N. lewisi Habitats and Behavior 105 



to cause little damage to the intruder. Attacks on N. punctatus, how- 

 ever, were more direct and resulted in serious wounds or death. 



Parzefall (1980) reported that N. maculosus and Proteus anguinus 

 responded to olfactory cues in water and those left on substrates. Nectu- 

 rus maculosus recognized and avoided retreat areas occupied by Pro- 

 teus. Necturus lewisi displayed similar responses to retreats that were 

 not physically occupied by the resident. This indicates that similar olfac- 

 tory cues may be involved in identifying territory. Adult N. lewisi rec- 

 ognized larvae and juveniles and did not display territorial aggression 

 nor did they attempt to eat them. 



Necturus lewisi was much more aggressive than the syntopic N. 

 punctatus, which never challenged intruding N. lewisi. Captive N. punc- 

 tatus were displaced by N. lewisi. This may indicate an amensalistic 

 relationship in the wild. 



Courtship in N. lewisi was similar to that reported for N. maculo- 

 sus. Bishop (1941) observed that the courting male repeatedly crossed 

 over the female, which became motionless with head erect, but he did 

 not report chin rubbing or trailing as was observed in N. lewisi. These 

 may indicate that pheromones, as well as the tactile senses, play an 

 important sexual role. The actual transfer of spermatophores from male 

 to female is still unobserved in Necturus. 



ACKNOWLEDGMENTS. — I express my appreciation to Keith 

 Everett and Ernie Flowers for their many long, cold, uncomfortable 

 hours in the field. I also thank Jesse Perry, Patricia Ashton, Dan Smith, 

 Paul Kumhyr, and the many others who assisted in the field. I thank, 

 too, Drs. John C. Clamp and John E. Cooper for identifying inverte- 

 brates, and further thank Dr. Cooper for the advice and assistance he 

 provided in all phases of the Necturus study. This study was partly 

 funded by a grant from the Carolina Conservationist Program, and con- 

 tract funds from a U.S. Fish and Wildlife Service (Office of Endangered 

 Species) cooperative agreement, both provided by the North Carolina 

 Wildlife Resources Commission. Other support was provided by the 

 North Carolina State Museum, a division of the N.C. Department of 

 Agriculture. Renaldo G. Kuhler, state museum scientific illustrator, did 

 the figures. 



LITERATURE CITED 



Ashton, Ray E., Jr. 1975. A study of movement, home ranges and winter behav- 

 ior of Desmognathus fuscus (Rafinesque). J. Herpetol. 9:85-91. 



Ashton, Ray E., Jr., and A. L. Braswell. 1979. Nest and larvae of the Neuse 

 River Waterdog, Necturus lewisi (Brimley) (Amphibia: Proteidae). Brim- 

 leyana 1:15-22. 



