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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.)

Zoology Series

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

The Natural History Museum

Cromwell Road

London SW7 5BD Issued 30 November 1995

Typeset by Ann Buchan (Typesetters), Middlesex

Printed in Great Britain at The Alden Press, Oxford

| an re ts it URAL

jsub ql ONOVAIWeR ESE UM

SANT ; 12 DEC 1995

Preliminary studies on a mandibulohyoid |, _..,,.-o

‘ligament’ and other intrabuccal connectiv@0°OGY LIBRAR’

tissue linkages in cirrhitid, latrid and

cheilodactylid fishes (Perciformes :

Cirrhitoidei)

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

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 |

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Ligamentous and other connective tissue linkages between the mandible, the palatoquadrate, the hyoid arch |

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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.

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

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.

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).

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

Awt: Tendinous aponeurosis of adductor mandibulae

Geh: Geniohyoideus

Im: Intermandibularis

St: Sternohyoideus

Skeletal elements:

Ang: Anguloarticular

Bb: 1st basibranchial

Ch: Ceratohyal

Dt: Dentary

Ect: Ectopterygoid

Ent: Entopterygoid

Epi: Epihyal

FB: Raised facet for articulation with epihyal

Ga: Gill arch

Hyom Hyomandibula

Hyp: Hypohyals

Thyl: Interhyal

Top: Interoperculum

Max: Maxilla

Mt: Metapterygoid

Pal: Palatine

Pop: Preoperculum

Q: 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

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.

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.

A |

Add. mand. [I u be. ier

Add. mand. I m 4 we

4 - |Z 2a

: eZ

Ga <—~. oS LA

C Lig. prim.

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 5 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 33 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

Pal

Lip

\ Ang

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 (5a 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 Sa 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 Sa has

fused. That none of the other cirrhitoids examined has a

quadrato-interopercular ligament would add credence to 5b

being a true branch of 5a, 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-

P.H. GREENWOOD

A\g

Nae

‘ D

‘

EAS

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; 5a: 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 5a 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

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 ef 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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Teleostei. Procedings of the Academy of Natural Sciences of Philadelphia

125; 225-317.

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-

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

105

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 1 and Figs 2-6).

In detail the dentition of C. attenuata differs in the follow-

ing aspects from that of C. hilliana: I' 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 L,; 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 2 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 6.72 6.96 6.91

SD 0.30 0.21 0.19 0.16 0.15 0.17

n 38 12 it) 5 23 11

Interorbital breadth 3.8-4.6 4.2-4.8 4.2-4.7 4.44.7 4.7-5.3 4.7-5.2

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 4 22 8

Braincase breadth 8.9-10.0 85-95 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 11.35 10.71 10.62 12.09 12.36 12.06

SD 0.55 0.46 0.42 0.24 0.35 0.41

n 72s 13 17 8 26 11

Mandible height 5.2-6.3 4.45.1 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 21 13 17 9 26

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 1.55

n 38 10 iS) 4 22 8

Length of M*: upper toothrow length 3.2-7.0 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 7.50 7.48 7.28 7.34

SD 0.51 0.16 0.28 0.42 0.42 0.35

n 34 12 17 5 24 10

than half of L, 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

‘tecently by Ruedi et al. (1990). Ruedi (in press) has

jattempted to correlate morphological features with these

‘chromosomal forms, in order to assign specific names to

jthem, reserving the name C. fuliginosa for those specimens

jwith 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

jspecimens 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

ito 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

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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Bull. nat. Hist. Mus. Lond. (Zool.) 61(2): 111-119

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 S. 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

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.IV.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 1 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, 1m (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

30.1V.1982, Ogoja, Cross River State, Im, 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.III.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 30.1V.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, 1f, coll. N. Cumberlidge. ZIM K3484,

Benin, 1m, 2f, xii.1909, coll. C. Manger. ZIM K30252, Njaba —

creek, 15.iii.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,

1m, 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, 1f, 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, 11.11.1917, coll. Elbert. ZMB 21308, Douala, If,

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, 1f, x.1912. AMNH

3358, affluents of Nepoko river, near Gamangui (Ituri For-

est), 3m, If. 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, If, iv.1912. AMNH 3448, Faradje, 1f, 1911.

| AMNH 3453, Poko, 1m, 4f, viti.1909. AMNH 3458, north of

Ganza, 1f (ovig), 16.xi1.1909. AMNH 3462, affluents of the

Tshope river, near Stanleyville. AMNH 3465, Yakukuku,

, 1m; 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 BP5048 (1m, CW 54.7, CL

136.0 mm), BP5047 (1f, CW 56.6, CL 39.5 mm) Landana,

\Cote de Loango, 4.ix.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 }), 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 j).

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 metus 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

caudal and superior views (Fig. 2 d,f), not visible from cephalic

view (Fig. 2 e). Subterminal segment of gonopod 1 broadened

conspicuously on outer margin, fringed with bristles (Fig. 2 d,e),

with raised flap extending halfway across segment in distal part,

tapering diagonally to point at junction with terminal segment,

forming roof of chamber for gonopod 2; subterminal segment

beneath flap forming lower floor of chamber for gonopod 2 (Fig.

2 d). Gonopod 2 (Fig. 2 g-i) shorter than gonopod 1 (reaching

only to the junction between last 2 segments of gonopod 1).

REDESCRIPTION OF AFRICAN FRESH-WATER CRAB 115

ig. 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).

erminal segment gonopod 2 cup-shaped, with pointed tip, ADULT FEMALE. Right, left chelipeds same proportions as

xtremely short, only 1/15 as long as subterminal segment. male of same size, unequal in both length, height. Mature

ubterminal segment gonopod 2 widest at base, then tapering female abdomen very wide reaching coxae of pereiopods 2-5.

harply inward, forming long, thin, pointed, upright process Segments of female abdomen becoming gradually longer

supporting short terminal segment. 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

Carapace Dimension (mm)

0 15

Carapace Length (mm)

20 25 30 35 40 45

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

X + SE X + SE x SE

Sudanonautes floweri 1.52+0.01 0.61+0.0 10.38+

0.003

(n = 65)

Sudanonautes aubryi 1.377 0101 (0527 0101 10:38 = 0.002

(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 0.517+0.01 0.417 + 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 0.51*+0.01 0.46% + 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

jalso 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

Mibutary 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,

)

117

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 B5048) 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 §. 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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Systematik, Geographie und Biologie der Thiere 37: 401-410.

— 1929. Potamonidae au Cameroon. Jn: Contribution a l'étude de la faune du

Cameroun. Faune Colonies frangaises 3: 115-129.

— 1936. Beitrage zur Kenntnis der Potamidae (Stisswasserkrabben) des

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Bott, R. 1955. Die Stisswasserkrabben von Afrika (Crust., Decap.) und ihre

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1959. Potamoniden aus West-Afrika. Bulletin de l'Institut Fondamental

D Afrique Noire, Série A 21(3):994-1008.

— 1964. Decapoden aus Angola unter besonderer Berticksichtigung der

Potamoniden (Crust. Decap.) und einem Anhang : Die Typen von Thel-

phusa pelii Herklots 1861. Publicagoes ©ulturais da Companhia de Dia-

mantes de Angola, Lisboa 69: 23-24.

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— 1993a. Redescription of Sudanonautes granulatus (Balss, 1929) (Potam-

oidea, Potamonautidae) from West Africa. Journal of Crustacean Biology

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— 1993b. Further remarks on the identification of Sudanonautes orthostylis

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eroon. Proceedings of the Biolological Society of Washington 106(3):

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Parisi, B. 1925. Un nuovo Potamonidi dell Abissinia. Atti Societi Italia Scienca

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REDESCRIPTION OF AFRICAN FRESH-WATER CRAB

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Bull. nat. Hist. Mus. Lond. (Zool.) 61(2): 121-137 Issued 30 November 1995

Association of epaxial musculature with

dorsal-fin pterygiophores in acanthomorph

fishes, and its phylogenetic significance

RANDALL D. MOOI

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

Latinae is given full familial status.

INTRODUCTION Stiassny, 1990), few workers using myological features have

a 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-

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 : : ; 6

(Stiassny, 1990; Stiassny & Moore, 1992) have shifted the phores, and interpreted this as evidence for uniting the two

focus somewhat from phylogenetic work on individual fami- Ce a 3 Les

lies to broader studies involving interrelationships of subor- Our oe studies on the phylogenetic positions of ye

ders and orders. Such studies are hampered by the difficulties Ces LIES RE UN 2 a aoe ee ee Souain

inherent in examining large numbers of taxa, determining families have failed to provide corroborating evidence for a

| appropriate character complexes, and interpreting homolo- SIS ENC EU Siu Seaeom 1G San EWE ane

gies among the variation within those complexes. In many Opistognathidae. Moreover, a preliminary survey of epaxial

instances, characters are too complex or difficult to survey morphology Mh perciforms | revealed that the reportedly

resulting in an incomplete understanding of their distribution Unique nassocialon of epaxial) musculature _ otGaiSe evi

within the included groups. During the course of investiga- pterygiophores described! By Mok eal((1920)smore widely

tions on the relationships among pseudochromoids (sensu distributed (Gill & Mooi, 1993: 333). Here we present an

Mooi, 1990), we began surveying the relation of dorsal extensive survey of Sau Done up Ue cine Soule.

epaxial myology to the dorsal-fin pterygiophores. Dorsal despite having a wider distribution than indicated by Mok et

epaxial myology appears to exhibit limited but sufficient al., epaxial muscle/dorsal-fin pterygicphore associations nev-

variation over a broad range of taxa and the character states ertheless aopeea ve we kuvely ESIC we ez ay

are simple enough to suggest it to be of high potential for morphs, and exhibit a number of recognizable morphologies.

phylogenetic analysis of higher relationships among acantho- we OcMuG ea BAe PRs spun gous

morphs. distribution of epaxial muscle insertions to dorsal-fin ptery-

Epaxial muscles, the dorsal component of the body muscu- giophores and their homology.

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;

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

Tays (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 Trachinoidei,

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; | — 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. Incertae 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

=a)

Srv

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, FMNH 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

Gobiidae

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 uneat., 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

Soro

Se EPS

NNONNNNNONNOCO

ooo

R.D. MOOI AND A.C. GILL

Taxon, Catalogue No., SL (mm) Type

Percoidei

Acropomatidae

Malakichthys griseus, USNM 184143, 60.2 0

Ambassidae

Ambassis sp., USNM 223376, 37.8 0

Chanda ranga, BMNH 1938. 12.22:132-141, 40.0 0

Apogonidae

Apogon angustatus, USNM 261750, 57.0 1

Apogonichthys ocellatus, AMNH 33808, 43.0 1

Cheilodipterus macrodon, AMNH 33714, 68.0 1

Bathyclupeidae

Bathyclupea malayana,BMNH 1982.9.6:106—107, 117.0 0

Callanthiidae

Callanthias australis, AMS 1.18709-002, 88.0 0

C. platei, USNM 307594, 93.0 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

Centrogentidae

Centrogenys vaigensis, USNM 245612, 70.0 1

Centropomidae

Centropomus armatus, USNM uncat., 108.7 0

C. ensiferus, ROM 61657, 47.7 0

C. pectinatus, ROM 61664, 61.0 0

C. undecimalis, ROM 40904, 118.5 0

Cepolidae

Cepola rubescens, BMNH 1970.4.18:3, 438.0 2

Owstonia totomiensis, BMNH 1986.10.6:61, 91.0 2

Chaetodontidae

Chaetodon multicinctus, MPM 13556, 89.7 0

C. miliaris, MPM 13466, 56.0 0

Datnioides quadrifasciatus, USNM 297256, 120.0 0

Dinolestidae

Dinolestes lewinii, USNM 59932, 138.5 0

Enoplosidae

Enoplosus armatus, USNM 48808, 77.5 0

Gerreidae

Eucinostomus gula, USNM 43216, 45.0 0

Glaucosomatidae

Glaucosoma scapulare, AMS 1.27325-002, 45.1 0

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 1

Conodon nobilis, MPM 13778, 104.5 1

Haemulon aurolineatum, MPM 23228, 64.2 1

Hapalogenys sp., BMNH 1984.1.13:76-82, 55.0 0

Hemilutjanus microphthalmus, USNM 77623, 138.0 0

Kuhlidae

Kuhlia rupestris, USNM 184110, 82.0 0

Kurtidae

Kurtus gulliveri, USNM 217310, 128.0 0

Kyphosidae

Girella tricuspidata, USNM 269547, 99.5 0

Sectator oxyurus, USNM 288880, 75.1 0 |

Lactariidae \

Lactarius delicatulus, BMNH 1895.2.28:51, 87.0 0

Lateolabrax japonicus, USNM 64630, 87.0 0

Latidae

Lates albertianus, ROM 26537, 141.1 1}

L. calcarifer, BMNH 1898.12.24:2, 113.5 1

L. mariae, ROM 28140, 125.2 1

L. niloticus, BMNH 1907.12.2:2959-2968, 48.5 1

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

NNNYNNYNN

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

NNOCNOCONNNY

SSS JS SS Sy SSS Ses SS Sy Ss ea SS Sy yes

i=) NN in)

126

R.D. MOOI AND A.C. GILL

Taxon, Catalogue No., SL (mm) Type Taxon, Catalogue No., SL (mm) Type

C. perplexus, USNM 258839, 51.5 1 Kali normani, USNM 207614, 159.6 0

Icelus hamatus, BMNH 1877.5.13:7-9, 60.3 1 Pseudoscopelus sp., ARC 8706465, 57.0 0

Myoxocephalus scorpis, BMNH 1981.2.10:629, 50.5 1 Creediidae

Taurulus bubalis, BMNH 1981.2.20:776-794, 38.5 1 Crystallodytes cookei, FANH 63619, 41.0 2

Cottocomephoridae Limnichthys fasciatus, AMNH 57282, 45.5 2

Cottocomephorus grewingkii, USNM 222075, 100.0 1 Percophididae

Cyclopteridae Bembrops anatirostris, AMNH 83323, 170.0 2

Cyclopterus lumpus, USNM 197582, 83.8 1 B. gobioides, FANH 67070, 112.0 2

Hoplichthyidae Pinguipedidae

Hoplichthys langsdorfi, USNM 309447, 123.0 1 Parapercis cephalopunctatus, FMNH 72471, 108.0 2

Liparididae P. montillai, AMNH 50585, 94.0 2,

Liparis agassizii, USNM 74697, 117.5 il Uranoscopidae

L. liparis, BMNH 1971.2.16:1757-1760, 99.5 1 Kathetostoma albiguttata, FMNH 45246, 99.0 2

Paraliparis hystrix, BMNH 1992.10.20:43-48, 87.8 1 Uranoscopus sp., USNM 113145, 80.0 2

Normanichthyidae Zoarcoidei

Normanichthys crockeri, USNM 176507, 56.7 1 Anarhichadidae

Pataecidae Anarrhichthys ocellatus, USNM 57832, 585.0 2.

Aetapcus maculatus, BMNH uncat., 118.0 1 Bathymasteridae

Pataecus fronto, BMNH 1914.8.20:282, 159.0 1 Bathymaster signatus, USNM 24004, 130.0 2

Platycephalidae Ronquilus jordani, MPM 394, 133.1 2

Thysanophrys japonica, USNM 70735, 119.0 1 Stichaeidae

Psychrolutidae Anoplarchus purpurescens, MPM 366, 94.2 2

Cottunculus microps, BMNH 1981.3.16:550—-553, 87.5 1 Zoarcidae

Psychrolues zebra, BMNH 1986.7.12:193, 41.5 1 Lycodopsis pacifica, MPM 408, 117.3 2

Scorpaenidae PLEURONECTIFORMES

Pterois radiata, USNM 140491, 64.8 1 Achiridae

Scorpaena sonorae, USNM 59463, 67.6 1 Achirus lineatus, MPM 13783, 95.0 0

Sebastes alutus, USNM 72461, 80.0 1 Bothidae

Triglidae Bothus lunatus, MPM 24885, 114.0 0

Bellator militaris, USNM 114793, 83.0 1 Cynoglossidae

Stromateoidei Symphurus plagiusa, MPM 10525, 113.0 0

Stromateidae Paralichthyidae

Peprilus burti, MPM 8291, 92.4 0 Citharichthys spilopterus, MPM 8951, 103.0 0

Trachinoidei Pleuronectinae

Ammodytidae Pseudopleuronectes americanus, AMNH 33401, 119.5 0

Ammodytes americanus, AMNH 36780, 72.0 0 Psettodidae

A. hexapterus, FMNH 80613, 106.0 0 Psettodes erumei, BMNH 1904.5.25: 197-8, 83.4 0

A. lanceolatus, FMNH 34257, 177.0 0 Poecilopsettinae

A. personatus, USNM 104499, 86.0 0 Poecilopsetta hawaiiensis, MPM 13604, 106.3 0

Champsodontidae Samarinae

Champsodon sp., USNM 150556, 64.2 1 Samariscus triocellata, MPM 13387, 67.0 0

Cheimarrhichthyidae TETRAODONTIFORMES

Cheimarrichthys fosteri, AMNH 98274, 71.0 1 Balistidae

Chiasmodontidae Rhinecanthus aculeatus, AMNH 50748, 52.5 2,

Chiasmodon sp., USNM 186139, 110.0 0 Monacanthidae

Dysalotus alcocki, MCZ 60806, 112.0 0 Pervagor spilosoma, MPM 13528, 78.4 2

ing on to no other pterygiophores, but becoming continuous DISCUSSION

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.

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

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.

EPAX ED

SP PM

rg DI

oat

y U, mee bbe AIA? ZLIL:

———— ——

SSS

GI on

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

SR1

SCA SP1 adh PM ED

SS eS =

Fig. 3 Type 1 epaxial musculature in the batrachoidid Opsanus

beta (MPM 8919, 139.5 mm SL). Insertions to the 11th dorsal-fin

pterygiophore. SP1, first spine; SR1, first segmented ray; other

abbreviations and methods of presentation as in Fig. 1. Scale bar

= 10 mm.

sp PM DI

SCA Lf,

Net Sp Pus oe RL INGRL

Fig. 4 Type 1 epaxial musculature in three percoids: a, Apogon

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),

Centrogeniidae, with insertions to the first through seventh

pterygiophores; c, Perca flavescens (MPM 25409, 79.2 mm SL),

Percidae, with insertions to the fourth through ninth

pterygiophores. Abbreviations and other methods of presentation

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

128

sc ,SN SP1

SCA Pe

SE SE oe Go

POZE

SSS SS

R.D. MOOI AND A.C. GILL

EPAX

hen GOES Foe zi

a/ ———— Se Se

UW, COMMA My <= N pyr

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

SP DI

SSC n et ag, 6A ¥¢ Ab léiah

SLZLGEG PLEA ERLE foe

LLL EE Ee SSsSssSc>}

LL SSSSS—SEaTHfixwxzqPcq»EPE-=

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

SCA

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

LEE eae

= \ nen boshficc EZ =

—SS=

R.D. MOOI AND A.C. GILL

SR1

——SS=—

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 et 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

SR1

=, 2 Sa f LE,

Dike NPA PAS PASI SIA) CAI

SS SS

xt DI ED

SCA SRi

33 pk Bs pE Se

hd, ZU 2 iG Sg

OA WAS = ——————— —— ———— DD

b EPAX ef oom tsi oe

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

= PM

SR1

SCM?

SSS

Lh gk 7

2f Pf 6 LEEE ZEEE AGREE

Zi, IN ia LE,

LEE

EPAX

————

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

relationships of some of the incertae sedis genera of the

Percoidei as identified by Johnson (1984: table 119). For

example, Siniperca has Type 2 musculature, which, although

a relatively common morphology, does circumscribe a

smaller perciform group from which possible relationships

could be initially explored. Johnson (1984) suggested a rela-

tionship between Symphysanodon and Synagrops based on

larval morphology. We find the former taxon to have Type 0

and the latter to exhibit Type 2 epaxial morphologies.

Although this does not refute a relationship, clearly more

work needs to be done. Other orphan percoid genera such as

Lateolabrax and Hapalogenys have Type 0 morphology,

which suggests they are unlikely to be included among Type 1

taxa such as the Serranidae and Haemulidae (where each

genus, respectively, had been traditionally placed).

Many percoid families have not had their close relatives

identified. Epaxial morphology might limit the search for

possibie relationships for some of these taxa. For example,

the Pholidichthyidae exhibit Type 2 morphology, and their

relationships might be narrowed to other taxa with this

morphology. Gill & Mooi (1993) summarized evidence sug-

gesting a possible relationship of the Notograptidae to acan-

thoclinine plesiopids. Notograptids and some acanthoclinines

share Type 2 morphology, which is absent in other plesiopids

(Table 1), and this perhaps provides additional support for

132

EARS

R.D. MOOI AND A.C. GILL

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 a EPAX

L———= SSS SSS SS 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

ace = ————S—SS=S=

sP1 RE. eo

Vie ZZ 2

Y my FI, 0 4

(Aa cniaisl WU pF

CY, hh L—_—=

a hin SS

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).

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-

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

=—=

——————

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

SP1

air

ar Wy oe Be,

Zi iN Mi

SSS

SSSA EGE

ED ‘DD ia

SCA

Zz Lube CLUE FX Hf (7,

Af Wi 2222 a2 bf, DD

——— =

SP1

SCA

LE “fd, Li

VAP OUD te

—— Dp

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.

A CAA ZZ

= oP

EPAX S| |

fa|

= —

ig. 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

SAIL Oe

Ss LIE og

OS LEZ

Lp ED

SSSSS% Ss

EPAX

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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4. Specify any unusual non-keyboard characters on the

front page of the hard copy.

5. Do not right-justify the text.

6. Do not set a left-hand margin.

7. Make sure you distinguish numerals from letters,

e.g. zero (0) from O; one (1) from | (el) and I.

8. Distinguish hyphen, en rule (longer than a hyphen,

used without a space at each end to signify ‘and’ or ‘to’, e.g.

the Harrison—Nelson technique, 91-95%, and increasingly

used with a space at each end parenthetically), and em rule

(longer than an en rule, used with a space at each end

| parenthetically) by: hyphen, two hyphens and three hyphens,

respectively. Be consistent with rule used parenthetically.

9. Use two carriage returns to indicate beginnings of

paragraphs.

10. Be consistent with the presentation of each grade of

heading (see Text below).

Title. The title page should be arranged with the full title;

name(s) of author(s) without academic titles; institutional

aes): suggested running title; address for correspon-

ence.

Synopsis. Each paper should have an abstract not exceeding

| 200 words. This should summarise the main results and conclu-

sions of the study, together with such other information to make

it suitable for publication in abstracting journals without change.

References must not be included in the abstract.

Text. All papers should have an Introduction, Acknowledge-

ments (where applicable) and References; Materials and Meth-

ods should be included unless inappropriate. Other major

headings are left to the author’s discretion and the requirements

of the paper, subject to the Editors’ approval. Three levels of

text headings and sub-headings should be followed. All should

be ranged left and be in upper and lower case. Supra-generic

systematic headings only should be in capitals; generic and

specific names are to be in italics, underlined. Authorities for

species names should be cited only in the first instance. Foot-

notes should be avoided if at all possible.

References. References should be listed alphabetically.

Authorities for species names should not be included under

References, unless clarification is relevant. The author’s

name, in bold and lower case except for the initial letter,

should immediately be followed by the date after a single

space. Where an author is listed more than once, the second

and subsequent entries should be denoted by a long dash.

These entries should be in date order. Joint authorship

papers follow the entries for the first author and an ‘&’ should

be used instead of ‘and’ to connect joint authors. Journal

titles should be entered in full. Examples: (i) Journals:

England, K.W. 1987. Certain Actinaria (Cnidaria, Antho-

zoa) from the Red Sea and tropical Indo-Pacific Ocean.

Bulletin of the British Museum (Natural History), Zoology 53:

206-292. (ii) Books: Jeon, K.W. 1973. The Biology of

Amoeba. 628 p. Academic Press, New York & London. (iii)

Articles from books: Hartman, W.D. 1981. Form and distri-

bution of silica in sponges. pp. 453-493. In: Simpson, T.L. &

Volcani, B.E. (eds) Silicon and Siliceous Structures in Bio-

logical Systems. Springer-Verlag, New York.

Tables. Each table should be typed on a separate sheet

designed to extend across a single or double column width of a

Journal page. It should have a brief specific title, be self-

explanatory and be supplementary to the text. Limited space in

the Journal means that only modest listing of primary data may

be accepted. Lengthy material, such as non-essential locality

lists, tables of measurements or details of mathematical deriva-

tions should be deposited in the Biological Data Collection of

the Department of Library Services, The Natural History

Museum, and reference should be made to them in the text.

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(84 mm wide) or double (174 mm wide) column width of the

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