4 John E. Cooper and Ray E. Ashton, Jr. 



mens led Bishop (1947:34) to suggest "an early-spring mating season for 

 this species, although some males of maculosus , which has a fall mating 

 season, are known to retain the swollen glands until spring." 



Although Schmidt (1953) retained the trinomial, Hecht (1953) 

 accepted Viosca's (1937) taxonomic change and placed N lewisi and N 

 beyeri in a Necturus lewisi superspecies group. Both species differed 

 from their congeners in having non-striped larvae and spotted medium- 

 sized adults. The N lewisi superspecies was considered intermediate 

 between the species N. maculosus and N punctatus. Hecht 's series con- 

 tained only 20 adult N. lewisi, so he could not address ontogenetic 

 changes in dentition, body proportions, and other features. He did note 

 a maximum SVL of at least 175 mm, a minimum breeding size between 

 100 and 105 mm SVL, and a change to adult pattern at 130 mm SVL. 

 Hecht (1958) opined that the species of Necturus appeared to be cold- 

 adapted salamanders, active only in the colder seasons and inactive dur- 

 ing hottest months. He further speculated that maximum and minimum 

 breeding size may be an adaptation to thermal regimes of the habitat, 

 concluding (p. 115): "natural selection has resulted in the adaptation of 

 the southern species to higher temperatures and a higher metabolism by 

 reduction of the minimum breeding and maximum size of the species." 

 He considered the lewisi group the most primitive in the genus, with N. 

 punctatus an early derivative of the proto-lewisi ancestor, and TV. macu- 

 losus a direct and recent (advanced) descendent of the lewisi group. He 

 stated that the striped larva of N maculosus is more specialized than 

 the primitive unstriped larval type of the lewisi and punctatus groups. 

 (See Ashton and Braswell, 1979, for discussion of the striped post- 

 hatchling larva of N. lewisi; also see Sessions and Wiley, this issue, for 

 chromosome evolution in Necturus.) 



Blair et al. (1968) included lewisi as a full species. Neill (1963:173), 

 defending species status for N. alabamensis Viosca, said that N. lewisi 

 "most resembles, and is probably most nearly allied to, N. beyeri (sensu 

 Viosca) even though the two inhabit well-separated portions of the 

 Coastal Plain. A distribution of this kind, in a group as ancient and 

 conservative as the waterdogs, suggests that lewisi and beyeri had a 

 common ancestor in the lowlands that bordered the shoreline of the old 

 Cretaceous Embayment. As the shoreline retreated southward, exposing 

 what is now the Coastal Plain, the range of the lewisi-beyeri animal was 

 fragmented." Brode (1970) revised the genus Necturus, using osteologi- 

 cal criteria to relegate N. lewisi to subspecies status under N. maculosus, 

 but this arrangement was not very widely employed. 



Fedak (1971), in the most thorough life history study of North 

 Carolina Necturus to that time, provided information on more than 600 

 Necturus (punctatus and lewisi) from 32 North Carolina localities, all 

 collected from the fall of 1966 through the summer of 1969. Of these, 



