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Hyobranchial Apparatus of the Cryptobranchoidea (Amphibia)

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Hyobranchial Apparatus of the Cryptobranchoidea (Amphibia) Great Basin Naturalist Volume 49 Number 4 Article 2 10-31-1989 Hyobranchial apparatus of the Cryptobranchoidea (Amphibia) Douglas C. Cox Brigham Young University Wilmer W. Tanner Brigham Young University Follow this and additional works at: https://scholarsarchive.byu.edu/gbn Recommended Citation Cox, Douglas C. and Tanner, Wilmer W. (1989) "Hyobranchial apparatus of the Cryptobranchoidea (Amphibia)," Great Basin Naturalist: Vol. 49 : No. 4 , Article 2. Available at: https://scholarsarchive.byu.edu/gbn/vol49/iss4/2 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. HYOBRANCHIAL APPARATUS OF THE CRYPTOBRANCHOIDEA (AMPHIBIA) Douglas C. Cox and Wilmer W. Tanner Abstract. —This report is a comparative study of the hyobranchial skeleton of the three species in the family Cryptobranchidae and four species of the family Hynohiidae, all of which are figured. Anatomical differences between the cryptobranchids and their relationship to hynobiids are noted and discussed. Reference is made to a relationship that may exist between these families and other salamander families, particularly the possible relationship of the Hynohiidae to the super families Ambystomatoidea or possibly Plethodontoidea. The primary purpose of this study was to Recent studies also indicate that the crypto- examine and compare the hyobranchial skele- branchids are primitive in that they have a ton of representatives of the families Crypto- large number of chromosomes (60). Mores- branchidae and Hynobiidae. We were con- calchi et al. (1977, 1979) give the structure and cerned not only with the anatomical rela- numbers of chromosomes for the three cryp- tionships of the two families, but also with the tobranchid species, with each having 60, and basic differences between available genera list the following hynobiid species and their within the families. Much has been done to chromosome numbers: Ranodon sibiricus, 2n establish an understanding of anatomical rela- = 66; Batrachuperus mustersi, 2n = 62; Ony- tionships primarily involving skeletal and soft chodactylus japonicus, 2n = 58 ± 2; Hyno- tissue comparative anatomy of other families, bius dunni, H. nebulosus, and H. tsuensis, particularly Ambystomatidae, Plethodonti- each with 2n = 56. A study by Sessions et al. dae, and Salamandridae. The studies by (1982) provides essentially the same informa- Humphry (1872) and Dunn (1923, 1926) were tion, listing 30 pairs of chromosomes in the followed by research of Noble (1931), Francis genus Andrias as compared to 11-14 pairs in (1934), Edgeworth (1935), Piatt (1935, 1939, other families of North American salaman- 1940), Taylor (1944), Tanner (1950, 1952), Fox ders. Taketa and Nickerson (1973) deter- (1954), Wake (1966), Wake and Ozeti (1969), mined the relative electrophoretic mobilities Nickerson and Mays (1972), Estes (1981), Du- of the hemoglobins of three families of sala- ellman and Trueb (1986), and others cited manders represented by Cryptobranchus, below, involving cytogenetic studies with the Necturus, and Hynobius, when compared intent to provide data showing familial rela- with adult human HbA, at pH 8.4. It is note- tionships. worthy that Cryptobranchus has a single com- The examination of the relationships of the ponent with greater mobility than the two Cryptobranchoidea by Dunn (1923) estab- components in Hynobius tsuensis. lished the primitive characters of external The relationship of the cryptobranchids to fertilization and retention of the angular bone other families of salamanders and particularly in the lower jaw. These characters have not to the hynobiids was noted by Dunn (1923). only placed the cryptobranchids and hynobi- He indicated that cryptobranchids are more ids as primitive groups of salamanders, but closely related to the hynobiids than perhaps also as possible descendants of ancestral stock to other salamander families. While this may similar to those from which other salamander yet be true, there are substantial differences lines might have arisen. Dunn (1923) also between the basic structures of the throat noted the following: both families possess anatomy of the hynobiids and the crypto- nasals that meet at the middorsal line, and the branchids. This is particularly evident in the premaxillary spines are short in contrast to bony and cartilaginous structures. If there other families with separate nasals and long is indeed a close relationship, it is apparent- spines. ly one based on such characters as external Life Science Museum, Brigham Young University, Provo, Utah 84602 482 October 1989 Cox, Tanner: Salamander Anatomy 483 fertilization, retention of the angular bone, and greater number of chromosomes, and not on the hyobranchial structures, which have undergone major modifications that would suggest not only an extended period of isola- tion, but certainly one in which major adap- tive modifications have occurred. Further- more, there has been a major radiation in the family Hynobiidae that has not occurred among the cryptobranchids. The fossil history of the development of the various families of salamanders indicates that there may have been a movement of repre- sentatives of two major families from North America into the Orient. Certainly North America has been a center for major salaman- der evolution, with all families (except Hyno- biidae) being represented. Only four of the seven families have extended their distribu- tions beyond the North American shores (Plethodontidae—Europe; Ambystomatidae, Fig. 1. Ventral view of the hyobranchial apparatus of = = Cryptobranchidae, and Hynobiidae—Asia). Cryptobranchus alleganiensis. M lower jaw; BH basihyal; Ul\ hypohyal; BB1 = basibranchial #1; BB2 The families Ambystomatidae and Crypto- basibranchial #2; CH = ceratohyal; CB1 = eerato- Asi- branchidae are represented on both the branchial #1; CB2 ceratobranchial #2; EB1 = epi- atic and American continents, with the Asiatic branchial #1; BB3 basibranchial #3; EB2 - epi- cryptobranchids having developed giantism branchial #2; EB3 epibranchial #3. and undergone a more complete morphologi- cal development than the American represen- tatives. The place of origin for the Cryptobran- choidea and particularly the family Crypto- branchidae is as yet uncertain. Meszoely (1967) described as new the genus Piceoer- peton from fossil vertebrae obtained from the early Eocene of Wyoming and included it in the family Cryptobranchidae. Within the fam- ily it is related to the genus Andrias. Estes (1969) maintained that the relationship of the genus Piceoerpeton is better placed in the family Scapherpetontidae, a representative of Ambystomatoidea. The occurrence of An- drias in Europe and Japan (Westphal 1958) suggests a previously wide distribution for the family and perhaps for the genus Andrias in North America, although this has not as yet been demonstrated. An examination of the throat skeleton sug- gests that Andrias has advanced beyond the semilarval condition seen in C ryptobranchas with its retained gill bars (Figs. 1-3). The two oriental species (A. davidianus and M.japoni- cus) have a greater similarity in hyobranchial structures than either has with adult C. alle- ganiensis, and both are much larger. 484 Great Basin Naturalist Vol. 49, No. 4 hypohyals. The hypohyals are enlarged and curved, meeting at the midventral line and also forming a broad joint with the cera- tohyals. The basihyals have been pushed caudad and lie on the medial border of the ceratohyals and the posterior border of the hypohyal. There is some doubt as to the proper names to be applied to these carti- lages; a final determination may require a careful examination of the embryonic devel- opment of these structures in this species. An examination of the position of the hy- obranchial apparatus in the two families (Figs. 1-4) indicates a noticeable posterior shift of all structures in the family Hynobiidae. This has raised a question as to whether the structure we have labeled the first basibranchial in Fig- Fig. 3. Ventral view of the hyobranchial apparatus of Andrias davidianus. ures 1 and 2 is a basibranchial or a separate segment of the first arch. In any case it is not possible to relate these structures (basi- on the primitive characters indicated above branchials) to homologous structures in hyno- and not on the skeletal characters of the biids using only adult specimens. throat. The structure of the hyobranchial apparatus The following is a description of the struc- has been illustrated as found it to occur in tures of the hyobranchial apparatus as ob- we the dissections of Andrias davidianus, Mega- served in some members of the superfamily Cryptobranchoidea. lobatrachus japonicus, and Cryptobranchus The hyobranchial apparatus of the family alleganiensis (Figs. 1-3). The semilarval con- Cryptobranchidae consists of cartilaginous dition of the American hellbender (C. alle- and bony elements. It has the following fea- ganiensis) is evident by the presence of gill tures: basibranchials, ceratohyals, hypohyals, bars, which are ossified, and by the remaining basihyals, ceratobranchials, and epibranchi- gill slit (Fig. 3). These structures were found als. The first basibranchial is a small cartilage in both C. a. alleganiensis and C. a. bishopi. located in the anterior median area of the
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