Environrnrntrrl Biology of Fishes 47: 399-405. 1996. O 1996 Klrrww Auuietnic Publishers. Printed it1 thc Nerherlirr~riv Structures associated with feeding in three broad-mouthed, benthic fish groups William A. Gosline Museum of Zoologx University of Michigan, Ann Arbor, MI 48/09, U.S.A. Received 28.7 1995 Accepted 26.2.1996 Key words: Feeding structures, Flatheads, Goosefishes. Toadfishes Synopsis The flatheads, toadfishes, and goosefishes discussed here hold certain features in common. All are bottom- living forms with depressed head areas and broad gapes, and all eat large food items: fishes andlor crabs. All have developed structural specializations in association with this diet. The three groups are at most distantly related, and their feeding specializations are different and have evolved from different bases. In flatheads the combination of large food items and depressed head regions seems to have led to the separation of the two halves of the pelvic girdle, a feature in which they differ from their scorpaenoid relatives. Toadfish peculiar- ities associated with feeding are various but most notable in those that pass crabs they eat through the gape and into the mouth. Goosefish feeding is centered around the use of a lure to attract prey to within striking distance. The three fish groups are discussed separately, but their feeding structures are compared to one another in the final section of the paper. Introduction traced in the flatheads. In the toadfishes a variety of modifications seem to be associated with feeding. The three groups under consideration here (Fig. 1) In the goosefishes the modifications are also vari- are the flatheads (family Platycephalidae, order ous, but all appear to be centered around the use of Scorpaeniformes), toadfishes (genera Opsanus and a lure to attract prey Batmchoides, order Batrachoidiformes), and goosefishes (family Lophiidae, order Lophii- formes). These groups are distantly related to one Flatheads (Fig. la) another but show certain general features in com- mon. All spend considerable time in contact with The family Platycephalidae contains about 12, the bottom; all have depressed head areas; and all mostly Indo-Pacific genera. The food of the best- eat large food items - fish and/or crabs. The mod- known genus, Platycephalus, consists of fishes and ifications in feeding-related structures in the three crabs, and Marais (1984) remarks on the large size groups are dealt with separatedly and in the context of a fish taken from a Platycephalus stomach. of close relatives. In the flatheads, the least special- Aside from the flattened head areas, flatheads ized of the three, the discussion centers on the sep- appear to be quite normal scorpaeniform fishes. aration of the two halves of the pelvic girdle. The two eyes are rather close together on the top of Though such a separation also occurs in the other the head but the eye pupils are near the base of the two groups, its development is most satisfactorily large eyeballs and presumably provide vision to the side of the fish. The two nostrils on each side are on Onigocia, Platycephalus, Rogadius, Sorsogonia, the upper surface of the head (unlike those of toad- Suggrundus and Thysanophrys, are not firmly unit- fishes). The lateral expansion of the head at the lev- ed posteriorly but merely overlap slightly if at all. el of the mouth provides a very wide gape. The low- This flexibility in the attachment of the two sides of er jaw is flat below and projects somewhat beyond the girdlis is interpreted here as a method of acco- the upper. The dentition is of a generalized type, modating large food items in the stomach directly with bands of fixed, conical teeth on the premaxil- above the girdle. lae, vomer, palatines, mandible, and pharyngeals. The closest relatives of the Platycephalidae are The genus Ratabulus is exceptional among platyce- the Bembridae, and the two groups are often com- phalide genera in having depressible teeth (Knapp bined into a single family (e.g. Masuda et al. 1984). personal communication). The gill openings extend The platycephalids have more flattened head re- far forward under the throat, and it seems clear that gions than the bembrids, longer anal fins without Platycephalus (unlike Opsanus, see below) can low- anal spines, and pelvic fins that are farther back un- er the anterior end of the hyoid arches relative to der the body and more widely spaced, but the most the mandible, for the sternohyoideus muscle is at- striking difference between the two families is in tached to a well-developed urohyal bone, as usual in pelvic girdle structure. Bembrids (Fig. 2b) have fishes. quite normal, scorpaeniform-type pelvic girdles, The most notable specialization of platycepha- with the two halves firmly united posteriorly. Ac- lids, and one in which they differ not only from their cording to the hypothesis advanced here the platy- scorpaeniform relatives but from most higher tele- cephalids should eat larger food items than the osts, is the structure of their pelvic girdles (Fig. 2a). bembrids. That the platycephalids eat large items The two halves of the pelvic girdle, examined in members of the genera Cociella, Grammoplites, Fig. I. Sketch of a typical: a = flathead, b = toadfish, and c = Fig. 2. Left half of the pelvic girdle from below of a = Platycepha- goosefish. lus indicus, and of whole girdle of b = Parabembras curtus. has been noted above. Unfortunately, the food of leosts. Though genera related to the toadfishes feed bembrids appear to be unknown, and the stomachs on a variety of things from sea urchins (Hoffman & of the bembrids opened were empty, but supporting Robertson 1983) to fishes (Lane 1967), Opsanus evidence for the idea that platycephalids eat larger (Schwartz & Dutcher 1963) and Batrachoides (Col- items than bembrids exists in the more capacious lette & Russo 1981) specialize in crabs. stomachs of platycephalis (Fig. 3). Toadfishes are normally in contact with the bot- The separated halves of the pelvic girdle in the tom. as are the crabs. The somewhat forwardly di- flatheads, and in the toadfishes and goosefishes, rected eyes and the anterior nostrils at the ends of represents a reversal in the usual course of teleos- horizontal tubes that project anteriorly to above the tean evolution, where the two halves are movably upper jaw are probably related to the fact that their articulated with one another in lower teleosts but major food items are directly in front of them. To usually firmly united at the percoid-scorpaenoid capture their larger prey toadfishes use a camou- level (Stiassny & Moore 1992). The separation is flage-and-ambush technique, and they must swim discussed in flatheads because there, unlike the directly forward over the substrate to capture such other two groups, it can be compared with the unit- items. Their mouth structures differ in several re- ed girdle halves in the closely-related bembrids. spects from bottom-resting fishes such as the goose- That the separation of the two halves of the pelvic fishes (Lophidae, see below) which seize prey swim- girdle is not caused by the flattening of the head ming overhead. or the goatfishes (Mullidae) which alone is suggested by the fact that in the very simi- swim over their benthic food below them. One of larly-shaped dragonets (Cnllionymus), which eat the toadfish differences is in the way that the man- smaller food items, the two halves of the pelvic gir- dible is lowered to open the mouth. To obtain such dle are united, as usual in higher teleosts. lowering while the mandible is maintained in a more or less horizontal position just above the sub- strate, there must bc a slight raising of the head rela- Toadfishes (Fig. lb) tive to the rest of the body. In toadfishes there is no joint-like arrangement between the head and the In a number of features associated with feeding the vertebral column. Indeed, the skull is effectively toadfishes (Opsarzus and Batrachoides) and their continued into the body by the firm attachment of allies among the batrachoidiform fishes are unusu- the neural arch of the first vertebra to the back of al, and in one of them, the hinge between the lateral the skull. Raising of the front end of the body is ethmoid and the maxilla (Fig. 4), unique among te- probably accomplished by lowering of the pelvic fins (as in Fig. Ib) against the substrate. These fins Fig. 4. Jaws and jaw musculature of Opsanus beta: A, and A,, = sections of them. adductor mandibulae (ap = ascending process of the premaxilla. ma =mandible. mx =maxilla, st = hinged strut Fig. 3. Outline of esophagus and stomach: a = Plarvcc~phalusin- between the ethmoid of the skull and the maxilla. and ta = dicus. b = Purubc~rnbrascurtus. toothed arm of the premaxilla). are well forward under the body and have a fibrous by the nature of the lower jaw, the two sides of pad along the front border of the leading ray. which flare outward in the coronoid areas (Fig. 5). If the front of the lower jaw is pushed forward The most widely flaring parts of these coronoid ar- over the substrate, the already somewhat prognath- eas remain just behind the maxillary attachments as ous mandible extends even farther ahead of the up- the mouth is opened. As a result the width of the per jaw when the head is raised. This discrepancy is gape is maintained as the mouth opens and may probably counteracted for purposes of an initial even be widened a little if the widest parts of the bite by protrusion of the premaxillae. The potential coromoid areas shift somewhat forward relative to problem here is somewhat alleviated by the fact the lateral ends of the maxillae.