PIETSCH: LOPHIIFORMES 



325 



inter-nested sets of vertical lines are used to indicate mono- 

 phyletic units. 



Order Lophiiformes 



Suborder Lophioidei 



Family Lophiidae 

 Suborder Antennarioidei 



Family Antennariidae 



Family Tetrabrachiidae 

 . Family Lophichthyidae 



Family Brachionichthyidae 

 Suborder Chaunacioidei New 



Family Chaunacidae 

 Suborder Ogcocephalioidei New 



Family Ogcocephalidae 

 Suborder Ceratioidei 



As a final note, the Lophiiformes has traditionally been allied 

 with the Batrachoidiformes, based primarily on osteological 

 characters ofthe cranium (Regan, 1 9 1 2a; Gregory, 1933; Rosen 

 and Patterson, 1969). However, this sister-group relationship 

 has yet to be shown conclusively, and 1 have not been able to 

 assess the significance of early life stages in supporting or refuting 

 this hypothesis. 



School of Fisheries WH- 1 0, College of Ocean and Fisheries 

 Sciences, University of Washington, Seattle, 

 Washington 98195. 



Ceratioidei: Development and Relationships 

 E. Bertelsen 



THE Ceratioidei differ most distinctly from all other mem- 

 bers ofthe order Lophiiformes in being meso- and bathy- 

 pelagic, lacking pelvic fins (except in larval Caulophrynidae) 

 and in having extreme sexual dimorphism. Males are dwarfed 

 and differ from females in lacking an external illicium and hav- 

 ing denticular teeth on the tips of the jaws and well-developed 

 eyes and/or olfactory organs. Furthermore, Ceratioidei differ 

 from other Lophiiformes, except the family Ogcocephalidae, in 

 lacking a third cephalic ray and its pterygiophore, and except 

 in the families Caulophrynidae, Neoceratiidae and the gigan- 

 tactinid genus Rhynchactis. females ofthe suborder differ from 

 all other Lophiiformes in having a bulbous swelling of the tip 

 of the illicium (escal bulb) containing a large globular photo- 

 phore. 



The suborder contains approximately 1 34 species placed in 

 34 genera and 1 1 families (Table 89). The taxonomy is based 

 mainly on studies of the females. Except for the larval stages 

 and the basic meristic and osteological characters shared by the 

 two sexes, descriptions require separate treatment of females 

 and males. 



The separation into families is based mainly on osteological 

 characters, of which some ofthe more important are compared 

 in Table 89. Most ofthe families form well-defined and mutually 

 very distinct taxa in which the females (especially) possess unique 

 morphological features which separate them from members of 

 all other families. 



The separation into genera is based mainly on characters 

 present only in females. Somewhat varying between families, 

 some ofthe most important of these characters are differences 

 in: (1) shape of skull and other bones of the head including 

 development of its spines; (2) jaw mechanisms, including den- 

 tition; (3) illicial apparatus, including basic patterns of escal 

 appendages; and (4) pigmentation of skin and development of 

 dermal spines. Some of the distinguishing osteological charac- 

 ters, especially in shape of opercular bones, are shared with the 



males, like the fin ray numbers which in some families show 

 distinct intergeneric differences. The special male structures, 

 such as denticular teeth, show distinct intrageneric differences 

 in full agreement with the separations based on characters of 

 the females. 



The species of Linophryne have been grouped into subgenera 

 and those of Himantolophus (in ms.), Oneirodes. and Gigan- 

 tactis into "species-groups'" based on shared minor differences 

 in one or more of the characters mentioned above. All intra- 

 generic separations of the females into species are based on 

 differences in pattern and shape of escal appendages, often com- 

 bined with differences in illicial lengths. In a majority of the 

 recognized species, no other separating characters have been 

 shown. In others, differences in meristic characters (numbers of 

 fin rays and teeth) and minor osteological characters (shape of 

 opercular bones, development and dentition of branchial arches, 

 etc.) have been observed supporting the separation into species. 

 A special opportunity to check the validity of the separations 

 based on escal characters is found in the genus Linophryne. in 

 which females have hyoid barbels which in pattern of branching 

 show very distinct differences between species and subgenera 

 (Bertelsen, 1982). 



In most cases it has not been possible to separate males into 

 taxa below the level of genera and subgenera. A few males 

 differing from their supposed congeners in special male char- 

 acters (especially denticular teeth) have been tentatively de- 

 scribed as representatives of separate species. Studies of males 

 attached to identified females have not revealed characters of 

 specific order. 



Development 



No spawned, fertilized eggs of Ceratioidei have been described 

 (re-examination by Bertelsen, 1980:66, of an egg referred to 

 Linophryne arborifera by Beebe, 1932:93, indicated that it rep- 

 resents a diodontid). 



