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AMERICAN MUSEUM Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3181, 18 pp., 9 figures November 22, 1996 Ontogeny of the Ethmoidal Region and Hyopalatine Arch in Macrognathus pancalus (Percomorpha, Mastacembeloidei), with Critical Remarks on Mastacembeloid Inter- and Intrarelationships RALF BRITZ' ABSTRACT Ontogenetic changes in head shape and struc- cartilaginous meniscus, which forms the articu- ture during development of Macrognathus pan- lation of the ecto- and entopterygoid with the lat- calus Hamilton, 1822, were investigated. The on- eral ethmoid in juvenile and adult specimens, de- togeny of the skeleton in the ethmoidal and hy- velops ontogenetically from the anterior tip ofthe opalatine complex is described. The following no- pars autopalatina. table features were discovered: pars autopalatina Hypotheses about inter- and intrarelationships chondrifies autogenously and lacks a maxillary of mastacembeloids are reviewed and tested using process at all developmental stages; the unusually ontogenetic data. The phylogenetic position ofSi- situated palatine bone develops without a carti- nobdella among mastacembeloids is reinvestigat- laginous precursor and is considered a dermopa- ed, and the genus is excluded from the Chaudhu- latine; the autopalatine is apparently lacking; the riidae and reassigned to the Mastacembelidae. INTRODUCTION The percomorph suborder Mastacembe- freshwaters of Africa, the Middle East, and loidei consists of the families Mastacembel- Asia (Nelson, 1994). Earlier authors consid- idae and Chaudhuriidae (Travers, 1984b). ered them to be closely related to true eels, The mastacembeloids currently comprise but Gunther (1861, 1880) demonstrated their about 67 species that are restricted to the affinities with acanthopterygian fishes. Nev- ' Postdoctoral Fellow, Department of Herpetology and Ichthyology, American Museum of Natural History. Copyright K) American Museum of Natural History 1996 ISSN 0003-0082 / Price $2.40 2 AMERICAN MUSEUM NOVITATES NO. 3181 ertheless, their relationships within Acan- Melanie Stiassny, Monica Toledo-Piza, Mar- thopterygii have remained obscure. Gosline celo Carvalho, and Maurice Kottelat also (1983) hypothesized that mastacembeloids commented on the manuscript, and the paper are the closest relatives of the swamp-eel was further improved by the thorough re- family Synbranchidae, and this view subse- views of Dave Johnson and Colin Patterson. quently was supported by Travers (1984a, This work was supported by a Kalbfleisch 1984b) and Johnson and Patterson (1993). Postdoctoral Fellowship in the Department Mastacembelids possess a peculiar snout, ofHerpetology and Ichthyology ofthe Amer- a feature noted by early ichthyologists (e.g., ican Museum ofNatural History, New York. Bloch, 1786). It consists of an unpaired me- dian trunklike tentacle flanked by a pair of prominent tubular nostrils. The skeleton of MATERIAL, METHODS, AND TERMINOLOGY the ethmoidal region, also remarkable in I follow Patterson (1977: 79) in the defi- structure and shape, has been described by nition of dermal bone, cartilage bone, and Regan (1912), Sufi (1956), Bhargava (1963), membrane bone, and the terminology I use Roberts (1980), Gosline (1983), and Travers for the different parts ofthe chondrocranium (1984a, 1984b). Although these authors com- is that of Gaupp (1906) and de Beer (1937). mented on the unusual shape and position of Pars autopalatina, pars quadrata, and pars the mastacembelid palatine bone, no studies metapterygoidea refer to parts of the carti- have been conducted to reveal its develop- laginous palatoquadrate, but the terms au- ment. Bhargava's (1958) ontogenetic inves- topalatine, quadrate, and metapterygoid are tigation of the cranium of Mastacembelus restricted to the cartilage bones of the pala- mastacembelus was restricted to the cartilag- toquadrate, as they are in Arratia and Schultze inous skull only. (1991). Autopalatine and dermopalatine re- This paper provides a detailed account of fer to a cartilage bone and a dermal bone, the ontogeny ofthe hyopalatine arch and eth- respectively. The term palatine is used when moidal region in Macrognathus pancalus, an dermo- and autopalatine have fused during Asian representative of the family Masta- ontogeny or when a distinction between the cembelidae. This ontogenetic information is two components cannot be made. then used to evaluate the homology of the Seven wild-caught specimens of Macrog- palatine bone in mastacembelids. Since the nathus pancalus Hamilton, 1822, were ob- study of ontogeny can also provide new in- tained from an aquarium fish importer in sights into the phylogenetic relationships be- Frankfurt, Germany. They were kept and bred tween taxa, Travers's (1984b) hypotheses in a 40-liter tank and fed mainly on aquatic about the intra- and interrelationships of insect larvae. When spawning first began- Mastacembeloidei then were tested with these usually in May or June-the eggs were re- new developmental data. moved, and the hatched larvae were reared in another tank. In the first few days after the beginning of external feeding, larvae fed ACKNOWLEDGMENTS mainly on Artemia nauplii. Their diet was I thank Wolfgang Maier for his support and later changed to Cyclops, Daphnia, and the for the opportunity to maintain the Macrog- larvae ofephemeropterans, plecopterans, and nathus at the Lehrstuhl fur Spezielle Zoolo- dipterans in accordance with their size. gie, Universitiit Tiibingen, and Horst Specimens were fixed in buffered formalin Schoppmann, Universitiit Tiubingen, for his at regular intervals. All measurements refer expert skills in SEM. I am also grateful to to total length. Kai-Erik Witte, who made the specimen of Five fixed specimens of M. pancalus Chaudhuria sp. available for my study, and (AMNH 217413: 8, 12, 17.8, 33, and 107 to Maurice Kottelat, who provided a copy of mm) were chosen for investigation ofthe gross his unpublished manuscript on chaudhuriid ontogenetic changes associated with head osteology. I extend special thanks to Jim Atz shape. They were stained for 10 to 40 seconds for his valuable criticism on the style and in a concentrated solution ofmethylene blue, contents of the manuscript. Gareth Nelson, then excess stain was removed by rinsing in 1 996 BRITZ: ONTOGENY OF MACROGNATHUS 3 water for one to two minutes. In the study of mtp metapterygoid skeletal development, 49 specimens of M. ns nasal septum pancalus between 3.3 mm (newly hatched) pap pars autopalatina and 36 mm (- 3 months) were cleared and pmt+pq pars metapterygoidea and In pars quadrata stained for cartilage and bone. addition, ppq posterior process of quadrate eight specimens ranging from 5.7 to 26 mm Ps parasphenoid were stained with alizarin only. Clearing and q quadrate staining followed the procedure outlined by sy symplectic Dingerkus and Uhler (1977), including the tmarg taenia marginalis modifications for larval material described tr trabecula by Taylor and vanDyke (1985). The eight trcom trabecula communis double-stained specimens ofM. pancalus fig- trh trabecular horn ured in this paper were deposited under v vomer AMNH 217414. For comparison, the following cleared and RESULTS double-stained material also was studied: MASTACEMBELOIDEI: Chaudhuria sp., 42 mm DEVELOPMENT OF HEAD SHAPE IN (AMNH 217415); Macrognathus maculatus, MACROGNATHUS PANCALUS 22, 32 mm (AMNH 217416); Macrognathus With the exception ofJob's (1941) account circumcinctus, 19 mm (AMNH 217412); Si- and some incidental remarks by Polder nobdella sinensis, 53 mm (AMNH 10259) (1963), the development of the highly pecu- and 113, 114, 117 mm (all AMNH 11078, liar snout structure of the mastacembelids caudal fins missing); "Mastacembelus pan- has received little attention, and no detailed calus," 15 mm head length (AMNH 3447); information is available. Mastacembelus armatus, 11 1, 130 mm (both The snout of the newly hatched embryo of AMNH 10274, caudal fins missing); Afro- Macrognathuspancalus is rounded and much mastacembelus congicus, 53,62 mm (AMNH different from that of the adult. Figure 1A 6157). SYNBRANCHOIDEI: Ophisternon aenig- shows the head of a larva of 8 mm in which maticum, 72 mm (AMNH 31573); Synbran- the snout is still rounded and the anterior and chus marmoratus, 47 mm (AMNH 74541), posterior openings ofthe nasal cavity are small 184 mm (AMNH 215280); Monopterus al- apertures in the skin. The lipfolds of the up- bus, 41 mm (AMNH 41579), 164 mm per and lower jaw are well developed. (AMNH 41579). At 12 mm, the snout region is slightly elon- All drawings were made with the aid of a gated (fig. IB). The anterior nasal openings camera lucida attached to a stereomicroscope are now located at the end of ventrally di- (Zeiss SV8). One 9-mm specimen of Ma- rected tubes. crognathus pancalus was prepared for scan- At 17.8 mm, the snout region has elongated ning electron microscopy (SEM) according to still further (fig. 1 C). Its anterior end projects the procedure described by Britz et al. (1995). as a median proboscis-like fleshy appendage with paired anterior nasal tubes on either side. ABBREVLATIONS At this stage the postorbital region ofthe head artmen articulatory meniscus also has undergone significant elongation. cartpr cartilaginous process on lateral At 33 mm (fig. 1 D), the snout region closely ethmoid resembles that of the adult. The entire eth- dp dermopalatine moidal region has continued to elongate and ectp ectopterygoid now projects in the long fleshy appendage
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