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Evolutionary Relationships in the Family Salamandridae Author(S): David B

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Evolutionary Relationships in the Family Salamandridae Author(S): David B Evolutionary Relationships in the Family Salamandridae Author(s): David B. Wake and Neclâ Özeti Reviewed work(s): Source: Copeia, Vol. 1969, No. 1 (Mar. 6, 1969), pp. 124-137 Published by: American Society of Ichthyologists and Herpetologists (ASIH) Stable URL: http://www.jstor.org/stable/1441702 . Accessed: 24/01/2012 13:21 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Society of Ichthyologists and Herpetologists (ASIH) is collaborating with JSTOR to digitize, preserve and extend access to Copeia. http://www.jstor.org Evolutionary Relationships in the Family Salamandridae DAVID B. WAKE AND NECLA OZETI Data on 40 characters, 27 relating to feeding mechanisms, were as- sembled in order to analyze evolutionary relationships in the family Salamandridae. In most instances, primitive and derived states were identified, and in many cases, state phylogenies for each character were hypothesized. Phenetic schemes of character analysis, including single and average linkage techniques and the results of inclusion and exclusion of primitive character states, were compared. There are significant dif- ferences, and exclusion of primitive character states is recommended. We put most emphasis on a combinatorial analysis of our own, which takes into account the changes in the feeding system as a single complex unit and also makes use of the phenetic methods in which primitive character states are excluded. The result is the placement of the 14 genera in four groups: A. Salamandra, Chioglossa, Salamandrina; B. Triturus, Euproctus, Neurergus, Paramesotriton, Cynops, Hypselotriton, Pachytriton, Taricha, Notophthalmus; C. Pleurodeles; D. Tylototriton. The very primitive groups C and D are slightly modified remnants of ancient groups. Group A is the most derived group. Chioglossa and Salamandrina are the most specialized genera. Of two computer methods for deducing phylogenies, one compares well with these results, but the other does not. INTRODUCTION which, for various reasons, appear to carry T HE family Salamandridae includes 14 useful information. Our data from feeding genera and about 43 species, distributed mechanisms do not indicate whether we broadly throughout the Holarctic region. are dealing with an integrated system under Considerable information concerning sala- a relatively simple genetic control, or whether mandrid anatomy and evolutionary relation- the parts (our 27 characters) are discrete ships appeared in the German literature of and evolving independently. Probably state the 1930's, and the monograph by Herre changes in several characters are due to re- (1935) remains the most authoritative ref- sponses to a single selection pressure, but erence on the family. The work of Herre this cannot now be determined. We think and others resulted in the inclusion in the that careful analysis of such an adaptive family Salamandridae of the genera pre- complex will provide useful information for viously placed in the family Pleurodelidae, the deduction of the phylogeny. Feeding is and the distribution of regionalized groups an extremely important activity and several of species of Triturus to several genera. Their clearly adaptive trends in the system are statements concerning salamandrid evolution associated with ecological shifts. In deducing have not been reevaluated, but von Wahlert phylogeny, knowledge of community of (1953) and others added new information descent is sought, and we feel that very that has, in general, tended to corroborate complex systems, where the chance of paral- their findings. Nevertheless, many aspects of lelism and convergence is reduced, offer the evolution in the family have not been best sources for such information. Biological studied, including feeding and locomotor information for systems other than the feed- systems. Data accumulated during our study ing system has been added, for we wish to on feeding mechanisms (Ozeti and Wake, make this study as comprehensive as our 1969) are used in this paper to reinvestigate present knowledge of the group permits. salamandrid relationships and to attempt to Our working hypothesis is that information deduce the phylogeny of the group. concerning relationship in primary groups Descriptive portions of our comparative (e.g., the family Salamandridae) is contained morphological paper are condensed here to only in derived character states. Resem- 27 characters. To these we add an additional blance that results from characters shared 13 that either have been used by others or in states that are primitive for the primary 124 WAKE AND OZETI-SALAMANDRID RELATIONSHIPS 125 group has no phylogenetic information. We salamander groups other than salamandrids. are interested in community of descent with Nevertheless, presence of the arch is con- change, and retention of primitive conditions sidered primitive (0), partial development a is essentially non-descent and provides no derived state (1), and absence a more de- information. Derived states which occur rived state (2). only in single taxa similarly have no infor- 3. Maxillary length. The toothed portion mation concerning relationships, but they of the maxillary bone of amphibians is do provide information concerning degree primitively very long. In the most primitive of derivation of a group; five of our 40 char- salamandrid state the maxillary extends vir- acters fall into this category. tually to the quadrate, or falls just short of Analysis of the 40 characters has been that bone (0). In a derived state the max- based on the premise that a character state illary extends past the eye but falls far of universal occurrence in related, primitive short of the quadrate (1). A more derived families, or in early fossil ancestors, is primi- state finds the maxillary falling short of the tive for the group under consideration. We posterior margin of the eye (2). have rigorously applied this standard, and 4. Nasal bone contact. In hynobiid and feel that the following analysis is conserva- cryptobranchid salamanders the nasals are tive in this regard. Where there is doubt, we in broad median contact. Such a condition is have not identified a primitive state, except considered to represent the primitive state where an apparently clear evolutionary trend in salamandrids (0). Narrow contact is con- exists within the primary group. To facili- sidered a derived condition (1), and no con- tate computer analysis we have coded the tact is a more derived state (2). characters in the following manner (Table 5. Operculum. Opercula are ossified in 1): primitive state (0), derived states (1, 2, primitive salamanders. In salamandrids os- etc.), character not present or inapplicable sification or mineralization is considered or no information (9). Reference to primi- primitive (0), and unmineralized cartilage is tive salamander families applies to the derived (1). families Hynobiidae, Cryptobranchidae, and 6. Caudosacral ribs. There is a general evo- Ambystomatidae, unless specified. lutionary trend in amphibians for reduc- In some instances data have been obtained tion and loss of postsacral ribs, borne on from the literature, especially in regard to caudosacral vertebrae. Accordingly, caudo- osteology (data from Bolkay, 1928; Herre, sacral ribs are considered to represent a 1932, 1933, 1935, 1939; Wolterstorff and primitive state (0) and absence is derived (1). Herre, 1935). 7. Fifth toe. Loss of the fifth toe and fifth distal tarsal is clearly a derived con- CHARACTER ANALYSIS dition (1), since all generalized salamanders 1. Premaxillary fusion. Premaxillary bones have five toes (0). are paired in primitive tetrapods and in 8. Lung reduction. Well developed lungs primitive salamanders. Two premaxillaries are primitively present in salamanders (0). are considered primitive (0) and fused pre- Lung reduction and loss is considered to be maxillaries derived (1). derived (1; see Liidike, 1955). 2. Frontosquamosal arch. Frontosquamosal 9. Skin 'texture. It is not possible to ascer- arches are found only in the "family Sala- tain whether or not rough skin such as mandridae. There are many fossil sala- that encountered in the terrestrial newts is mandrids, but most are represented only by primitive or derived. However, since it seems vertebral fragments. Some are known from to be associated with a primitive way?:of complete skulls and the earliest of these life and is absent only in those groups that (Tylototriton weigelti, Middle Eocene; Paleo- have diverged from that pattern, we con- taricha oligocenia, Upper Oligocene; Oligo- sider roughened, keratinized skin to be semia spinosa, Oligocene; Heliarchon fur- primitive (0) and smooth skin to be derived cillatus, Lower Miocene) have well developed in one direction in terrestrial environments arches. The general trend is for bone re- (1) and in another in aquatic habitats (2). duction in salamanders, and the arches are 10. Egg size. Primitively salamanders lay undergoing apparent reduction in certain large numbers of small eggs. Such a pattern genera (e.g., Triturus). However, fronto- is considered primitive (0), and the laying squamosal arches
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