Zoology Series SZ | THE NATURAL | HISTORY MUSEUM | - VOLUME61 NUMBER2 30 NOVEMBER 1995 — THE NATURAL | HISTORY MNCTUIM 12 ‘Rh PRESENTED ZOOLOGY LISA \RY The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Zoology Series is edited in the Museum’s Department of Zoology Keeper of Zoology: Dr C.R. Curds Editor of Bulletin: Dr N.R. Merrett Assistant Editor: Dr B.T. Clarke Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. 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(Zool.) © The Natural History Museum, 1995 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 | ME COCLUIG TH OM aa tata a ait gs 5 weactoiceesicealanMesicseomidee stowage sr Gseeii< ann do ddtgaanB ted. = Sige Oeomindtnr ceMeoninwel- Sacgmeusech se cewestwaoctinaee rs 91 IM Fier Gilliaval Ini (vd 1CoyotS Migeseacec Ceneee Sepchonc eRe nase SapObtact CE aenaeie: SE Ee ear rneEC eee pneec or cere roscratorteramemercenc, bobcbccrcoane 92 | Ligamentous and other connective tissue linkages between the mandible, the palatoquadrate, the hyoid arch | ANG theEJOPEKCMAar SEMES IN THTES!CITILOIG LAMMNICS Foo ss connie nt cise nces cu cinn orinseseie sina ete tarnetee aioe one Ras sees ee see 93 | Re arnt CMe Ae aria cs aly acetic aie sate ieee Oe ORIN oa «wn atverorematmacaseresaneartaalarateortaefelate Beara acta ovewr torte at tara tol ra 93 URSA T VA AUC ACen otan, MEA TAE ee a. oni A Se. dely dle Dass ss nn oisejan doin ste eiveislesigoaieaiie dele sie,siesikamne ot se omialnaiadisine siduae saeahtels 93 hie tantly! CHeilOdaCwvldsewees. Mea Met cs case rss scaiscoss neste sdscanetes eons ot eeles cassie isda celsmaeissi cleasieie eatansiesies seat 97 DI SEMISSION ANG CONGIISIONS Pe eRe det: Medien sce ecaenacttednensescareetswadeis cies slags ans geetasis jess naieeiiesadajeiicaaee anosise seecseotes 98 ZNOMTION LEC PCIOM SEP rpR eee deere Relates do van. on'o os nt ticle «sisivaien een atom ohn veteeitane «een swine Seelcien wee seeeR Meee cc acd 100 | REL ORENICE Sie eae tee nee ime Stee oan OG MRM wise sues aos enw ence auth seatieuiecetec saccaen Mercer nen Sceteetncese Soesc ents ee eeeRe Meets 101 | Synopsis. In certain taxa of at least three groups of percomorph fishes belonging to the cirrhitoid families Cirrhitidae and Latridae, there is a connective tissue linkage between the mandible and the hyoid arch, suggestive of the mandibulohyoid ligament described in certain sub-percomorph groups. This ligament is generally thought to be a feature of lower teleost fishes, although a mandibulohyoid connection has also been identified in a few more derived taxa. The mandibulohyoid connection in the Cirrhitidae examined would appear to originate from the tendinous aponeurosis associated with the Aw division of the adductor mandibulae muscles, but its derivation in the latrid species Acantholatris monodactylus remains undetermined. The Aw aponeurosis in A. monodactylus, as well as in the latrid Mendosoma, and in two genera of Cheilodactylidae (viz. Chirodactylus and Cheilodactylus) ramifies extensively over the paladoquadrate arch and part of the opercular series. This system, together with various intrabuccal ligaments is described from representatives of the three cirrhitoid families studied. It is concluded that, contrary to several earlier ideas, a mandibulohyoid linkage is of taxonomically and phylogenetically widespread occurrence in teleosts but that it might be derived from different connective tissue sources. The value of this connective tissue complex in phylogenetic studies has yet to be established, but it appears to be of use in at least establishing intragroup relationships within the Cirrhitoidei. 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. © The Natural History Museum, 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). 92 The nature of the hyoid-mandibular connection in the cirrhitoids, a derived percomorph group (Greenwood, 1995), and that described in other teleosts and in the Lepisosteidae, raises doubts about the strict homology of the connection in these various taxa. Functionally, it would seem that a mandibulohyoid connec- tion is of importance both in adults (see Aerts et al., 1987) and in early larval stages (Verraes, 1977); its phylogenetic history in this context is discussed at length by Lauder (1982). Regrettably, the feeding studies on cirrhitoids, from which this anatomical study arose, are not sufficiently advanced or refined to allow informed speculation on any correlation between the morphology and the feeding habits of these fishes. Furthermore, many other cirrhitoid taxa remain to be studied before it will be possible to evaluate what significance the different types of intragroup mandibulohyoid and other linkages (or absence thereof) may have in unravelling the taxonomy and phyletic relationships of the group. Neverthe- less, there are indications from this study, and from informa- tion in the literature on various forms of mandibulohyoid connections, that further investigations may yield insight into the biomechanics of feeding and into historical relationships. MATERIALS AND METHODS Material Clupeidae: Etrumeus terres. RUSI 34140 (1 speci- men) Salmonidae: Oncorhynchus mykiss RUSI 36417 (3 specimens) Characidae: Hydrocynus vittatus RUSI 19355 (1 specimen) Cirrhitidae: Amblycirrhitus bimacula RUSI 77-20 (3 specimens) Cirrhitichthys oxycephalus RUSI 40526 (3 specimens) Cirrhitops fasciatus RUSI 2375 (1 speci- men; ex Hawaii) Cyprinocirrhites polyactis RUSI 12339 (3 specimens) Paracirrhites arcatus RUSI 30975 (2 specimens) Paracirrhites forsteri RUSI 39419 (3 specimens) Latridae: Acantholatris | monodactylus RUSI 33485 (2 specimens) Mendosoma lineatum RUSI 33613 (1 specimen) RUSI 33626 (1 specimen) RUSI 26176 (1 specimen) Cheilodactylidae: | Cheilodactylus fasciatus DIFS unreg. (2 specimens) Cheilodactylus pixi DIFS unreg. (3 specimens) Chirodactylus brachydactylus DIFS unreg. (3 specimens) DIFS: Department of Ichthyology and Fisheries Science, Rhodes University, Grahamstown. RUSI: J.L.B. Smith Insti- tute of Ichthyology, Grahamstown. P.H. GREENWOOD Method The entire opercular series, palatopterygoid arch and hyoid arch of one side, together with the mandible, premaxilla and maxilla of that side, were dissected away from the head. Muscles, tendons and ligaments were examined on the dis- sected side, and checked on the contralateral aspect which was left in situ. All specimens had been fixed in formalin and preserved in either ethyl or propyl alcohol. In the absence of ontogenetical information on the devel- opment of ligamentous and tendinous systems in these fishes, ligaments and tendons in the various taxa are presumed to be homologous if their places of origin and insertion are similar. ABBREVIATIONS FOR FIGURES Abbreviations for tendons and ligaments are given separately with each figure. Muscles: Add. mand. Iu&m: Adductor mandibulae A, upper and main divi- sions respectively Aw: Aw division of adductor mandibulae muscle Awt: Tendinous aponeurosis of adductor mandibulae Aw 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 93 that division of the muscle inserting on the maxilla through both the ligamentum primordium and the maxillary ligament of adductor mandibulae A, muscle. In all three groups the major (ie lower) division of the muscle is attached to the ligamentum primordium and the maxillary ligament, the latter inserting on the ventral aspect of the maxilla, and the former on the bone’s dorsolateral aspect. Other intergroup differences involve the epihyal- interopercular and the interhyal-interopercular ligaments (For comparison of these and other ligaments with the situation in other cirrhitoid families, see pp. 94 and pp. 97-98 and Figs 2-4). Group II species have the latter ligament partly associated with the epihyal as well as the interhyal, as does the single Group III species dissected; in Group I taxa, however, the ligament is confined to the interhyal. The epihyal-interopercular ligament shows more marked inter- group differences, especially when species of Group I are compared with those of the other two groups, a difference possibly associated with the manner in which the epihyal contacts the interoperculum. In Group II taxa, the lateral face of the epihyal head articulates with a well-defined, prominently raised and posteriorly directed facet situated a little below the dorsal margin of the interoperculum and slightly behind the bone’s midpoint. The epihyal- interopercular ligament in these fishes is short and stout, originates on the lateral face of the epihyal near its dorsal tip, and runs forward at approximately 45 to the sagital plane. It inserts on the upper and anterior faces of the prominence supporting the facet on the interoperculum against which the epihyal articulates.A similar epihyal-interopercular ligament occurs in the single Group III species examined, viz. Ambly- cirrhitus bimacula. However, in this species, unlike those of Group II, the interopercular facet is located on a relatively lower base. The epihyal-interopercular ligament is most distinctive in Group II. In species of this group (unlike the other groups) the epihyal articulates directly with the medial face of the interoperculum and not with a facet carried on a distinct and elevated base (albeit only slightly so in the single Group III species examined). The ligament itself is a prominent feature originating (as in other groups) on the dorsal tip of the epihyal’s lateral face, from which it runs anteriorly onto the dorsal margin of the interoperculum at a point near the bone’s anterior tip, where it is narrowly separated from the attachment point of the mandibulo-interopercular ligament (cf. Acantholatris monodactylus Fig.3 & p. 94). A short interhyal-metapterygoid ligament is present in all three groups. No interhyal-preopercular ligament is present in any exam- ined species of the three groups (cf. the other cirrhitoids described below). A stout mandibulo-interopercular ligament (Fig. 1C; lig. 3) is present in taxa of the three groups. It is confined to the lateral face of both elements in all species except Cyprinocir- rhites polyactis and Cirrhitichthys oxycephalus (both members of Group I). In these two species it divides anteriorly to insert on both the lateral and the medial aspects of the anguloarticu- lar bone. Also common to species of the three groups is a short and deep, ventrally located ligament connecting the anguloarticu- lar and dentary. 94 The family Latridae There are clear-cut differences in certain aspects of the ligamentous and other connective systems in the two latrid species examined, namely the monotypic genus Mendosoma lineatum (Gay) and the species Acantholatris monodactylus (Carmichael) of that polytypic genus. Also, an upper division of the adductor mandibulae muscle A, is absent in A. monodactylus whereas in Mendosoma it is an elongate, rather thin element which lies lateral to the major part of the muscle and extends over the greater part of its length. The minor division, unlike the major one, has no direct connection with the ligamentum primordium and inserts on the maxilla, together with the major division, via the maxillary tendon of the A, muscle. In Mendosoma the adductor mandibulae Aw division is a very thin muscle, largely tendinous and with a single posterior extension of its tendinous aponeurosis. This runs slightly below the upper point of the articulation of the lower jaw with the quadrate, to which bone it is attached a short distance from the anterior border (Fig. 2; tendon 3). In other words, it is of the basic percomorph type sensu Gosline (1986), except that in Mendosoma it has one prolongation extending along the symplectic, another running ventrally to attach to the medial aspect of the preoperculum, a third, directed dorsally to insert on the quadrate, and a fourth directed obliquely backwards to attach to the ventral aspect of the interoperculum medially and anteriorly. In all essentials, the adductor mandibulae Aw tendon system’s extension onto the interoperculum and preopercu- lum in Mendosoma is very similar to that in Acantholatris, with that in Mendosoma, as it were, foreshadowing the more clearly differentiated condition in Acantholatris (cf. Figs 2 & 3). There is no ligament-like connection between the mandible and hyoid arch in Mendosoma (cf. Acantholatris; Fig. 3). The epihyal-interopercular ligament is stout and short, connecting the lateral aspect of the epihyal with the dorsal margin of the interoperculum a short distance anterior to its slightly raised facet for articulation with the epihyal. Unlike the backward-facing facet in those cirrhitids in which it occurs, that in M. lineatum faces forward (Fig. 2), as it does in the other latrid examined (Acantholatris monodactylus; and in the cheilodactylids dissected). A discrete interhyal-interopercular ligament (present in cirrhitids) is apparently lacking in M. lineatum (as it is also in Acantholatris, and Cheilodactylus). Like the two latter genera, but not in the cirrhitids exam- ined, Mendosoma has a_ well-developed _ interhyal- preopercular ligament and another, more dorsally placed ligament between the interhyal and the metapterygoid (Fig. P.H. GREENWOOD 2; lig. 7). This latter ligament I consider to be the homologue of the interhyal-quadrate ligament in Cheilodactylus, and the ligament in Acantholatris which runs from the interhyal to both the quadrate and the entopterygoid (see p. 98 & p. 99 respectively). Mendosoma has discrete lateral and medial divisions of the mandibulo-interopercular ligament, with the medial division terminating a short distance behind the anterior tip of the interoperculum (Fig. 2; lig. 6), and the lateral division extending much further posteriorly. The anguloarticular-dentary ligament is short and stout, markedly stouter than in any cirrhitid species examined, and stouter than that in Acantholatris. As compared with the ligamentous and other connective tissue systems in Mendosoma lineatum, those in Acantholatris monodactylus are considerably more complex, (as they are when compared with the cirrhitid species studied). As was noted earlier (p. 94), there is no obvious sub-division of the adductor mandibulae A, muscle in A. monodactylus. How- ever, anteriorly the upper third of the muscle, unlike the other two-thirds, is free from the ligamentum primordium and inserts on the maxilla only through the maxillary tendon, to which the major part of the muscle is also attached. Acantholatris monodactylus has a substantial Aw portion of the adductor mandibulae muscle. From the muscle’s mediolateral tendinous aponeurosis a stout and relatively short branch (tendon 3 in Fig. 3) runs posteriorly to insert on the anteromedial aspect of the preoperculum’s horizontal limb. A second stout and much longer tendon from the Aw muscle (tendon 5 in Fig. 3) extends from the ventral margin of the muscle above the anguloarticular bone, and runs obliquely backwards to attach to the medial aspect of the interoperculum a short distance from that bone’s anterior tip. This tendon, unlike tendon 3, is not derived from the aponeurosis of the adductor mandibulae Aw muscle but originates directly from the muscle itself. Immediately after its origin, tendon 5 is attached to the anterodorsal aspect of the anguloarticular’s medial face. It then passes over that face of the retroarticular, and attaches to the medial aspect of the interoperculum a short distance from the bone’s anterior tip. Since this tendon links the mandible with the interoperculum it would appear to be the functional equivalent of the mandibulo-interopercular ligament in the other species described above. However, a true and very long mandibulo- interopercular ligament is also present in A. monodactylus (Fig. 3; lig. 6). Anteriorly it has an extensive attachment to the lateral face of the anguloarticular and retroarticular bones, as well as another on the posterior face of the — retroarticular. From here the ligament extends across to, and Fig. 1 A: Cyprinocirrhites polyactic (Group I species) Medial aspect of the left lower jaw, cheek region and hyoid arch, viewed obliquely from above, to show the mandibulohyoid connection (semi-schematic). The branchial skeleton is displaced to the right. About times natural size. 1: Mandibulohyoid connection; 2a: tendon from lower part of adductor mandibulae A, muscle to maxilla; 2b: continuation of tendon 2a, joining tendinous aponeurosis of adductor mandibulae muscle Aw. Lig. prim: Ligamentum primordium. B: Cirrhitops fasciatus (Group I species) Diagramatic representation of mandibulohyoid connection and related tendons; medial aspect of left side to demonstrate the second form of tendinous relationships within species of Group I. Abbreviations as in Fig. 1A. C: Paracirrhites forsteri (Group II species). Medial aspect of the right lower jaw, cheek region and hyoid arch, viewed somewhat dorsally; the branchial skeleton and hyoid arch considerably displaced to the left and posteriorly in order to reveal the mandibulohyoid connection. (Semi-schematic). About times natural size. 1: Posterior portion of mandibulohyoid connection, inserting partly on the quadrate, and partly | continuous with tendinous aponeurosis of the adductor mandibulae muscle Aw (Awt); 2b: ventral continuation of maxillary tendon of adductor mandibulae muscle A,; 3: interopercular-mandibular ligament; 4: tendon of adductor mandibulae A, muscle; Lig. prim: ligamentum primordium. MANDIBULOHYOID CONNECTION IN CIRRHITOID FISHES Lig. prim. A | Add. mand. [I u be. ier Add. mand. I m 4 we 4 - |Z 2a : eZ Ga <—~. oS LA C Lig. prim. 95 96 P.H. GREENWOOD Pal Fig. 2. Mendosoma lineatum Medial aspect of left lower jaw, cheek region, and hyoid arch. Scale = 2mm. 2a: Maxillary tendon of adductor mandibulae muscle A,; 2b: extension of tendon 2a, joining tendinous aponeurosis of adductor mandibulae muscle Aw; 2c: tendon of adductor mandibulae muscle A,; 3a & b: extensions of adductor mandibulae muscle Aw’s tendinous aponeurosis; 6: interopercular-mandibular ligament; 7: interhyal-metapterygoid ligament. along, the dorsal and dorsolateral margins of the interopercu- lum, ending at a point about midway between the bone’s anterior tip and the face of the prominent, forward-facing articulatory facet for the epihyal (cf pp. 94). Here it attaches to a slight eminence on the dorsal margin of the interopercu- lum. At first sight the ligament appears to be continuous with the epihyal-interopercular ligament (Fig. 3; lig. 4) which also inserts at that point. Careful dissection reveals, however, that the two are separate entities (see also p. 93 and p. 94 respectively for the situation in cirrhitids and the latrid - Mendosoma). Apart from the more complex condition in cheilodactylids, this double linkage of the mandible with the interoperculum, one involving both tendons and ligaments, seemingly has not been recorded in any other teleosts. However, it also occurs in Mendosoma (see p. 94 and Fig. 2) where the lowermost arm of the Aw aponeurosis is attached to the anteromedial aspect of the interoperculum, and in Cheilodactylus (see below, and tendon 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 97 Pal Lip \ Ang R 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- 98 P.H. GREENWOOD 2a TP A\g Nae WS ‘ D ‘ ‘ \ EAS NS Ee 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 99 adductor mandibulae Aw tendon system seems less likely in the latrid Acantholatris monodactylus. Here the linkage extends from the posterior tip of the dentary’s lower arm (not, as in the cirrhitids, from its coronoid process or the anguloarticular) to the upper part of the ceratohyal’s lateral face (Fig. 3). At no point has this apparent ligament in Acantholatris any association with the adductor Aw muscle or any part of its tendon system. With regard to its attachment to the lower aspect of the dentary, the connection is compa- rable both with the loosely compacted and fibrous linkage between the dentary and ceratohyal identified by Aerts 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. REFERENCES Aerts, P., Osse, J.W.M. & Verraes, W. 1987. Model of jaw depression during feeding in Astatotilapia elegans (Teleostei: Cichlidae). Mechanisms for energy storage and triggering. Journal of Morphology, 194: 85-109. Anker, G.CH. 1974. Morphology and kinetics of the head of the stickleback, Gasterosteus aculeatus. Transactions of the Zoological Society of London, 32: 311-416. — 1978. The morphology of the head-muscles of a generalized Haplochromis species: H. elegans Trewavas 1933 (Pisces, Cichlidae). Netherlands Journal of Zoology, 28(2): 234-271. Edgeworth, F.H. 1935. The cranial muscles of the vertebrates, viit+493 pp. Cambridge University Press, Cambridge. Elshoud-Oldenhave, M.J.W. & Osse, J.W.M. 1976. Functional morphology of the feeding system in the Ruff-Gymnocephalus cernua (L. 1758) — Teleostei, Percidae). Journal of Morphology, 150: 399-422. Gosline, W.A. 1986. Jaw muscle configuration in some higher teleostean fishes. Copeia, 1986: 705-713. Greenwood, P.H. 1985. Notes on the anatomy and phyletic relationships of Hemichromis Peters, 1858. Bulletin of the British Museum (Natural History), (Zoology), 48(3): 131-171. — 1995. A revised familial classification for certain cirrhitoid genera (Teleostei, Percoidei, Cirrhitoidea), with comments on the group’s mono- phyly and taxonomic ranking. Bulletin of the Natural History Museum (Zoology). Lauder, G.V. 1982. Patterns of evolution in the feeding mechanism of actinopterygian fishes. American Zoologist, 22: 275-285. 101 —— & Liem, K.F. 1980. The feeding mechanism and cephalic myology of Salvelinus fontinalis: form, function, and evolutionary significance. pp. 365-390. In: Balon, E.K. (ed.) Charrs: salmonid fishes of the genus Salveli- nus. W. Junk by Publishers, The Hague. — 1989. The role of historical factors in the evolution of complex organismal functions. pp. 63-78. In: Wake, D.B. & Roth, G. (eds) Complex organismal functions: integration and evolution in vertebrates. John Wiley & Sons Ltd. Chichester Liem, K.F. 1978. Modulatory multiplicity in the functional repertoire of the feeding mechanism in cichlid fishes. Part 1: Piscivores. Journal of Morphol- ogy, 158: 323-360. — 1991. Functional morphology. pp.129-150. Jn: Keenleyside, M.H.A. (ed.) Cichlid fishes: behaviour, ecology and evolution. Chapman & Hall, London. Osse, J.W.M. 1969. Functional morphology of the head of the Perch (Perca fluviatilis L.): an electromyographic study. Netherlands Journal of Zoology, 19(3): 289-392. Rosen, D.E. 1973. Interrelationships of higher euteleostean fishes. pp. 397-513. In: Greenwood, P.H., Miles, R.S., and Patterson, C. (eds) /nterrelationships of fishes. Academic Press, London. Springer, V.G., Smith, C.L., and Fraser, T.H. 1977. Anisochromis straussi, new species of protogynous hermaphroditic fish, and synonymy of Anisoch- romidae, Pseudoplesiopidae, and Pseudochromidae. Smithsonian Contribu- tions to Zoology. No. 252: iii + 1-13. Tchernavin, V.V. 1953. The feeding mechanisms of a deep sea fish, Chauliodus sloani Schneider, vii + 101 + 10 plates. British Museum (Natural History), London. Verraes, W. 1977. Postembryonic ontogeny and functional anatomy of the ligamentum mandibulo-hyoideum and the ligamentum interoperculo- mandibulare, with notes on the opercular bones and some other cranial elements in Salmo gairdneri Richardson, 1836 (Teleostei: Salmonidae). Journal of Morphology, 151: 111-120. Wiley, E.O. 1976. The phylogeny and biogeography of fossil and recent Gars (Actinopterygii: Lepisosteidae). Miscellaneous Publications, University of Kansas Museum of Natural History, No. 64: 1-111. Winterbottom, R. 1974. A descriptive synonymy of the striated muscles of the 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- © The Natural History Museum, 1995 plete maxillary and mandibular dentition. Extracted from an owl pellet from a roost at Wat Tham Maho Lan, Ban Nong Hin, 48 km south of Loei, Loei Province, northeastern Thailand, 17°06'N 101°53’E, altitude 575m. PARATYPES. Eighteen specimens of crania with mandibles and thirteen specimens of crania only, all from owl pellets at the same locality as the holotype. Three specimens of crania with mandibles and five specimens of crania only from Wat Tham Pha Phu, 7 km north of Loei, Loei Province, 17°34'N 101°42’E, altitude 542m. Diagnosis Zygomatic process of maxilla broad and angular, interorbital region narrow; coronoid process broad and deep. Upper and lower first incisors robust, first upper unicuspid large and broad relative to the other unicuspids, talonid of the third lower molar (M,) reduced to a single cusp. Description Overlapping in cranial size with medium to large specimens of Crocidura attenuata and smaller specimens of C. fuliginosa but differing from both species in proportions (see Table 1 and Figs 1-6). Cranium and mandible robust; cranium angu- lar in appearance, especially in dorsal view. The rostrum is moderately deep and obliquely sloping anteriorly; the maxil- lary region is broad, the zygomatic process of the maxilla is broad and angular; the interorbital region is long and narrow, its width increasing only slightly from anterior to posterior; the zygomatic plate is positioned above the first upper molar (M!) and the anterior of the second upper molar (M7), its posterior face is semi-circular and deeper than the anterior face; the braincase is angular, with pronounced angular superior articular facets in dorsal view, a squamosal crest is present and lambdoid crests are well developed, meeting at an acute angle at the midline. The horizontal ramus of the mandible is moderately robust; the coronoid process is broad and deep (see Fig. 4); the ascending ramus is long and low; the condyle is nearly as broad or broader than deep (ratio of 104 P.D. JENKINS AND A.L. SMITH Fig. 1 BM(NH)1933.4.1.183. condyle width to height 91.2-113.3), the postero-internal ramal fossa has a broad base and is approximately as broad as deep; the mental foramen is positioned below the anterior part of the first lower molar (M,). The dentition is illustrated in Figs 2-6. The first upper incisor (I') is robust, slightly proodont with well developed posterolingual and posterobuccal cingula; the upper unicus- pids are overlapping and crowded; the first upper unicuspid (Un') is large and broad in comparison with the other unicuspids, its breadth is equal to or greater than the distance between the two first unicuspids and this tooth is more than two thirds the height of I' and P*; the second and third unicuspids (Un? and Un*) are subequal in size; the upper premolar (P*) has a moderately small parastyle and robust metacone; the first and second upper molars (M! and M7?) show no significant distinguishing features; the third upper molar (M®*) is short and slender with a slightly compressed lingual basin. The first lower incisor (I,) is robust, long, deep and curved, and the anterolingual ridge extends for circa three quarters of the length of the tooth, diverging from the ventral border, the posterior border of I, lies below the middle of the lower premolar (P,); two thirds of the second lower incisor (I,) are in contact with I, and one third of the tooth is overlapped by P,; the postentoconid ledge is very narrow in the first lower molar (M,) and yet more reduced in the second lower molar (M,); the talonid of the third lower Dorsal view of cranium from left to right of C. attenuata BM(NH)1911.9.8.26, C. hilliana BM(NH)1994.113 and C. fuliginosa molar (M,) is reduced to a single cusp. Etymology This species is named in honour of John Edwards Hill, who taught one of the authors (PDJ) the basics of mammalogy and who also provided invaluable help in the identification of some of the skull fragments of bats found during the survey. Comparison with other species Five species of Crocidura have been recorded from Thailand (Lekagul & McNeely, 1977, Davison, 1984): C. fuliginosa (including C. dracula Thomas, 1912 listed as a separate species by Lekagul and McNeely), C. attenuata, C. pullata vorax Allen, 1923 (listed as C. russula vorax), C. horsfieldii indochinensis Robinson & Kloss, 1922 and C. monticola Peters, 1870. Crocidura hilliana is separated from most specimens of C. fuliginosa dracula by its smaller size (see Table 1), while it is considerably larger than C. p. vorax (condylobasal length <17.5), C. horsfieldii (condyloincisive length <17.9, data taken from Heaney & Timm (1983) for specimens from Vietnam) or C. monticola (condylobasal length <17.4). Crocidura hilliana falls at the middle to upper part of the | cranial size range of C. attenuata and the lower part of the size NEW SPECIES OF CROCIDURA 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. 7 | 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. References Allen, G.M. 1923. New Chinese insectivores. American Museum Novitates (100): 1-11. Allen, G.M. & Coolidge, H.J. 1940. Mammal and bird collections of the Asiatic Primate Expedition: mammals. Bulletin of the Museum of Comparative Zoology 87 (3): 131-166. Andrews, P. 1990. Owls, caves and fossils. Natural History Museum Publica- tions, London. Blyth, E. 1855. Proceedings of the Asiatic Society of Bengal for May 1855. Journal of the Asiatic Society of Bengal 24 (4): 354-364. Bonhote, J.L. 1903. Report on the mammals. Fasciculi Malayenses, Zoology 1: 1-44, pl. 1-4. Bunn, D.S., Warburton, A.B. & Wilson, R.D.S. 1982. The barn owl. T. & A.D. Poyser, Calton. 109 Butler, P.M. & Greenwood, M. 1979. Soricidae (Mammalia) from the carly Pleistocene of Olduvai Gorge, Tanzania. Zoological Journal of the Linnean Society 67: 329-379. Corbet, G.B. & Hill, J.E. 1992. The mammals of the Indomalayan region. Natural History Museum Publications and Oxford University Press, Oxford. Davison, G.W.H. 1984. New records of peninsular Malayan and Thai shrews. Malayan Nature Journal 36 (4): 211-215. Geoffroy Saint-Hilaire, E. 1810. Sur les phyllostomes et les mégadermes. Annales du Muséum d'Histoire Naturelle. 15: 157-198. Glue, D.E. 1967. Prey taken by the barn owl in England and Wales. Bird Study 14 (3): 169-183. Heaney, L.R. & Timm, R.M. 1983. Systematics and distribution of shrews of the genus Crocidura (Mammalia: Insectivora) in Vietnam. Proceedings of the Biological Society of Washington 96 (1): 115-120. Hill, J.E. 1960. The Robinson collection of Malaysian mammals. Bulletin of the Raffles Museum, Singapore (29): 1-112. Hutterer, R., 1993. Order Insectivora.: 69-130. Jn D. E. Wilson & D. M. Reeder (Editors) Mammal species of the world. Smithsonian Institution Press, Washington and London. Jenkins, P.D. 1976. Variation in Eurasian shrews of the genus Crocidura (Insectivora: Soricidae). Bulletin of the British Museum (Natural History), Zoology 30 (7): 271-309. 1982. A discussion of Malayan and Indonesian shrews of the genus Crocidura (Insectivora: Soricidae). Zoologische Mededelingen 56 (21): 267-279. Kloss, C.B. 1917. On five new mammals from Siam. Journal of the Natural History Society of Siam 2: 282-287. — 1919. On mammals collected in Siam. Journal of the Natural History Society of Siam 3 (4): 333-407. Lekagul, B. & McNeely, J.A. 1977. Mammals of Thailand. Kurusapha Ladprao Press, Bangkok. Medway, Lord 1965. Mammals of Borneo. Field keys and an annotated checklist. Journal of the Malayan Branch of the Royal Asiatic Society 36 (3): 1-193. — 1977. Mammals of Borneo. Field keys and an annotated checklist. Monographs of the Malayan Branch of the Royal Asiatic Society (7): 1-172. Meester, J. 1963. A systematic revision of the shrew genus Crocidura in Southern Africa. Transvaal Museum Memoir (13): 1-127. Milne-Edwards, A. 1872. Recherches pour servir a histoire naturelle des Mammiferes. Paris. Mills, J.R.E. 1966. The functional occlusion of the teeth of Insectivora. Journal of the Linnean Society (Zoology) 46 (308): 1-25. Peters, W. 1870. Uber neue Arten von Spitzmausen des Konig]. zoologischen Museums aus Ceylon, Malacca, Borneo, China, Luzon und Ostafrika. Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin. : 584-596. Robinson, H.C. & Kloss, C.B. 1914. On new mammals mainly from Bandon and the adjacent islands, east coast of the Malay Peninsula. Annals and Magazine of Natural History (8) 13: 223-234. & 1922. New mammals from French Indo-China and Siam. Annals and Magazine of Natural History (9) 9: 87-99. & 1923. Mammals and birds from the hills of Nakon Sri Tamarat, peninsular Siam. Journal of the Federal Malay States Museums. 11: 58-63. Robinson, M.F. & Smith, A.L. (in prep.) Small mammals from Northeast Thailand. Ruedi, M., Maddalena, T., Yong, H.- S. & Vogel, P. 1990. The Crocidura fuliginosa species complex (Mammalia: Insectivora) in peninsular Malaysia: biological, karyological and genetical evidence. Biochemical Systematics and Ecology. 18 (7/8): 573-581. (in press) Taxonomic revision of Crocidura shrews from the Sunda Shelf and Sulawesi with description of two new species (Mammalia: Soricidae). Zoological Journal of the Linnean Society. Savi, P. 1822. Osservazioni sopra il mustietto o mustiolo, nuova specie di topo ragno Toscano Sorex etruscus. Nuovo Giornale dei Letterati, Pisa. 1: 60-71. Scopoli, J.A. 1769. Annus I [— 5] Historico Naturalis. Lipsiae. Smith, A.L. & Robinson, M.F. (in prep.) Observations on the diet of the barn owl, Tyto alba, in Northeast Thailand. Swindler, D.R. 1976. Dentition of living primates. Academic Press, London. Thomas, [M.R.] O. 1912. New species of Crocidura and Petaurista from Yunnan. Annals and Magazine of Natural History. (8) 9: 686-688. 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 MT CHL CEO Ute Satter era Sate cacao ceiosteci ch mete ack Ae ah PP ain ein cate Bh sraiga Iainjea’s ele acloas.s srastgolant-wmneiesgaduastieee> demeasmaee iesk 111 De SECM ACI CHAN C COLIN ae asco latte oe nies sate ast cat canoes icles ao-ols Sis gs aa le Sloss oie a ome claee Sate lames naisinciows eres amas ebecinesekE es 112 AMON GUICSLOVEFTA ECA NACI wo ONG nseixctas ctaclas ape@arls aides usiapast cia yediaseerssccesancewaie eose ease cas vosaGenssea senescent 1, DAS Ul ELON arctan create ate tanto snc canis oseGionlocttasis clganecue dole Gone ares sais sciqos gate aces caver seaes soem sar Secon 112 DOM estt A ere ena tana Se teri nectar ean ee ae ae ceense ena coaee see Shas rece Mee kes hamae Josenns ra seeu anemtae vioamblotcctoeaececa ase sete 112 DESCRIP OM een cncn cs anecaacneo etn ene res caseNn dean cnassece-ncaccecarcscesibanaucastiagt tis anctsic wecinesieeioeesecteye~ sep Magee er asicaean see eR 1S SCONOPIGAL NOLES” ocr crs-sraraermacecotctegaec esse See ates re sie cincis Meee fot ac sete seteiastion: setcesncumtissnedstecarettettcs 117 SAO MOMMCINCIIAUK SMM Meentes morse me edtcsr cece tentthetccar stone ssunvat anmosumceeer Seton cneemecsadtte acana cacti ce te wiaenains scaueeeete 117 PACKNOWICUMEMEMUS tere terecseteaserecetcctscs ccacceadestsarstesscarecasecathtosececuerstureassusterpedmain cauaseded: + eau ecansce ns ot 118 IREECKELICES eee ere ae MR RSNA sc ete TCA FESS E NO Roc eee ta nines qolgestnadeshidbignedemedea meus Gtoancge athae cute eukebepenatede ns eaenet 118 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 © The Natural History Museum, 1995 S. aubryi in the literature (A. Milne Edwards, 1853; De Man, 1901; Bott, 1955; Monod, 1977, 1980) have led to the misidentification of specimens of S. floweri as S. aubryi by parasitologists (Voelker, et al., 1975, fig. 6; Voelker & Sachs, 1977, fig. 4). The right mandible and the right first and second gonopods of the type of S floweri were removed to illustrate these structures from different angles and under magnification (Fig. 2 a-i). Specimens of S. floweri from Nigeria collected by the author were either dug from their burrows at the sides of streams, or were trapped in fishing nets set overnight in ponds. One specimen (NMU 9.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. REFERENCES Balss, H. 1914. Potamonidenstudien. Zoologische Jahrbiicher, Abteilung fiir 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 Kongogebeites. Revue du Zoologie et Botanie dAfrique 28: 65-204. Bott, R. 1955. Die Stisswasserkrabben von Afrika (Crust., Decap.) und ihre Stammesgeschichte. Annales du Musée Royal du Congo Belge, (Tervuren, Belgique,) C-Zoologie Série III, III 1(3): 209-352. 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. Capart, A. 1954. Révision des Types des especes de Potamonidae de I’ Afrique Tropicale conservés au Museum d’Histoire Naturelle de Paris. Volume Jubilaire Victor Van Strallen, Director de l'Institut Royale des Sciences Naturelles de Belgique, 1925-1934, II: 819-847. Chace, F. A. 1942. Scientific results of a fourth expedition to forested areas in eastern Africa, III: Decapod Crustacea. Bulletin of the Museum of Compara- tive Zoology, Harvard College 91(3): 1285-233. Cumberlidge, N. 1991. Sudanonautes kagoroensis, a new species of fresh-water crab (Decapoda: Potamoidea: Potamonautidae) from Nigeria. Canadian Journal of Zoology 69: 1938-1944. — 1993a. Redescription of Sudanonautes granulatus (Balss, 1929) (Potam- oidea, Potamonautidae) from West Africa. Journal of Crustacean Biology 13(4): 805-816. — 1993b. Further remarks on the identification of Sudanonautes orthostylis (Bott, 1955),with comparisons with other species from Nigeria and Cam- eroon. Proceedings of the Biolological Society of Washington 106(3): 514-522. De Man, J. G. 1901. Description of a new fresh-water Crustacea from the Soudan; followed by some remarks on an allied species. Proceedings of the Zoological Society of London: 94-104. Flower, S. S. 1931. Notes on Fresh-water crabs in Egypt, Sinai, and the Sudan. Proceedings of the Zoological Society of London: 729-735. Herklots, J. A. 1861. Symbolae carcinologicae. Etudes sur la classe des Crustacés: 1-43. Leiden. Parisi, B. 1925. Un nuovo Potamonidi dell Abissinia. Atti Societi Italia Scienca Naturello, Museo civice Milano 61: 332-334. Milne Edwards, A. 1869. Révision du genre Thelphusa et description de quelques especes nouvelles faisant partie de la collection du Muséum. Nouvelles Archives du Muséum dHistoire naturelle, Paris 5: 161-191. Milne Edwards, H. 1853. Observations sur les affinitiés zoologiques et la classification naturelle des Crustacés. Annales des Sciences Naturelles, Zoologie, Paris, Série 3, 20: 163-28. Monod, T. 1977. Sur quelques crustacés Decapodes africaines (Sahel, Soudan). Bulletin de Muséum national d Histoire naturelle, Paris 3, 500: 1201-1236. 1980 Décapodes. In: Flore et Faune Aquatiques de I’Afrique Sahelo- Soudanienne, 1: 369-389. Ed. J-R. Durand and C. Léveque, ORSTOM, I.D T. 44, Paris. Nozais, J. P., Doucet, J., Dunan, J. & Assale N’Dri, G. 1980. Les paragoni- moses en Afrique Noire. A propos d’un foyer recent de Cote d'Ivoire. Bulletin de la Sociétié de Pathologie exotique, 73: 155-163. Rathbun, M. J. 1904. Les crabes d’eau douce (Potamonidae). Nouvelles Archives du Muséum d'Histoire naturelle (Paris) 6(4): 255-312. REDESCRIPTION OF AFRICAN FRESH-WATER CRAB — 1905. Les crabes d’eau douce (Potamonidae). Nouvelles Archives du Muséum d'Histoire naturelle (Paris) 7(4): 159-322. — 1921. The brachyuran crabs collected by the American Museum Congo expedition 1909-1915. Bulletin of the American Museum of Natural History 43: 379-474. Voelker, J. & Sachs, R. 1977. Uber die Verbreitung von Lungenegeln 119 (Paragonimus africanus und P. uterobilateralis) in West-Kamerun und Ost-Nigeria auf Grund von Untersuchungen an Siisswasserkrabben auf Befall mit Metazerkarien. Tropenmedizin und Parasitologie 28: 129-133. Voelker, J., Sachs, R., Volkmer, K. L., & Braband, H. 1975. On the epidemiology of paragonimiaisis in man and in animals in Nigeria, West Africa. Veterinary Medical Review 1-2: 161-175. 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; © The Natural History Museum, 1995 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) oo 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) No 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 A oat y sc 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 z= SE SE oe Go Ai 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 SP 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 A oS ——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 DD =, 2 Sa f LE, Dike NPA PAS PASI SIA) CAI = SS SS > =e PM 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 DD SCM? SSS Lh gk 7 2f Pf 6 LEEE ZEEE AGREE Zi, IN ia LE, LEE z= 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 PM DI EARS DI PM R.D. MOOI AND A.C. GILL b Fig. 12 Epaxial muscle morphology in: a, Lates niloticus (ROM 28524, 80.8 mm SL); b, Psammoperca waigiensis (ROM 46627, 91.2 mm SL). Note the insertions on to the second pterygiophore just ventral to the spine/pterygiophore articulation. Abbreviations and other methods of presentation as in Fig. 1. Scale bars = 5 mm. SCA ///. SP 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 PM 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 a NW Al =—= —————— 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 a SCA Zz Lube CLUE FX Hf (7, Af Wi 2222 a2 bf, DD ——— = b SP1 SCA ED 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| | l) f} f fa| a = — SJ a) if 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 Zz Lp ED Zu 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). REFERENCES Allen, G.R. & Swainston, R. 1988. The marine fishes of north-western Australia. 201 pp. Western Australian Museum, Perth. Anderson, M.E. 1984. 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Bulletin of Marine Science 52(1): 395-414. Yabe, M. 1985. Comparative osteology and myology of the superfamily Cottoidea (Pisces: Scorpaeniformes), and its phylogenetic classification. Memoirs of the Faculty of Fisheries, Hokkaido University 32(1): 1-130. wat GUIDE FOR AUTHORS Policy. The Bulletin of the British Museum (Natural His- tory) Zoology, was established specifically to accommodate manuscripts relevant to the Collections in the Department of Zoology. It provides an outlet for the publication of taxo- nomic papers which, because of their length, prove difficult to publish elsewhere. Preference is given to original contribu- tions in English whose contents are based on the Collections, or the description of specimens which are being donated to enhance them. Acceptance of manuscripts is at the discretion of the Editor, on the understanding that they have not been submitted or published elsewhere and become the copyright of the Trustees of the Natural History Museum. 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Smith 111 Redescription of Sudanonautes floweri (De Man, 1901) (Brachyura: Potamoidea: Potamo- nautidae) from Nigeria and Central Africa N. Cumberlidge 121 Association of epaxial musculature with dorsal-fin pterygiophores in acanthomorph fishes, and its phylogenetic significance R.D. Mooi and A.C. Gill Bulletin of The Natural History Museum ZOOLOGY SERIES Vol. 61, No. 2, November 1995