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Descriptive Osteology of Gymnocorymbus Ternetzi (Teleostei: Characiformes: Characidae)

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Descriptive Osteology of Gymnocorymbus Ternetzi (Teleostei: Characiformes: Characidae) © Zoological Institute, St. Petersburg, 2008 Descriptive osteology of Gymnocorymbus ternetzi (Teleostei: Characiformes: Characidae) N.G. Bogutskaya, A.M. Naseka & I.V. Golovanova Bogutskaya, N.G., Naseka, A.M. & Golovanova, I.V. 2008. Descriptive osteology of Gymnocorymbus ternetzi (Teleostei: Characiformes: Characidae). Zoosystematica Rossica, 17(2): 111-128. The purpose of this paper is giving an extensive overview of the cranial skeleton of Gym- nocorymbus ternetzi (Boulenger, 1895) in a form of a formalized scheme that reflect its Bauplan (German for building plan, blueprint; plural: baupläne or bauplaene), a term in biology referring to the common new and original [homologous] properties of the members of a systematic group [taxon]). Each element of the Bauplan can be described by a set of parameters, i.e., size, shape, structure, material composition and position. Though Bauplan is undoubtedly an abstraction, it is a necessary abstraction to be used in phylogenetic analy- sis with preference to “ingroup” and “outgroup” comparisons.The cranial osteology of black tetra Gymnocorymbus ternetzi (Boulenger, 1895) is described based on the study of larvae, juvenile and adult specimens. Provided are Latin terms and some English equiva- lents as well remarks on origin, homology and terminology for each cranial bone discussed. N.G. Bogutskaya, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg 199034, Russia. E-mail: [email protected] A.M. Naseka, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg 199034, Russia. E-mail: [email protected] I.V. Golovanova, St. Petersburg State University, Universitetskaya nab. 7/9, St. Petersburg 199034, Russia. Introduction group. A close relationship between the group, until then referred to as the Ostariophysi, and the Fishes are known to have a morphologically Gonorhynchiformes, was demonstrated by Rosen & complex and highly kinetic skull (Ferry-Graham & Greenwood (1970), based on evidence of the caudal Lauder, 2001). The cranial musculo-skeletal system skeleton, the presence of a fright reaction mecha- of adult teleost fishes consists of about 60 intercon- nism, swimbladder morphology, presence of nuptial nected skeletal parts that are moved by an approxi- tubercles and a striking similarity in the mouth mately equal number of muscles (Aerts, 1991). opening mechanism in Phractolaemus ansorgii Morphological studies on the skeletal morphology (Gonorhynchiformes, Phractolaemidae) and Bivi- of Ostariophysi have already been done by many branchia (Characiformes, Hemiodontidae). The authors, although the nomenclature of the skeletal, fact, however, that in Gonorhynchiformes no “real especially cranial, structures in these papers demon- sign of vertebral differentiation that would suggest a strated a high degree of inconsistencies, which re- condition paralleling the development of Weberian quired an initial priority of this study: to provide a ossicles”, could be observed that convinced them of conclusive nomenclature, which allowed compari- similar functionality, in a broad sense, as the Webe- son with other taxa. Besides, the ontogeny of the rian apparatus, resulted in the subdivision of the skull could provide substantial information concern- Ostariophysi in the Anotophysi (Gonorhynchifor- ing the origin of bones, whereas the study of the mes) and the Otophysi (Ostariophysi s.s.) (Rosen & cranial lateral-line system could give data on the Greenwood, 1970). A first detailed survey of the true nature of canal bones. ostariophysan interrelationships was given by Rob- Regan (1911a, 1911b) considered the Ostario- erts (1973). He stated that “evidence from caudal physi to be a group of species which share the skeleton morphology supports relationships be- common feature of the Weberian apparatus. The tween Clupeomorpha and Ostariophysi, and be- Ostariophysi, according to Regan (1911a, 1911b) tween Clupeomorpha and Gonorhynchiformes, as comprised the suborders Cyprinoidea and the well as between Gonorhynchiformes and Ostario- Siluroidea. The Characiformes, belonging to the physi.” He opposed against the notion that characins Cyprinoidea, were regarded as the least specialised and cyprinids would be more closely related to each 112 N.G. Bogutskaya et al.: Osteology of Gymnocorymbus ternetzi • ZOOSYST. ROSSICA Vol. 17 other than to the catfishes, consequently raising the al., 1992; Cubbage & Mabee, 1996; Bird & Mabee, latter to the same level as the former two. His order 2003), characids (e.g. Bertmar, 1959; Vandewalle et of Cypriniformes than corresponds with the Oto- al., 2005; Miquelarena et al., 2005) and catfish (e.g. physi of Rosen & Greenwood (1970), containing Bamford, 1948; Geerinckx et al., 2005, 2008) (see the three suborders: (1) Characoidei, (2) Cyprinoidei also references cited in these publications). and (3) Siluroidei. A more extensive comparison of The purpose of this paper is giving an extensive ostariophysan groups by Fink & Fink (1991, 1996) overview of the skeleton of Gymnocorymbus ter- lead to a new classification of ostariophysans. Fink netzi (Boulenger, 1895) in a form of a formalized & Fink (1996) gave strong morphological evidence scheme that reflect its Bauplan (German for for the phylogenetic relationships of the Otophysi building plan, blueprint; plural: baupläne or (comprising herein four orders, the Cypriniformes, bauplaene), a term in biology referring to the Characiformes, Siluriformes, and Gymnotiformes). common new and original [homologous] properties Their results show, of the major clades, the fossil of the members of a systematic group [taxon]). Each Chanoides to be sister to all remaining members of element of the Bauplan can be described by a set of otophysans (placed in the Cypriniphysi), with the parameters, i.e., size, shape, structure, material Cypriniformes being the primitive sister taxa to the composition and position (Verraes, 1981; Adriaens, remaining three groups (the Characiphysi), and 1998). Though Bauplan is undoubtedly an abstrac- Characiformes being sister to the clade (the tion, it is a necessary abstraction to be used in phy- Siluriphysi) of Siluriformes and its sister group the logenetic analysis with preference to “ingroup” and Gymnotiformes. “outgroup” comparisons. Several hypotheses have been proposed concern- The nomenclature used for the developing skele- ing the evolutionary distribution pattern of ostario- tal structures is based principally on the works of de physan fishes.As for today, not all groups of Beer (1937), Harrington (1955), Daget (1964), Pat- ostariophysans show an identical distribution. The terson (1977) and Geerinckx et al. (2005). A discus- Siluriformesshow a cosmopolitan distribution, sion on homologies and terminology of the bones is whereas the Gymnotiformes are confined to South given below in respective places. America.The majority of the ostariophysans can be categorised as primary freshwater fishes. Secondary Material and Methods fresh water fishescan be found in Cypriniformes (Cyprinidae), Characiformes and Siluriformes (e.g., The ontogeny of the bones was observed in the Clariidae, Siluridae,Claroteidae, Pangasiidae, Lori- developmental stages of Gymnocorymbus ternetzi. cariidae), whereas peripheral species have been Specimens, obtained from an aquarium shop, were observed in Gonorhynchiformes(Chanidae, fixed in a 4% buffered formaldehyde solution, and Gonorhynchidae) and Siluriformes (Aspredinidae, used for in toto clearing and staining at different Ariidae and Plotosidae) only (Roberts, 1975; time intervals according to Hanken & Wassersug Teugels, 1996; Nelson, 2006). (1981), with trypsine replaced by a 3% KOH solu- The Otophysi included (as calculated in 2005, tion. The examined material ranges from 4.0 mm Nelson, 2006) 64 families, 1,068 genera, and 7,894 norochord length (NL) larvae (9 days after hatch- species. It is now included four orders, Cyprinifor- ing) to 65.0 mm standard length (SL) adults (three mes (eight families: Cyprinidae, Psilorhynchidae, months after hatching). Gyrinocheilidae, Catostomidae, Balitoridae, Cobiti- dae, Nemacheilidae and Botiidae; two latter families Categories of skeletal structures in terms of had earlier been treated as subfamilies of Cobitidae, bone histology and embryonic development later of Balitoridae, and now as distinct family, e.g. Nalbant & Bianco, 1998; Tang et al., 2006; Šlech- Several different types of cartilage could be dis- tová et al., 2008), Characiformes (18 families), Siluriformes (about 15 families). About 64% of all tinguished in Teleostei: (1) cell rich hyaline carti- freshwater fishes can be grouped within the lage, (2) matrix rich hyaline cartilage, (3) fibro/cell- Ostariophysi, where they can be found world wide, rich cartilage, (4) elastic/cell rich cartilage, (5) with exception of Antarctica, Greenland and New ‘Zellknorpel’ (could be observed in the oral barbells Zealand. and the gill filaments), (6) scleral cartilage (support- ing the eye ball) (Benjamin, 1989, 1990). The larval There are many publications, including classical skeleton of fishes is constituted of cartilaginous ones, on descriptive and comparative osteology of fishes of the group Otophysi (e.g. Nawar, 1954; structures. In the whole skeleton, there are about 60 Harrington, 1955; Weitzman, 1962; Vandewalle, cartilage precursors that further develop into ossi- 1977; Cabuy et al., 1999; Huysentruyt & Adriaens, fied structures. The cranial cartilage may be divided 2005; Serra & Langeani, 2006).
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