1 Pacific Science (1984), vol. 38, no by the University of Hawaii Press. All rights reserved Structure, Function, and Ecology in the Goatfishes (Family Mullidae)! WILLIAM A. GOSLINE 2 ABSTRACT: The Mullidae differ from other percoid families in a number of structural features. Th e most notable ofthese is a pair ofhighly developed hyoid barbels used in feeding. Many ofthe other goatfish specializations seem to be associated in one way or another with the use made of these barbels. Structural peculiarities of the Mullidae are described and their functional and ecological implications suggested where possible. The hypothesis is made that the goatfishes have evolved a distincti ve ecological niche for themselves based on their use of the barbels in hunting. MOST OF THE50 ORSOfamilies of percoid fishes are separated from one another by minor structural features (Regan 1913). The goatfishes (Family Mullidae) differ dra stically from this general pattern. They have developed a structur al peculiarity unique among percoids, the hyoid barbels, and a number of other specializations, several of which appear to be functionally or structurally associated with the barbels.the adaptations of the Mullid ae seem to have been rather successful, for goatfi shes are represented circumtropically in inshore areas, often by large numbers of individual s. Within the family, there appears to have been a minor adaptive radi ation based on the exploitation of the barbels for various types of feeding because the principal differences between the six otherwise quite similar genera of the family are in dentition. A pair of well-developed barbels is present in all adult goatfishes. Th e two members of each pair are independently movable and bear numerou s sense organs (Sato 1937). When a goatfish is not feeding, the barbels are folded back under the rims of the chin and gill covers. In feeding, the barbels are lowered to touch the bottom under the fish. Probably all goatfishes use their barbels to detect food items on or slightly below the surface of the substrate (Figu re 1). When an area with such items is located, different goatfishes adopt various methods for obtaining the prey. Some, such as I Manu script accepted I July University of Michigan, Museum of Zoology, Ann Arbor, Michigan Upeneus tragula, shown in Figure 2, use their barbels as an excavating device (see also the comments of Hiatt and Strasburg [1960: 84] concerning Parupeneus cyclostomus). In Hawaii Parupeneus cyclostomus (formerly P. chryserydros) pokes its barbels into holes and crevices in hard substrates to dislodge small an imals (Hobson 1974). Mulloides fiavolineatus (Mulloidichthys samoensis auctorum) may back off from an item discovered by the barbel s (Figure 1) and " blow" away sand to uncover it (pers. obs.); the same species may also burrow into the sand with its snout to retrieve a food item (Hobson, pers. comm.). Mullus surmuletus on occasion ploughs up the mud with its snout in feeding and appa rently expels water from its mouth while doing so (LoBianco 1907). These feeding methods are only used by half-grown and adult goatfishes, for all mullids so far as known have pelagic eggs, larvae, and ju veniles (Caldwell 1962). Because of the use that goatfishes make of their barbels and functionally associated characters, the interrelationship between structure, function, and ecology in the Mullid ae is particularly close and is stre ssed here. On the oth er hand, the distinctiveness of the family seems to have obscured evidence for relationship between the Mullidae and other percoid s, and an attempt to determine these relationships has not been very successful. In the present pap er the various peculiarities of the goa tfishes are described beginning with the barbels. The se peculia rities are compared with similar developments elsewhere in teleosts and where possible are interpreted in terms of
2 Structure, Function, and Ecology in Goatfishes-GosLINE 313 FIGURE 1. Mulloidesflavolineatus probing for food (Kona, Hawaii). Photograph by E. S. Hobson. FIGURE 2. Upeneus tragula using its barbelsas an excavatingtool (UjungoffPandang, Sulawesi, formerly Makassar, Celebes). Photograph by J. E. Randall. function. Comments on mullid genera, family relationships, and ecology conclude the paper. The material on which the article is based consists of partly dissected and whole specimens ofall six mullid genera in the University of Michigan collections. The scientific names used in the paper are those suggested by Randall (pers. comm., March 4, 1984). The name changes of concern here are as follows: Parupeneus chryserydros is placed in the synonymy ofp. cyclostomus and Upeneus arge in that of U. taeniopterus; Mulloides is considered a valid generic name, replacing the more widely used substitute Mulloidichthys. The question ofwhether the genus Parupeneus should be synonymized with the closely related Pseudupeneus is left open. THE BARBEL APPARATUS There are two mechanically separate components to barbel manipulation in goatfishes (Figure 3 A, B). One has to do with the movements of the barbels (A, AI' and A z) relative to the hyoid bars with which they articulate. Each of the two barbels has its own musculature that controls such movements, thus enabling the two to move independently. The second mechanical component (B, B I, and
3 314 PACIFIC SCIENCE, Volume 38, October FIGURE 3. Diagram to illustrate goatfish barbel mechanics, Lateral view from right side. Stippled areas represent bone. Arrows indicate direction of movement. A, barbel with the ligaments that (A I) retract and (At) lower it. B, hyoid bar with the urohyal (B I ) that lowers and the M. prot ractor hyoideus (B t ) that raises it. B z) cau ses the lowering and raising of the anterior ends ofthe pair of hyoid bars relative to the head of the fish. The barbel itselfconsists ofa fleshy, sensory part that enclo ses a long, tapering, distally flexible branchiostegal ray. A large nerve enters the base of the fleshy part and innervates the numerous sense organs on the surface of the barbel. The modified branchiostegal ray (La Bianco 1907) provides a strengthening support for the barbel. It originates in a cap of fibro-cartilage which forms a socket over the tip of a forward projection from the anterior ceratohyal of the hyoid bar (Figure 4). Two ligaments insert on the cap of fibro-cartilage at the base of the barbel, one above the ceratohyal projection and one below. These ligaments lead back into musculature originating along the hyoid bar (not shown in Figure 3). Contraction of the muscle to the upper ligament (Figure 3A z) lowers the barbel whereas contractionofthe muscle to the lower ligament (A 1) retracts the barbel. The muscles to both ligaments appear to be modified parts of the hyohyoideus system (Winterbottom 1974a). The second component of the barbel apparatus governs the movement of the anterior ends of the hyoid bars. Generally in teleosts the hyoid bars are movable at both ends. An teriorly they are connected to the symphyseal area of the chin by musculature (the M. protractor hyoideus of Winterbottom 1974a; the M. genioh yoideus of Osse 1969); posteriorly to the inner surface of the suspensorium by the intercalation of a movable interhyal bone. In the Mullidae the freedom of movement ofthe anterior ends ofthe hyoid of the hyoid bars has been increased and that of their posterior ends reduced to rot ation in a firm socket. The structural changes associated with the
4 Structure, Function, and Ecology in Goatfishes-e-Go suxs 315 Ac Uh B2Hy B1 Uh Lh Ac FIGURE 4. Bones associated with the forward end of the hyoid apparatus in Pseudupeneus macula/ us. A, lateral and B, dorsal view. Only the basal part of the branchiostegal ray is shown. Ac, anterior ceratohyal; BI, first basibranchial; B2, second basibranchial; Br, branchiostegal; GI, glossohyal; Hy, first hypobranchial; Lh, lower hypoh yal; and Uh, upper hypohyai. hyoid bar movements of goatfishes are as follows. The posterior end of each hyoid bar is restricted to a socket formed primarily by a semicircular, medianly projecting flange on the inner surface of the interopercle (Figures 3, 5). In front of this flange, the posterior end of the bar is propped from above by the interhyal bone. Anteriorly, the forward end of the pair of hyoid bars is connected with the chin only by flexible membrane. The protractor hyoideus muscle that is usually in this area has moved its posterior attachment ' well back along the shaft ofthe hyoid bar (Figure 3B 2 ). It still serves to raise the hyoid bar, but the posterior displacement of its attachment permits considerably greater movement of the anteriorends ofthe bars than in other teleosts. The cause of such lowering is, as usual, contraction of the sternohyoideus muscles acting via the urohyal bone (Figure 3B 1 ). Lh A GI Teleosts have evolved ventrally directed projections ofvarious types, usually or always equipped with sense organs. Such projections range from barbels on the chin to extensions ofthe pectoral rays. However, sensory probes developed in association with branchiostegal rays are rare and occur, to my knowledge, only in the Mullidae and in the beryciform genus Polymixia. That the hyoid barbels in these two taxa have evolved independently is clear from a number ofstructural differences between them. Furthermore, the barbel apparatus of the Mullidae is much more specialized than that ofpolymixia. In Polymixia three modified branchiostegal rays support the barbel rather than the one ray in the Mullidae (Starks 1904). In Polymixia the hyoid bar has the usualmovable attachment at both ends rather than the mod ifications described above in the Mullidae. There is also a considerable difference between the two taxa in the basal articulation of the barbel and in the bones at the anterior end of the hyoid bar. In Polymixia the hypohyal bones hold their usual position, one under the other, at the anterior end of the hyoid bar, and the three branchiostegal rays supporting the barbel have simply moved forward to an articulation with the under surface of the lower hypohyal (Starks 1904: fig. 4). In the larval goatfish Mullus surmuletus the branchiostegal rays have their normal serial arrangement (LoBianco 1907), but during growth the anterior branchiostegal moves forward along the hyoid bar and becomes widely separated from the other branchiostegal rays. This forward movement ofthe anterior branchiostegal ray is apparently accompanied by the development of a forward projection from the lower part ofthe anterior ceratohyal, because in the adultgoatfish, this projectionextends anterior to the two hypohyals which themselves have become horizontally aligned (Figure 4; see also Starks 1904: fig. 5). The result of these changesis that in goatfishes the barbel articulation has moved forward to about the level of the tip of the glossohyal (concealed under the membrane of the floor of the mouth) and forward of the hypohyals (Figure 4).
5 316 PACIFIC SCIENCE, Volume 38, October t--0 p ---,~--\--,t p 0 /I..;.-----,,~ j'-- Ih Qu 8y Hy 10 F IGURE 5. Internal view of the suspensorium of Mullus barbatus with the posterior end of the hyoid bar and the interhyal included. A w, part of the A w section of the M. adducto r mandi bulae; ca, cartilage; Ec, ectopterygoid; Hm, hyomandibular; Hy, posterior part of the hyoid bar; Ih, interhyal; 10, interopercle; Ms, mesopterygoid; M t, metapterygoid; Op, opercle; Pa, palatine; Po, preope rcle; Qu, quadrate; So, subopercle; st, strut on the mesopterygoid that abuts against the back of the lateral ethmoid; and Sy, symplectic. There are two points to be made about the forward articulation of the barbel in goatfishes. First, the greater the distance of the articulation from the fulcrum at the posterior end of the hyoid bar, the greater the amount the barbel will be lowered when the bar itself is rot ated through a given angle. Presumably, then, the extent to which the hyoid bar can move the barbel do wnward has been increased by the forward displacement of the barbel articulation. The second point concerns the possible structural relationship between the anterior elongation ofthe hyoid bar and the lengthening of the preorbital part of the head that appears to have occurred in the Mullidae. In goatfishes the snout region above the mouth has apparently become extended as have the hyoid bars below the mouth, and the possibility seems high that the two features are related. THE LENGTHENIN G OF THE SNOUT REGION Th e snout length in various goatfishes is quite different, ranging from short and blunt in Mullus to con siderably extended in Parupeneus and Pseudup eneus. Nevertheless two osteological features present in all goatfishes suggest that the preorbital part of the head has been lengthened in the mullid ancestral stock. One is in the superficial circumorbital series of bon es. Usually in teleosts, the anteriormost member of the series, the lacrimal, articulates with the lateral ethmoid in fron t of the orbit, and the infr aorbital bone behind it is well separa ted from the lateral ethmoid. In the Mullidae, the lacrimal seems to have shifted forward, for it has only a slight contact with the lateral ethmoid, wherea s the infraorbital behind it has the main contact with the lateral ethmoid (Smith and Bailey 1962).
6 Structure, Function, and Ecology in Goatfishes-GosLINE 317 The other apparently forward shift in the anterior head structures occurs in the suspensorium. Normally, the anteriormost bone in the suspensorium, the palatine, provides the only articulation between the forward part of the suspensorium and the skull. As with the circumorbital series, in the Mullidae a bone behind the palatine, the mesopterygoid, has developed an articulation with the skull (Figure 5). The apparent forward dislocation of the lacrimal and palatine bones in the Mullidae is almost exactly duplicated in another very different group of percoids, the Pentacerotidae (Quinquarius examined). Some pentacerotids, like mullids, also have barbels, but those of the pentacerotids are on the chin, not the hyoid bars. In Quinquarius the anterior (palatine) part of the suspensorium appears to have a normal, firm attachment to the rest of the suspensorium, and the whole compound structure seems to move as a single unit. In the Mullidae, however, the upper part of the palatine is joined to the mesopterygoid strut behind it by a flexible area of cartilage (Figure 5), providing ajoint between the anterior and posterior components of the suspensorium. Such a joint is common in percoids. Furthermore, in the Mullidae the M. adductor arcus palatini, or a more or less separate section of it, extends forward along the inner surface of the palatine. I have not seen such a forward extension elsewhere. It may have to do with the expulsion ofajet of water from the mouth in these fishes. OTHER STRU CTURES ASSOCIATED WITH THE MOUTH In addition to the goatfish specializations already discussed, there are a numberof other peculiarities associated with the mouth that range in distribution from one which is widely held among percoid fishes to at least one feature that appears to be entirely restricted to the Mullidae. Adult goatfishes feed on benthic animals and their mouths open forward and downward. Some ofthe specializations in the snout region appear to be related to their bottom feeding. The lacrimal bone, as in many bottomfeeding percoids, (e.g., the lutjanoid-sparoidhaemuloid series, Johnson 1980), is extended downward so as to overlap a large part of the maxillary shaft. Also, as in the same and other percoid groups, the originally movable nasal bone has in goatfishes a rigid posterior attachment to the skull. LoBianco (1907) suggests that in Mullus this increased rigidity of the anterior part ofthe skull is associated with the habit in this genus of ploughing up the bottom with its snout. Goatfishes have gone one step further along this same line of development than many fishes with rigidly attached nasal bones. They have lost the usual ethmoid-maxillary ligament and have functionally replaced it with a ligament that passes anteriorly from the lateral border of the nasal bone over the palatine prong to an attachment on the forward rim of the maxillary (Figure 6A). This same substitution of ligaments occurs elsewhere in percoids in such fishes as the pentacerotid Quinquarius and in lethrinids (pers. obs. ; see also Johnson 1980). In goatfishes the premaxillae protrude in a more or less downward direction, and a number of specializations seem to be associated with this. The anterior face ofthe vomer has a raised, median, cartilaginous ridge (Starks 1926) over which the ascending premaxillary processes slide. There is a similar ridge in, for example, Menticirrhus, a sciaenid with a downwardly projecting mouth opening. In goatfishes the shaft of the maxilla is about equally deep throughout, the primordial ligament does not pass across it to an attachment on its outer rim, and the palatine prong extends down over the maxilla and has a more or less expanded tip (Figure 6A). These features are all present in the sparoid genus Lethrinus. Other peculiarities associated with the mouth ofgoatfishes cannot readily, if at all, be functionally associated with the downward protrusion of the premaxillae. The most distinctive of these is what will here be called a supramaxillary scale (Figure 6A). Like the supramaxillary bone of many lower percoids this specialized scale, present in all Mullidae,
7 318 PACIFIC SCIENCE, Volume 38, October nm --\-\ Po Ay---lr- --~~-'--- M x - -:::j",-...,...:r-----,--7' sm Md- ~ ~_~=::::..--- :;~= rb --rm Mx----_+_ Md----1~ ph cp.figure 6. The adductor mandibulae muscle and topographically associated features in Upeneus moluccensis. A, externa l and B, interna l view. A I ' A 3, and A w ' thre e of the sections of the M. adductor mand ibula e; Ax and A y, the two part s of the usual A 2 section; ap, aponeurosis; Md, mandible; Mx, maxilla; nm, nasal-maxillary ligament; Pa, palatine; ph, upper end of the M. protractor hyoideus; rb, lower branch of the ramu s mand ibularis nerve; rm, ramus mandibularis; and sm, supramaxillary scale. extends upward from the distal end of the maxillary bone. It frequently takes on some of the characteristics of the distal end of the maxillary bone (e.g., sculpturing). Furthermore, it may underlie other, more normally developed scales. It differs from the supramaxillary bone in that its base overlaps the maxilla rather than riding on its upper rim. Furthermore, in Upeneus par vus, which has a particularly large supramaxillary scale, the re are circuli around its anterior surface. Usually in goatfishes there are a number of scales on the maxilla, but in some species of Upeneus only the supramaxillary scale ap pears to be present. I do not know ofa scale of this type elsewhere, but in a number of percoids with a scaleless maxilla there is a cartilagelike upward extension along the dorsal rim of the
8 Structure, Function, and Ecology in Goatfishes-GosLINE 319 distal end of the maxilla (e.g., in the sciaenid Menticirrhus). The configuration of the adductor mandibulae muscle, though inadequately investigated in percoids up to now, can be a good systematic character (see, for example, Johnson 1980); it certainly is within the Mullidae. As in so many small-mouthed higher teleosts (see, for example, Winterbottom 1974b) the M. adductor mandibulae of all goatfishes is complex (Figure 6). In Upeneus molueeensis, which seems to represent the basal goatfish condition in this feature, there are five more or less separate sections of this muscle passing between the suspensorium arid the jaws. The uppermost section in external view (Figure 6A) is undoubtedly an extended maxillaris (A 1) component. It originates on the posterior part of the suspensorium and passes forward to an insertion on the inner surface of the maxilla just behind the proximal head of that bone. The usual A 2 component of the M. adductor mandibulae appears to be represented by two sections (Figure 6 Ax, A y). The upper of the two in external view passes forward to an insertion along the inside of the mandible above A w (Figure 6B). The lower of the A 2 sections in external view almost completely underlies the upper. It passes forward into the aponeurosis with A w and then is continued forward by A w on the inside of the mandible. The mandibular ramus of nerve V passes forward medial to A 1 and the upper section of A, 2 then forward into the mandible between the two sections of A 2, sometimes, as in Figure 6A, with a branch that passes down across the base of the lower section of A 2 ' A 3 is the usual small muscle visible only in internal view. In addition, A w, normally limited to the mandible, has a part attached by ligament to the inner surface of the quadrate bone of the suspensorium (Figure 5). Pseudupeneus (see Smith and Bailey 1962) and Parupeneus differ notably from other goatfishes in that most or all of the A 1 section ofthe adductor mandibulae is attached to the under surface of the infraorbital bones. Such an attachment also occurs in the percoid genus Lutjanus (Smith and Bailey 1962) and certain other members of the Lutjanidae (Johnson 1980), but I do not know of such an attachment elsewhere. THE SPH ENOTI C PROJECTION Goatfishes all have one peculiarity in the bony postorbital rim. In percoids with more or less cylindrical heads (e.g., Perea, Osse 1969), the M. levator arcus palatini originates. on the under surface of the postorbital part of the skull. In fishes with high, compressed heads this muscle usually originates on the posteroventral surface of a lateral projection from the sphenotic which, with the frontal bone above it, forms a continuous rim around the posterodorsal part of the eye. In goatfishes there is a gap, crossed externally by the dermosphenotic, between the sphenotic projection and the frontal borderofthe orbit above it. In some goatfishes (e.g., Mullus, Mulloides, Upeneiehthy s) the M. dilatatlor operculi passes forward through this gap to an attachment on the innersurface ofthe orbitalwall. A similar gap in the postorbital rim occurs in some sparoid fishes (e.g., Calamus, Archosargus) but by no means all. The functional significance ofthis gap is unclear, and its systematic significance, if any, has been too little investigated to determine. STRUCTURES ASSOCIATED WITH TH E FINS Goatfishes always have two, well-separated dorsal fins. They also have a single, minute anal spine and a well-developed scaly proces s in the axil of the pelvic fin. These features, held in common with other percoids, mayor may not have value as indicators of relationship. In two other features associated with the fins, the goatfishes are highly peculiar. The scapula of the pectoral girdle contains either two or three foramina (Starks 1930). Most other teleosts ha ve only one (Starks, 1930, record s two in cirrhitids) through which a major nerve passes (see, for example, Freihofer 1963). Examination of the scapularforamina of goatfishes indicates that the anteriormost corresponds to that of other fishes, and that the other one or two transmit nothing. Their significance is unclear.
9 320 Goatfishes have 13 instead of the usual 15 branchedcaudal rays ofpercoids. A reduction in branched ray count occurs sporadically elsewhere in higher teleosts, for example in anabantoids and Scatophagus. More notable is the pattern of fusion in the caudal skeleton. Externally the caudal fin ofgoatfishes is essentially symmetrical but the caudal skeleton is more than usually asymmetrical (see figures in Monod 1968). The hypurals to the upper caudal lobe are fused to one another and to the urostylar process, but the hypurals to the lower lobe remain separate from one another and from the urostyle. I have not seen this type of fusion in the caudal skeleton elsewhere. The reasons behindthe differentialfusion in the upper and lower caudal lobes of goatfishes can only be a matter for speculation. However, the following possibility seems worth suggesting. Goatfishes have the forked tails of strong swimmers, which they undoubtedly are. Both lobes of the caudal fin are swung from side to side during rapid locomotion, and the fusion in the parts of the caudal skeleton associated with the upper caudal lobe may increase the strengthof the tail beat. However, during hunting (Figure 1) or feeding (Figure 2) it is necessary to adjust the head precisely with reference to the bottom. The relative flexibility of the lower caudal lobe may be used for this purpose by enabling differential movements between the upper and lower caudal lobes during slow forward movements or for lowering or raising the head during feeding. COMMENTS ON GENERA Dentition is the principal character used for differentiating goatfish genera (see, for example, Lachner 1960). Certain rectifications in descriptions of this feature may be noted here. Upeneus has the most extensive dentition among goatfishes and one that is of a generalized percoid type. The genus is said to have teeth on the jaws, vomer, and palatines (e.g., Lachner 1954). Though there are normally teeth on the vomer, they may be few. In a specimen of U. sundaicus I cannot find any at PACIFIC SCIENCE, Volume 38, October 1984 all. The teeth on the palatines, on the other hand, are always well represented. Furthermore, the palatine patch seems always to be continued back on to the ectopterygoid to a greater or lesser extent. Mullus is usually described as lacking teeth in the upper jaw. This, as Caldwell (1962) notes, is only true of adults. The vomer of Mullus is expanded into a broad, medially grooved plate bearing pebblelike teeth. Though the palatines are sometimes said to provide part of the patch of teeth on the roof of the mouth (e.g., Lachner 1960), they are toothless and quite separate from the vomer. Lachner (1960: 3)states that Pseudupeneus differs from Parupeneus in the presence on the upper jaw of a second, outer row of one to several teeth. These may be somewhat reverted and tusklike. What apparently happens here is that the teeth in the single row of juveniles develop irregularly as the fish grows, for I can find no second row of teeth in available specimens smaller than 164mm in standard length. Specimens available seem to have a single row of teeth in the lower jaw, not two or more as stated by Lachner (1960). Two other aspects of differentiation between goatfish genera deserve comment. Upeneichthys, by comparison with other genera, appears to have emphasized perception by the lateralis system, as the old name porosus applied to one of the species indicates. In U. lineatus the whole snout region is covered by a series of moderately large pores and lacks the embedded scales of other goatfish genera. Parupeneus and Pseudupeneus, two closely related genera (or subgenera), differ more radically from other goatfishes and appear to be the most specialized genera in the family. The following specializations may all be associated with their relatively elongate snouts. In addition to the attachment ofthe A 1 section ofm. adductor mandibulae to the infraorbital bones noted above, they have the epaxial body musculature extending farther forward over the top of the head than in other goatfishes (except possibly for some species of Upeneus) and have lost the superficial connection between the preopercular and temporallateralis canals present in other mullids.
10 Structure, Function, and Ecology in Goatfishcs-e-Gostrxs 321 FAMILY RELATIONSHIPS The systematic position of the Mullidae among percoid families is obscure. As already noted, several goatfish specializations appear to be unique among percoids. One of these, the increase in the number of scapular foramina, seems to offer no indication ofrelationship. Two others suggest something about mullid ancestry by implication; Morphologically, the hyoid barbels have no counterpart, rudimentary or well-developed, elsewhere among percoids, but their position and the use made of them suggest that they originated in a bottom-feeding fish. The supramaxillary scale again seems to have no counterpart elsewhere, but its position suggests a substitute development for the supramaxillary bone of many lower percoids and lower teleosts generally. If this interpretation is correct, two implications are at least worth considering: that goatfishes evolved from a stock that (1) had lost its supramaxillary bone and (2) had scales on the maxilla. Other goatfish specializations, or even suites of them, seem to have evolved independently in other percoid groups. This is true of the attachment of the A 1 section of the M. adductor mandibulae to the infraorbital bones in certain mullids and lutjanids. It also appears to be true ofthe several similarities in the anterior part of the head between mullids and the pentacerotid genus Quinquarius: the articulations of both the first infraorbital and the mesopterygoid with the lateral ethmoid, and the loss of the ethmoid-maxillary ligament. Two suggestions concerning mullid relationships have been made. One is with the Lutjanidae by Regan (1913) and the other with the Sparidae by Boulenger (1904). There are far more similarities between goatfishes and the sparoid fishes than between mullids and lutjanids. These similarities are listed below for what they are worth, but to me they do not furnish very convincing evidence of relationship. The possibility of their independent development in two different bottomfeeding percoid groups seems strong. A caveat in this regard is that some of the more notable similarities do not occur between goatfishes and all sparoids, or even with the most generalized sparoids, but rather with now one segment and now another of the sparoid group (Johnson 1980). To start with the least diagnostic, the features held in common are: 1. a well-developed scaly process in the axil of the pelvic fin; vertebrae; 3. two widely separated nostrils on each side ofthe head with the posterior one a valved slit near the orbit as in such sparids as Calamus; 4. the lacrimal bone broadly overlapping the maxilla; 5. the ethmoid-maxillary ligament functionally replaced by a ligament from the nasal bone to the maxilla as in many sparoids; 6. the maxillary blade ofabout equal width for most of its length and overlapped anteriorly by a palatine prong that tends to be expanded at its tip as in the lethrinid sparoids; 7. a gap between the sphenotic projection and the frontal border of the orbit above it as in such sparid genera as Calamus; and 8. the distally expanded symplectic. There are of course numerous differences between the mullids and the sparoids. Perhaps particularly significant among these, so far as possible relationships are concerned, are two characters of the Mullidae that might be expected in a related group but which do not occur in sparoids, namely the two wellseparated dorsal fins and the squamation of the maxilla. To me, a real relationship between mull ids and sparoids seems far from established, but I am unable to suggest a more convincing relative for the Mullidae. STRUcrUR E AND ECOLOGY The structural peculiarities of goatfishes seem to be reflected at several points in their ecology, or vice versa. Goatfishes appear to have evolved a particular feeding niche for themselves. This niche depends on a specialized method of hunting, not on the exploita-
11 322 tion of any particular food source. To judge from reports of goatfish stomach contents (see, for example, Hiatt and Strasburg 1960, Hobson 1974), mullids eat a wide variety of small to medium-sized animals. No goatfish species appears to be a food specialist. (In this. regard the gill rakers and pharyngeal dentition of all goatfishes seem to be of normal, percoid type.) The highly developed hyoid barbels seem to provide the base from which goatfishes have evolved their ecological peculiarities. Man y fishes have sensory probes by means ofwhich they locate food on or in the bottom. The goatfi sh barbel apparatus, however, has evolved beyond analagous structures in other fishes in a number of respects. The hyoid barbels of the Mullid ae are relatively long, strong, and highly movable. The y can be and are used as an excavating device (Figure 2) or for harassing and dislodging prey from holes and crevices in hard substrates. Ind eed, other fishes often take advantage of these mullid barbel activities. Hobson (1974: 962) notes that in H awaii Parupeneus chryse rydros (now P. cyclostom us) is frequently followed around by the jack, Caran x melampygus, "probably as a tactic to capture pre y driven from cover as the foraging goatfi sh disturb th e substratum," and Randall has a photograph ofajack investigating the possibilities of Mulloides foraging in the West Indie s. Equally or more important is the goatfish abilit y to raise or lower the barbel appa ratus (Figure 3) while the fish maintains a hori zontal position abo ve the substrate (Figures 1, 2). Though members of the relatively specialized Parupeneus forage in reef ar eas, the more generalized Upeneus, Mullus, and Mulloides feed over sand or mud. Sandy or muddy bottoms may occur as pockets in hard substrates but are more frequently present as bro ad, unprotected areas. There appear to be two aspects of the relationship between the goatfish barbel apparatus and feeding in such areas. One has to do with escape from predators. Goatfishes do not have effective spines; they do not bury themselve s in the substrate; and they are often far from holes in which they can take refuge. To escape, they simply swim off into open water, a system that PACIFIC SCIENCE, Volume 38, October 1984 depends on rapid acceleration and susta ined speed among other things. The horizontal position of the body above the substrate while hunting or excavating appears to be advantageous here. In moving off, goatfishes do not have first to leave the substrate or to shift from a head-down bod y position (except when they are actually feeding rather than hunting). The same horizontal position of the body above the substra te while locating food also appears advantageous as a method ofhunting over wide areas with occasional sampling of the bottom for food. Indeed, on e goatfi sh uses, at least on occasion, a roving method of hunting in which the barbels are not used at all. Hiatt and Strasburg (1960 : 87) state regarding Upeneus arge (now U. taeniopterus): " It does not probe the sand with its barbels as do oth er goatfishes but rather seeks out benthic crustaceans living exposed on the bottom. Perhaps because it does not probe, it seems swifter than oth er goatfishes." However, U. taeniop terus has weli-developed barbels which it probably uses at least on occasion, as U. tragula (Figure 2) certainl y does. To summarize briefly, goatfish feeding over sand or mud seems to combine attributes of roving predators such as the bon efish (A lbula) and the pompano (Trachinotus) with some of those of excavators such as ophichthid eels and flatfishes. As noted in the introduction, there appears to have been a minor adaptive radiation within the Mullidae. Morphological differences between gener a, together with what is known of their ecology, indicate variations in feeding methods. The relatively extensive dentition of Upeneus suggests a generalized carnivore from which the genera Mullus on the one hand and Parupeneus and Pseudupeneus on the other have become specialized in two different directions. In Mullus the absence of teeth in the adult upper jaw suggests a suction feeder, and its habit of ploughing up the mud is in line with such a feeding method. At the other extreme, the strong jaw teeth near the end of a relati vely long snout in Parupeneus and Pseudup eneus suggest that their teeth are used for immediately seizing active prey that might otherwise escape or be eaten by other fishes. That at least
12 Structure, Function, and Ecology in Goatfi shes-gosline 323 some of the prey of these two mullid genera are relatively large, active animals is indicated by the higher proportion offishes in the stomachs of Parup eneus chryserydros (now P. eyclostomus) than in the stomachs of any of the Hawaiian goatfishes Hobson (1974) investigated. ACKNOWLEDGMENTS I am greatly indebted to three people for help with this paper: John Paxton supplied specimens of Upeneichthys and Quinquarius hendeeaeanthus from the Australian Museum; Jack Randall brought me up to date on mullid nomenclature and provided Figure 2 as well as general information; and to Ted Hobson I am indebted for Figure I and a discussion of goatfish ecology. LITERATURE CITED BOULENGER, G. A Fishes (Systematic account of Teleostei). Pages , figs in The Cambridge natural history, vol. vii. Macmillan, London. CALDWELL, M. K Development and distribution oflarval and juvenile fishes ofthe family Mullidae of the western North Atlantic. Fish. Bull. Fish and Wildl. Ser. 62: , 43 figs. FREIHOFER, W. C Patterns ofthe ramus lateralis accessorius and their systema tic significance in teleostean fishes. Stanford Ichthyol. Bull. 8: , 29 figs. HIATT, R. W., and D. W. STRASBURG Ecological relationships ofthe fish fauna on coral reefs of the Marshall Islands. Eco!' Monogr. 30:65-127, 7 figs. HOBSON, E. S Feeding relationships of teleostean fishes on coral reefs in Kona, Hawaii. Fish. Bull. Fish and Wildl. Ser. 72: ,42 figs. JOHNSON, G. D The limits and relationships of the Lutjanidae and associated families. Bull. Scripps In st. Oceanogr. 24 : I-Il4, 37 figs. LACHNER, E. S A revision of the goatfish genus Upeneus with descriptions of two new species. Proc. U.S. Natl. Mus. 103: ,pls.13, Family Mullidae: Goatfishes. Pages 1-46, pis in L. P. Schultz et ai., Fishes of the Marshall and Marianas Islands, vol. 2. U.S. Natl. Mus. Bull LoBIANCO, S L'origine dei barbigli tattili nel genere Mullus. Atti della Reale Accademia dei Lincei, Ser. 5, Rendiconti, Classe di Scienzi fisiche, matematiche e naturali, 16: , 8 figs. MONOD, T Le complexe urophore des poissons teleosteens, Memoires de l'institut fondamental d'afrique noire 81: 1-702, 989 figs. OSSE, J. W. M Functional morphology of the head of the perch (Perea fluviatilis L.): An electromyographic study. Netherlands. J. Zool. 19: ,15 pis., 24 text figs. R EGAN, C. T The classification of the percoid fishes. Ann. Mag. Nat. Hist., ser. 8, 12 : SATO, M Preliminary report on the barbels ofa Japanese goatfish, Upeneoides bensasi (Temminck & Schlegel). Sci. Repts. Tohoku Imperial University, 4th Ser. (Biology), II: , 3 figs. SMITH, C. L., and R. M. BAILEY The subocular shelf of fishes. J. Morph. 110: 1-18,3 pis. STARKS, E. C The osteology of some berycoid fishes. Proc. U.S. Nat. Mus. 27: ,10 figs Bones of the ethmoid region of the fish skull. Stanford Univ. Pub!., Univ. Series, BioI. Sci. 4: , 58 figs The primary shoulder girdle of the bony fishes. Stanford Univ. Publ., Univ. Series, BioI. Sci. 6 : ,38 figs. WINTERBOTTOM, R. 1974a. A descriptive synonymy of the striated muscles of the Teleostei. Proc. Acad. Nat. Sci. Philadelphia 125: , 56 figs b. The familial phylogeny of the Tetraodontiformes (Acanthopterygii: Pisces) as evidenced by their comparative myology. Smithsonian Contr. Zool. 155, 201 pp., 185 figs.
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