwood (1976)
hypothesized the monophyly of the Centropomidae, with its
two subfamilies as sister taxa, on the basis of two synapomor-
phies: pored lateral-line scales extending to posterior margin
of caudal fin, and neural spine of second vertebra markedly
expanded in an ‘anteroposterior direction.’ Pored lateral-line
scales extend well on to the caudal fin in many acanthomorph
fishes, and reach, or nearly reach, the posterior margin of the
fin in several families, including sciaenids (Greenwood,
1976), moronids (G.D. Johnson, pers. comm.), most pem-
pheridids, rhyacichthyids (Springer, 1983) and polynemids.
Therefore, this character does not provide convincing evi-
dence of relationship, and may be plesiomorphic within
perciforms. We also are not convinced that Greenwood’s
second character (also noted by Gosline, 1966), expansion of
the second neural arch, is homologous in centropomines and
latines. In adult centropomines (see Fraser, 1968: 455 for
discussion of ontogenetic variation), the second neural spine
is broadly expanded over most of its length (resulting in a
truncated or rounded distal tip to the spine) and closely
applied to the first neural spine, which is narrow and sharply
pointed (see Fraser, 1968: fig. 14; Greenwood, 1976: fig. 25d;
Rosen, 1985: fig. 39B). In contrast, the anterior neural spine
morphology of the latines does not differ markedly from the
conditions found in various basal perciforms; the second
neural spine is only expanded proximally, and is not closely
applied to the first neural spine (see Greenwood, 1976: figs
25a-c). Given the lack of convincing synapomorphies to unite
the subfamilies Latinae and Centropominae, and considering
the differences in epaxial morphology (as well as various
other anatomical differences listed by Greenwood, 1976),
there is no justification for placing them in a single family.
Based on their modified Type 1 epaxial morphology, we here
remove the African/Indo-Australian genera Lates and Psam-
moperca from the Centropomidae to a separate family,
Latidae. Hypopterus (Western Australia) and Eolates (Italy
[Monte Bolca]), included as latines by Greenwood (1976),
presumably also belong to the newly created Latidae. Green-
wood (1976) considered Psammoperca macroptera, the type
species of Hypopterus, to be a synonym of P. waigiensis, the
single species he recognized in Psammoperca; however,
recent authors (e.g., Allen & Swainston, 1988: 62; Paxton et

Fig. 7 Type 1 epaxial musculature in the scorpaeniform Pterois radiata (USNM 140493, 63.3 mm SL). Note the insertion of the epaxial
muscle on to elements of the second pterygiophore and those posterior to the ninth pterygiophore. Abreviations and other methods of

presentation as in Fig. 1. Scale bar = 5 mm.

130

Pa a | = ee

PM ,DI

oe LOE aa

R.D. MOOI AND A.C. GILL

Oss ates ————

—_ZZ ld —S

cee

RPT ED SRI
wee SS SS ss BEEZ ————
———— ———— eee <

——S——_ =

Fig. 8 An unusual Type 1 epaxial morphology in Normanichthys crockeri (USNM 176507, 63.4 mm SL). I - portion of the epaxial that
inserts on to the anterior pterygiophores largely separate from the main body of the epaxial, with only a few fibres shared from each
myoseptal section. The exceptions are the insertions on the two anteriormost pterygiophores which have many of their fibres originating
from the main epaxial muscle body. II — portion inserting on to pterygiophores that is not separate from the main epaxial body. III —
portion inserting on to the ptergygiophores bearing segmented rays, is mostly separate until just beyond the last ray where it merges with
the rest of the epaxial musculature. RPT, rayless pterygiophore; other abbreviations and methods of presentation as in Figs 1, 3. Scale bar

= 5mm.

al., 1989: 482) have regarded Hypopterus as a valid, mono-
typic genus. We provisionally retain the Centropomidae
(Centropomus only) until its relationships are better under-
stood.

The Trachinoidei as defined by Pietsch & Zabetian (1990)
exhibit a variety of epaxial morphologies (Table 1).
Ammodytids and chiasmodontids have Type 0, champsodon-
tids and cheimarrichthyids have Type 1, and Type 2 is found
in the creediids, percophidids, pinguipedids and ura-
noscopids. Considering the discussion by Johnson (1993:
13-15), this epaxial character distribution casts further doubt
on the integrity of this suborder as currently constituted.
Although it seems likely that the epaxial morphologies as
defined here have evolved more than once among acantho-
morphs, it is difficult to reconcile their distribution with the
phylogeny provided by Pietsch & Zabetian (1990). One of
their phylogenetic hypotheses is a sister group relationship
between the Chiasmodontidae and the Champsodontidae.
The Chiasmodontidae do not exhibit any muscle insertions on
the dorsal-fin pterygiophores, whereas the Champsodontidae
have a Type 1 condition very similar to that of scorpaenoids
and serranids. Ammodytids, considered a derived trachinoid
group, exhibit the primitive Type 0 condition, while a puta-
tive basal taxon, Cheimarrichthys, has Type 1, usually a
derived morphology. Reversals are possible and structural
homologies are uncertain (as discussed below), but the incon-
sistencies among these taxa suggest a more thorough investi-
gation of the composition of the Trachinoidei sensu Pietsch &
Zabetian (1990) is warranted.

There are differences even among those trachinoids that
share a Type 2 morphology. Parapercis has a separate muscle
that runs the entire length of the dorsal fin, with only
intermittent epaxial fibres contributing to the muscle body.
The posterior end of this separate muscle has some fibre and
fascia connection with the supracarinalis posterior and only
very weak attachments to the dorsal-fin pterygiophores that
bear segmented rays. These pterygiophore insertions become
strong anteriorly on spine-bearing pterygiophores, and the
muscle is continuous with the supracarinalis anterior. This
morphology is reminiscent of that of the Mullidae, described
above, but shows an even closer association with the supra-
carinalis muscles, suggesting a supracarinalis derivation,
rather than an epaxial one, for these pterygiophore inser-
tions. This is completely different from the condition in

percophidids (Bembrops), which have a more typical Type 2
morphology with epaxial insertions on to the five pterygio-
phores of the anterior dorsal fin and to the first pterygiophore
of the second, and with the anterior and posterior supracari-
nalis muscles entirely separate from the epaxial musculature.
Of course, such differences can be interpreted as autapomor-
phies for families and genera among the trachinoids, but can
also be considered suggestive of non-relationship.

Epaxial/pterygiophore associations can also strengthen
hypotheses about monophyly of currently recognized groups.
Although not unique among perciforms, the occurrence of
the Type 1 attachment in Niphon spinosus (Fig. 13) and its
proposed relatives, the serranids, lends support to Johnson’s
(1983) placement of Niphon within this family based on other
characters. Niphon had previously been aligned with the
Percichthyidae, a family that exhibits Type 0 epaxial mor-
phology.

Among blennioids (sensu Springer, 1993), the Type 1
epaxial morphology has been found in all examined taxa, but
there is some variation in details. Tripterygiids, dacty-
loscopids, clinids, chaenopsids and blenniids have a separate,
more-or-less fan-shaped, anterior slip of the epaxial muscle
bundle that inserts on to the distal portions of the anterior
dorsal-fin pterygiophores and extends forward to the skull
(Fig. 14a-c). We have not found this anterior slip elsewhere
among acanthomorphs with epaxial attachments to dorsal-fin
pterygiophores, and interpret it as a synapomorphy of the
Blennioidei. This corroborates Springer’s (1993) hypoth-
esized monophyly of the suborder. However, labrisomids are
an exception among blennioids in exhibiting a more typical
Type 1 morphology, without an anterior slip to the skull (Fig.
14d). On the basis of molecular work, Stepien ef al. (1993)
hypothesized that the Labrisomidae are nested within the
Blennioidei. Morphological characters provided by Springer
(1993) also suggest that the Labrisomidae are not a basal
blennioid family; for example, labrisomids, clinids, blenniids,
and chaenopsids are more derived than tripterygiids and
dactyloscopids in having the dorsalmost pectoral-fin ray
articulating only with the dorsalmost proximal radial (vs with
the scapula). Therefore, the absence of an anterior extension
of the dorsal epaxial slip to the skull is most parsimoniously
interpreted as a reversal, and a synapomorphy of the Labriso-
midae.

It is

also possible that the discovery of epaxial/

EPAXIAL MUSCLES AND ACANTHOMORPH RELATIONSHIPS 131

PM
SP1 DI ED

SCA SR1

; eZ LS us fi = ee
DZ iL SOE —————SSSSS——SE SS ee ==577'S

FE —_—— Be

a

YH FALE

f
A

Fig. 9 Type 2 epaxial musculature as exemplified by: a, Opistognathus maxilloxus (MPM 30098, 98.3 mm SL); b, Ronquilus jordani (MPM
394, 133.1 mm SL). Abbreviations and other methods of presentation as in Figs 1, 3. Scale bars = 5 mm.

Fig. 10 Epaxial insertions via long tendons of Sparisoma rubripinne (MPM 30040, 62.6 mm SL), typical of some Type 2 epaxial muscles.
Abbreviations and other methods of presentation as in Figs 1, 3. Scale bar = 5 mm.

SP DI
SN SCM? 4th

Spe PM igi ED SCP
ai ODE LAGE ECE LL BS RSS
7d AN ein AW ZA — = Ki KF Le» By SS Ae EZ

Zl EE ESS SSS CEE

EPAX

LEE

Fig. 11 Type 3 epaxial musculature as exemplified by Parupeneus multifasciatus (MPM 13530, 79.0 mm SL). In contrast to Types 1 and 2,
the dorsal epaxial has direct fibre insertion to only one anterior pterygiophore, and ventral to the articulation with the spine. These anterior
fibres merge with what is possibly a modified supracarinalis medius (SCM?), which has a similar anterior insertion and tendonous insertions
to a few posterior pterygiophores more dorsally. The epaxial muscle shares only a few fibres with the supracarinalis medius near the
posterior end of the first dorsal fin. The supracarinalis medius is contiuous with the supracarinalis posterior. SCM?, possible supracarinalis
medius; other abbreviations and methods of presentation as in Figs 1, 3. Scale bar = 5 mm.

pterygiophore morphologies could help to determine the which suggests they are unlikely to be included among Type 1
relationships of some of the incertae sedis genera of the taxa such as the Serranidae and Haemulidae (where each
Percoidei as identified by Johnson (1984: table 119). For genus, respectively, had been traditionally placed).

example, Siniperca has Type 2 musculature, which, although Many percoid families have not had their close relatives
a relatively common morphology, does circumscribe a identified. Epaxial morphology might limit the search for
smaller perciform group from which possible relationships possible relationships for some of these taxa. For example,
could be initially explored. Johnson (1984) suggested a rela- the Pholidichthyidae exhibit Type 2 morphology, and their
tionship between Symphysanodon and Synagrops based on relationships might be narrowed to other taxa with this
larval morphology. We find the former taxon to have Type 0 morphology. Gill & Mooi (1993) summarized evidence sug-
and the latter to exhibit Type 2 epaxial morphologies. gesting a possible relationship of the Notograptidae to acan-
Although this does not refute a relationship, clearly more thoclinine plesiopids. Notograptids and some acanthoclinines
work needs to be done. Other orphan percoid genera such as share Type 2 morphology, which is absent in other plesiopids
Lateolabrax and Hapalogenys have Type 0 morphology, (Table 1), and this perhaps provides additional support for

R.D. MOOI AND A.C. GILL

132
a
DI PM
PSSSSSENA
I
EPAX
b
Fig. 12

Epaxial muscle morphology in: a, Lates niloticus (ROM 28524, 80.8 mm SL); b, Psammoperca waigiensis (ROM 46627, 91.2 mm

SL). Note the insertions on to the second pterygiophore just ventral to the spine/pterygiophore articulation. Abbreviations and other

methods of presentation as in Fig. 1. Scale bars = 5 mm.

SCA SP PM

d —— SEE
a> SSSS=

EPAX

ee

SSS SSS
{SS SSS SSE
iii ———SSSS

EEE

<=

SSS

Fig. 13 Type 1 epaxial musculature in Niphon spinosus (USNM 59739, 128 mm SL). Note the separate slip of muscle inserting on to the
second dorsal-fin pterygiophore and insertions to the 2nd through 8th pterygiophore, as in Epinephelus (Fig. 2). A separate bundle of fibres
originates tendonously from the 10th pterygiophore to merge with those from the main epaxial muscle body. Abbreviations and other

methods of presentation as in Fig. 1. Scale bar = 10 mm.

their relationship, or at least does not contradict such a
conclusion.

Variation within families exhibiting a particular morpho-
type has considerable potential for exploring internal rela-
tionships. Among serranids, the anthiines Hypoplectrodes,
Acanthistius, and Plectranthias all have very similar epaxial
morphologies (Fig. 15), in which a short and not highly
differentiated slip of muscle inserts on to the second pterygio-
phore, and a weak tendon extends from the myoseptum to
the first pterygiophore. This differs notably from the condi-
tion in more typical anthiines, such as Pseudanthias, where a
completely separate slip of epaxial muscle extends from
below the fifth pterygiophore to insert on to the first through
fourth pterygiophores (Fig. 16). These differences could
provide evidence to unite members of one or another of these
anthiine groups. If epinephelines are the sister group of
anthiines as implied by Johnson (1988) and supported by

Baldwin & Johnson (1993), decisions concerning homology
and character definition become crucial; primitive epi-
nephelines (Niphon, Epinephelus) have a separate slip of
muscle inserting on to the second pterygiophore, but no weak
tendon to the first pterygiophore, a combination of features
found in the two anthiine groups (cf. Figs 2, 13, 15, 16).
Variation in morphology of epaxial musculature might
prove useful in other taxonomic groups. Insertion patterns of
epaxial fibres to pterygiophores, the portions of the pterygio-
phore involved in the insertion, the degree of separation of
the involved musculature from the main body of the epaxial,
and the relationship of the muscle with the supracarinalis all
vary. Among the haemulids examined, Anisotremus has a
limited number of attachments involving only the fourth and
fifth pterygiophores, Conodon exhibits a more robust con-
tinuous series of insertions extending from the third to
seventh pterygiophores more typical of Type 1, and Haemu-

EPAXIAL MUSCLES AND ACANTHOMORPH RELATIONSHIPS

EAs;
<7

LA

_——

Fig. 14 Epaxial musculature of blennioids: a, Tripterygiidae,
Enneanectes pectoralis (MPM 22463, 26.5 mm SL), insertions to
ninth pterygiophore; b, Chaenopsidae, Acanthemblemaria
greenfieldi (MPM 24876, 30.4 mm SL), insertions to 13th
pterygiophore; c, Blenniidae, Entomacrodus nigricans (MPM
18256, 55.4 mm SL), insertions to 11th pterygiophore; d,
Labrisomidae, Labrisomus bucciferus (MPM 31163, 57.0 mm SL),
insertions to 13th pterygiophores. F, fan-shaped anterior slip of
epaxial to skull; other abbreviations and methods of presentation
as in Figs 1, 3. Scale bars = 1 mm (a,b), 5 mm (c,d).

SP
S deutoton

133

lon has an almost completely separate series of muscle fibres
that insert on to the third to ninth pterygiophores (Fig. 5).
Type 1 appears to be the primitive condition for the cirrhi-
toids (Fig. 17), with a secondary change to an epaxial/
pterygiophore association resembling more closely a Type 2
morphology among some cirrhitids, which could be indicative
of close relationship (Table 1). Among sciaenids both epaxial
muscle Types 0 and 2 occur, although their distributions are
difficult to interpret with our current understanding of sci-
aenid relationships (Table 1; Sasaki, 1989). Within scor-
paenoids there is variation in epaxial morphology among the
higher taxa. More extensive surveys within these and other
groups with epaxial/pterygiophore insertions could help to
elucidate some of their intrarelationships.

Basal taxa (Embiotocidae, Pomacentridae, and Cichlidae)
of the suborder Labroidei (Kaufman & Liem, 1982; Stiassny
& Jensen, 1987) exhibits Type 0 morphology, whereas some
labrid taxa exhibit Type 2 (Table 1). It is most parsimonious
to interpret Type 2 epaxial muscle as independently derived
within labrids. This interpretation places Bodianus, Choero-
don, and Tautoga as basal genera among the Labridae, and
might be helpful for determining the polarization of other
characters for phylogeny reconstruction in this confusing
group.

Some tetraodontiforms exhibit epaxial insertions on to the
distal tips of the dorsal-fin pterygiophores that resemble Type
2: Balistidae (Rhinecanthus, pers. obs.; probably Balistes,
Balistapus, Melichthys, and Odonus from figs 78, 86, 88 and
90 in Winterbottom, 1974b), Monacanthidae (Pervagor, pers.
obs.; probably Aluterus, Cantherines, Chaetoderma, Paralu-
teres, Paramonacanthus, and Stephanolepis from figs 100,
102-105 and 108 in Winterbottom, 1974b), probably Tria-
canthidae (Triacanthus, Tripodichthys, Trixiphichthys from
figs 66, 76-77 in Winterbottom, 1974b), and perhaps some
Triacanthodidae (Triacanthodes, Tydemania, and Mac-
rorhamphosodes but not Hollardia or Parahollardia from figs
49, 57-58, 61 and 64 in Winterbottom, 1974b). Consideration
of the overall anterodorsal morphology of balistids, mona-
canthids, and triacanthids suggests that these insertions are
likely to have been derived independently of (and non-
homologous with) those found in the Perciformes. In these
tetraodontiforms, the anterior spinous dorsal fin is closely
associated with the back of the skull and separated from the
soft dorsal fin. It seems that the robust pterygiophores of the
spinous dorsal fin act functionally as a supraoccipital crest
and that the epaxial musculature inserts on to these elements
as it would to such a crest. If triacanthodids, which possess a
more conventional arrangement of spinous dorsal fin and
posterior skull, do have epaxial/dorsal pterygiophore inser-

U cb vdt Ga
Ai ie

Zu SS

EPAX

Fig. 15 Type 1 epaxial musculature in Acanthistius sebastoides (USNM 246689, 96.5 mm SL). A weak tendon extends from a myoseptum to
the first pterygiophore and a short and not highly differentiated muscle slip inserts on to the second pterygiophore. Abbreviations and other

methods of presentation as in Fig. 1. Scale bar = 5 mm.

134

SP1
SCA

SZ, ED

CK
SS
EPAX

Fig. 16 Type 1 epaxial musculature in Pseudanthias taeniatus
(USNM 279782, 44.8 mm SL). A separate slip of the epaxial
inserts on to the first to fourth dorsal-fin pterygiophore, and
epaxial insertions occur as far posteriorly as the eighth
pterygiophore. Abbreviations and other methods of presentation
as in Figs 1, 3. Scale bar = 5 mm.

tions, an argument could be made for homology with a Type
2 morphology found among the perciforms, and implied
relationships should be investigated. Optimizing epaxial char-
acter distribution on existing phylogenies of the tetraodonti-
forms (Winterbottom, 1974b; Leis, 1984) implies that the
Type 2 morphology is the primitive condition for the order.
Unfortunately, the character does not provide additional
evidence for intrarelationships because the remaining extant
families of tetraodontiforms do not possess a spinous dorsal
fin.

Even among taxa that do not exhibit epaxial insertions on
to the distal portions of the proximal-middle pterygiophores
or on to the distal radials, we did observe some possibly
significant variation in other muscle morphology. As noted
above, most (if not all) acanthomorphs have epaxial muscle
insertions on to the proximal ends or along the shafts of the
dorsal-fin pterygiophores. In most pleuronectiforms the
epaxial muscle inserts via bundles of muscle fibres that pass
underneath the depressores dorsales. Psettodes, usually con-
sidered the sister group of other pleuronectiforms, has the
epaxial muscles overlying most of the length of the pterygio-
phores, with very short connections extending under the
depressors to the pterygiophore shafts just ventral to the
spine articulations. These connections only occur on the first
12 pterygiophores. Psettodes is the only genus with dorsal-fin
spines; all other flatfishes have epaxial insertions on to a
higher number of pterygiophores, although most of the
examined taxa have dorsal fins extending over the head. The
extent to which the epaxials overlie the pterygiophores in
remaining flatfishes varies considerably and might be of
interest for determining relationships. The few examined
bothids, paralichthyids and samarines have the epaxials cov-
ering about half the length of the pterygiophores before short
fibres attach to these bones. In available achirids the arrange-
ment is similar to that described for bothids for the most
posterior insertions, but anteriorly there are separate, elon-
gate muscle slips that insert high on to the pterygiophore
shafts just ventral to the ray articulations (Fig. 18). The
cynoglossids, considered close relatives of the achirids (Chap-
leau, 1993), have an epaxial morphology more similar to that
of Psettodes in the one species examined. Poecilopsetta
(Poecilopsettinae) has epaxial muscles that lie only as far
dorsally as the proximal tips of the dorsal-fin pterygiophores,
a condition that appears derived among pleuronectiforms and
could provide evidence for relationship if observed in other
taxa. Additional taxa need to be surveyed and character
definitions must be clarified before epaxial morphology can

R.D. MOOI AND A.C. GILL

contribute to an hypothesis of pleuronectiform phylogeny,
but such an investigation appears worthy of pursuit.

A similar, though less extensive, series of epaxial insertions
under the depressors is found in Urophycis of the Gadidae
(Fig. 19). Gadoids have not been thoroughly surveyed, but
variation in epaxial muscle morphology, which is relatively
simple to observe, might be useful for defining broad groups
among gadoids, and paracanthopterygians in general. The
occurrence of a Type 1 epaxial morphology among batra-
choidids also suggests that a further survey of paracanthop-
terygians could contribute to the understanding of
relationships within this taxon.

Of course, epaxial muscle morphology is not informative in
all cases. For example, the Callionymoidei have a highly
modified Type 2 condition consisting of a complex series of
epaxial insertions on to the pterygiophores and modified
neural spines. This will not help determine whether the
Callionymoidei and Gobiesocidae are sister taxa, as hypoth-
esized by Gosline (1970) and Winterbottom (1993: 409),
because the latter taxon does not have a spine-bearing dorsal
fin. It would be reasonable to suggest that any epaxial muscle
associated with the fin would also have disappeared or have
become reduced. Like any other feature, epaxial morphology
can undergo secondary loss or autapomorphic modification.

The homology of the three epaxial muscle morphotypes
identified remains uncertain. It is unlikely that they form a
nested set of character states. That a single morphotype can
be independently derived from a Type 0 condition is illus-
trated by the independent development of Type 2 in some
labrids, and similarly in the Acanthoclininae, a derived taxon
within the Plesiopidae which otherwise exhibit Type 0 (Table
1). The occurrence of a Type 1 morphology in some paracan-
thopterygians, usually considered unrelated to perciforms,
also indicates non-homology of the character state as recog-
nized here. These examples suggest that the morphologies
themselves require better definition. With more sophisticated
inquiry through ontogenetic or neurological studies, it is
possible that these cases of non-homology can be dismissed as
inappropriately recognized character state equivalence. In
the apparently unique morphology of the Mullidae, Type 3,
the pterygiophore insertions involve both epaxial and supra-
carinalis fibres (Fig. 11). The muscle is essentially separate
from the main epaxial muscle body over its entire length, a
condition very different from that found in the Type 1 or 2
morphologies. It appears that the Type 3 musculature is
directly derived from the supracarinalis muscles, rather than
from the epaxial muscles. This also seems likely in the
pinguipedid trachinoid Parapercis, where the muscle bundle
inserting on to the dorsal-fin pterygiophores is continuous
with the supracarinalis anterior and posterior. The condition
in mullids and Parapercis could provide evidence that, in at
least these taxa, the sheet of muscle inserting on to dorsal-fin
pterygiophores is actually derived from the supracarinalis,
and only secondarily shares muscle fibres from the epaxialis.
These problems of homology and ontogeny of the muscle are
beyond the scope of this paper.

Despite these concerns, we are confident that epaxial
morphology is useful for exploring the relationships of acan-
thomorph taxa. Of course, this one character complex must
be taken in the context of other characters before any
definitive statements can be made regarding, for example,
percoid/scorpaenoid relationships, or before making gener-
alizations concerning the integrity of such groups as the
trachinoids. However, one important concept that the inves-

EPAXIAL MUSCLES AND ACANTHOMORPH RELATIONSHIPS 135

1 VST >
<== SS
ED ‘DD EPAX
a
SP1 PM epax Ee tee
SCA DI

——— <———1

ye Ad hiiedd ab UVELLE fy
if) —$$—$—SSSASSESSS J

SP1 PM

WA Za Z| ED
4 5) RE ae,

Ti thd Le
NEG

EPAX

Fig. 17 Epaxial musculature in cirrhitoids: a, Type 1 in Aplodactylidae, Aplodactylus punctatus (USNM 227298, 58.0 mm SL); b, modified
Type 1 in Cirrhitidae, Paracirrhitus arcatus (MPM 13587, 66.7 mm SL); c, Type 2 in Cirrhitidae, Amblycirrhitus bimacula (MPM 13509,

56.9 mm SL). Abbreviations and other methods of presentation as in Figs 1, 3. Scale bars = 5 mm.

/ Ro \?

A De
L_E i FA
ZZ SE TG7z=z=
ZZ |
EPAX7 ——

4

Fig. 18 Epaxial musculature of the pleuronectiform Achirus lineatus (MPM 13783, 95.0 mm SL). Individual slips of epaxialis insert on to the
dorsal third of the dorsal-fin pterygiophore shafts under the depressores dorsales. Abbreviations and other methods of presentation as in

Fig. 1. Scale bar = 10 mm.

136

SR
Ss ZLLL_
OC EZ,

ZH
VG "Wi EB

Fig. 19 Epaxial musculature in the gadid Urophycis regia (MPM
31175, 133.0 mm SL). Individual slips of muscle extend from the
main epaxialis body to insert on the dorsal-fin pterygiophore
shafts under the depressores dorsales. Abbreviations and other
methods of presentation as in Figs 1, 3. Scale bars = 5 mm.

tigation of epaxial muscle variation elucidates is the need to
shrug off the straitjacket of present classifications when
investigating phylogeny of higher taxa. This is particularly
true when the taxa are already recognized as non-
monophyletic, undefined, or poorly defined (e.g., Percoidei,
Perciformes, Paracanthopterygii), but have in essence been
reified over time. It is necessary to look beyond the tradi-
tional taxonomic boundaries, not only when dealing with
undefined groups such as the percoids, but also when investi-
gating apparently well-defined or well-established taxa such
as the scorpaenoids and trachinoids. Epaxial muscle inser-
tions to dorsal-fin pterygiophores provide one character
complex that illustrates the potential and novel relationships
that such an approach can suggest. These possible relation-
ships await rejection or corroboration from similar studies of
additional characters.

ACKNOWLEDGEMENTS. Specimens were kindly made available by:
Mary Anne Rogers, Kevin Swagel, Mark Westneat (FMNH), Tony
Harold, Marty Rouse, Rick Winterbottom (ROM), Susan Jewett,
Lisa Palmer, Dave Johnson (USNM), Norma Feinberg, Melanie
Stiassny (AMNH), Mark McGrouther, Sally Reader, Tom Trnski
(AMS), Oliver Crimmen, Anne-Marie Hodges (BMNH). Earlier
drafts of this manuscript were reviewed by the late Humphry
Greenwood, Dave Johnson, Jeff Leis, Nigel Merrett, Colin Patter-
son, Darrell Siebert and Vic Springer; their comments were greatly
appreciated. This material is based upon work supported by Smithso-
nian postdoctoral fellowships (RDM, ACG), a _ Lerner-Gray
Research Fellowship at the American Museum of Natural History
(ACG), and the National Science Foundation under Grant No.
DEB-9317695 (RDM).

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— 1984. Percoidei: development and relationships. Jn Moser, H.G., Rich-
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CONTENTS

91 ‘Preliminary studies on a mandibulohyoid ‘ligament’ and other intrabuccal connective tissue
linkages in cirrhitid, latrid and cheilodactylid fishes (Perciformes: Cirrhitoidei)
P.H. Greenwood

103 Anew species of Crocidura (insectivora: Soricidae) recovered from owl pellets in Thailand
P.D. Jenkins and A.L. Smith

111 Redescription of Sudanonautes floweri (De Man, 1901) (Brachyura: Potamoidea: Potamo-
nautidae) from Nigeria and Central Africa
N. Cumberlidge

121 Association of epaxial musculature with dorsal-fin pterygiophores in acanthomorph fishes,
and its phylogenetic significance
R.D. Mooi and A.C. Gill

Bulletin of The Natural History Museum

ZOOLOGY SERIES
Vol. 61, No. 2, November 1995