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1 Society for the Study of Amphibians and Reptiles Male Secondary Sexual Characters of the Eurycea bislineata (Amphibia, Urodela, Plethodontidae) Complex in the Sourthern Appalachian Mountains Author(s): David M. Sever Source: Journal of Herpetology, Vol. 13, No. 3 (Jul. 30, 1979), pp Published by: Society for the Study of Amphibians and Reptiles Stable URL: Accessed: :42 UTC REFERENCES Linked references are available on JSTOR for this article: You may need to log in to JSTOR to access the linked references. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at 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 support@jstor.org. Society for the Study of Amphibians and Reptiles is collaborating with JSTOR to digitize, preserve and extend access to Journal of Herpetology.

2 1979 JOURNAL OF Ht-HtIOLOGY 13(3): Male Secondary Sexual Characters of the Eurycea bislineata (Amphibia, Urodela, Plethodontidae) Complex in the Sourthem Appalachian Mountains David M. Sever Department of Biology, Saint Mary's College, Notre Dame, Indiana 46556, USA ABSTRACT-Polymorphism of male secondary sexual characters occurs among populations nominally assignable to Eurycea bislineata in the southern Appalachian Mountains. Two male morphs are recognized, one identified as E. b. wilderae and the other designated morph A. Male E. b. wilderae possess mental hedonic glands, seasonally enlarged premaxillary teeth, cirri and reduced temporal musculature. Male morph A lack mental hedonic glands, seasonally enlarged premaxillary teeth and cirri and possess strongly developed temporal musculature. * * * INTRODUCTION In the southern Appalachian Mountains there are two morphs nominally assignable to Eurycea bislineata (Green) that show extreme differences in their male secondary sexual characters. One of these morphs agrees with descriptions and type material of E. b. wilderae Dunn while the other is referred to herein as morph A. Male secondary sexual characters and several other characters will be contrasted between the two forms, and comparisons will be made with E. junaluska, E. aquatica and E. bislineata from elsewhere in its range. MATERIALS AND METHODS Specimens collected by myself were initially preserved in 10% formalin. For light microscopy, tissue specimens were dehydrated in ethanol, decalcified with Jenkin's solution (when appropriate) and embedded in paraffin (Humason, 1972:46). Sections were cut at 10,m and usually stained in Mallory's triple stain although all testes were stained in hematoxylin-eosin. Testes were removed and prepared histologically from all specimens from which other tissue was examined by light microscopy. Collection sites in the southern Appalachians are shown in Fig. 1 and listed below. Except for two specimens of E junaluska in my own collection, all specimens from these localities are deposited in major museums, and collection numbers are listed in Appendix I. Typical specimens of male E. b. wilderae, morph A and male E. junaluska from these localities are shown in Fig. 2. Locality 1: Road Prong, Indian Gap, Great Smoky Mts., Sevier Co., Tenn., 1600 m elev. Locality 2: Fighting Crk., Sugarlands Park Hdqs., Great Smoky Mts., Sevier Co., Tenn., m elev. Locality 3: Mid's Branch, Elkmont, Great Smoky Mts., Sevier Co., Tenn., 687 m elev. Locality 4: Rush Branch of Little R., 1.2 km SE Townson, Great Smoky Mts., Blount Co., Tenn., m elev. Locality 5: Abrams Crk. and tribs., Cades Cove, Great Smoky Mts., Blount Co., Tenn., m elev. Locality 6: Calderwood Dam, Blount Co., Tenn. Locality 7: Cheoah R., km SE Tapoco, Graham Co., N. Car., m elev. Locality 8: Straton Bald, Graham Co., N. Car., 1600 m elev. Locality 9: Rattler's Ford, Santeetlah Crk. at Co. Rt. 1127, Graham Co., N. 245

3 246 DAVID M. SEVER YTENN R PIGEON R - L T ENN R, A 1 d G SMOKY B 'd UNI 6 CHEOAH,N /,, ~ ' ( I)~ COWE E FIGURE 2. Typical appearance of male E. b. wilderae, morph EN SNOWBIRD TENN R A and E. junaluska in the southern Appalachian Mountains. A. NC SOWBIRD Morph A (MVZ , 44.8 mm SVL) collected 23 October, TAHALA ) 1977 at locality 1. B. E. junaluska (DMS 3367, 45.3 mm SVL) NANTAHALA collected 25 October, 1977 at locality 7. C. E b. wilderae (USNM , 37.1 mm SVL) collected 27 October, 1977 at 10km IY? / L locality 11. FIGURE 1. Study localities for male E. b. wilderae (circle with line at 2 o'clock), morph A (circle with line at 10 o'clock) and Car., m elev. Locality 10: Snowbird E. junaluska (circle with line at 6 o'clock) in the southern Crk. at Co. Rt. 1115, Graham Co., N. Car., Appalachian Mountains. Locality 7 is the type locality of E. junaluska. Major mountain ranges and some major rivers (R) 670 m elev. Locality 11: Panther Crk. Teyahale are indicated. Bald, Graham Co., N. Car., m elev. Locality 12: Tululah Crk., km SE Robbinsville, Graham Co., N. Car., m elev. Locality 13: km N Shooting Creek (Town), Clay Co., N. Car. Locality 14: vicinity of Kyle and Aquone Macon Co., N. Car. Locality 15: Wayah Bald and Jarrett Crk. area, Macon Co., N. Car. Locality 16: 10.5 km NE Cullasaja, Macon Co., N. Car. Only E. b. wilderae was collected at localities 3, 11 and 15, only morph A at localities 6, 8, 10, 13 and 16, and only E. junaluska at locality 7. All three forms were sympatric at localities 2 and 12, morph A and E junaluska were sympatric at locality 9, and E. b. wilderae and morph A were sympatric at localities 1, 4, 5 and 14 (Fig. 1, Appendix I). In all, 122 male E. b. wilderae were examined from March, October, November and December collections, and 114 morph A were examined from March, May, July, August, September, October, November and December collections. All specimens referred to morph A are males. Besides the type series of 23 E. junaluska (11 d :12 Y) collected in May, July and August from localities 7, 9 and 12, I1 examined additional single male specimens collected in March at localities 2 and 9 and in October at locality 7. The mental hedonic gland region was studied in all male E. b. wilderae and morph A from these localities by gross dissection or light microscopy. In all, 111 male E b. wilderae and 100 morph A were dissected. Sections through the lower jaw were made on 11 male E. b. wilderae from localities 1, 2, 5 and 11 including three specimens from locality 5 from both March and October samples. Lower jaws of 14 morph A from localities 1, 2, 8 and 9 were sectioned including three specimens from locality 9 from both March and October samples and two from a July collection. The lower jaw was sectioned in single specimens of male E., junaluska from localities 2, 7 and 9. All male specimens from the October samples were examined for penetration of the premaxillary teeth through the upper lip anterior to the apex of the lower jaw as described by Noble (1931:123) and Arnold (1977:155). Gross dissections of temporal musculature were made on 16 specimens of E. b. wilderae and 31 of morph A from all localities combined and on one male E. junaluska from locality 7.

4 SECONDARY SEXUAL CHARACTERS OF EURYCEA 247 Anatomy of the temporal musculature and number of costal grooves between toes of the adpressed limbs were examined in all females collected with either male E. b. wilderae or morph A from the 16 localities. Altogether 126 male specimens of E. bislineata were examined from other parts of its range (Fig. 3, Appendix II). Included among these were types of E. b. wilderae and topotypes of E. b. bislineata and E. b. cirrigera. The mental hedonic gland region was dissected or sectioned in all specimens. Slides resulting from histological preparation of museum material are on deposit with the specimen. The mental hedonic gland region was also examined in males of the type series of E aquatica from Jefferson Co., Alabama. Temporal musculature was dissected in samples of male E b. bislineata in breeding condition from Hocking Co., Ohio (N = 16), Carter Co., Kentucky (N = 14) and Monongalia Co., West Virginia (N = 13). RESULTS Male Reproductive Cycle.-Spermatozoa occur in all vasa deferentia examined from male E b. wilderae, morph A and male E. junaluska collected from October through March. Testes from March specimens are largely or completely filled with spermatogonia. Morph A from July still have spermatogonia in the anterior tip of the testes followed by primary and secondary spermatocytes in the middle portion and spermatids and spermatozoa in the posterior tip. October E. b. wilderae and morph A have spermatozoa in the anterior portion of the testes and spermatogonia in the posterior end. Cirri.-No specimen of morph A has measurable cirri. A male E. junaluska from October possesses cirri.4 mm long. Male E. junaluska from March have short "bumps" at the ventral border of the nasolabial groove. Males of the type series of E. junaluska from May, July and August lack cirri (Sever et al., 1976:28). All male E. b. wilderae examined from the 16 localities possess cirri. Cirri seem best developed in October samples. Cirri of 67 males ( mm SVL, x = 36.8 mm, SD = 2.22) from October are mm long (x =.92 mm, SD =.14), and cirrus length is positively correlated with SVL (r =.45, 95% C. L. = ). Mental Hedonic Glands.-All male E. b. wilderae examined from the 16 localities possess a large and distinctive mental hedonic gland cluster (Figs. 4A, 5). These clusters agree with the description for the species given by Sever (1976: ). Pores occur ventral and immediately posterior to the mandibular symphysis. The distal portions of the glands pass posteriorly in a thin sheath of dermal FIGURE 3. Localities from which Eurycea aquatica (small open circle) and E bislineata (stars and closed circles) were examined outside the area shown in Fig. 1 (large open circle). Open star = topotypes of E. b. bislineata, open star in circle = type locality for E. b. wilderae and closed star = topotypes of E. b. cirrigera. Not shown is a locality 32 km N Quebec, Quebec Province, Canada. A B FIGURE 4. Gross appearance of mental glands. A. E. b. wilderae (USNM , 38.3mm SVL) collected 27 October, 1977 at locality 11. B. Morph A (AMNH 99653, 40.4 mm SVL) collected 28 October, 1977 at locality 8. Of 114 specimens of morph A, this is the only one with enlarged dermal glands in the chin region. MG = mental gland, P = secretory pores, and PM = premaxillary teeth. a

5 248 DAVID M. SEVER B FIGURE 5. Transverse sections through the lower jaw posterior to the mandibular symphysis in male E. b. wilderae. Scale in lower right hand corner = 250 Lm. A. MCZ 5843, 33.0 mm SVL, collected 9-11 July, 1919 at Grayson Co., Virginia. B. USNM , 36.0 mm SVL, collected 23 October, 1977 at locality 1. C. USNM , 36.0 mm SVL, collected 23 October, 1977 at locality 2. D. USNM , 40.4 mm SVL, collected 27 October, 1977 at locality 11. E. USNM , 41.5 mm SVL, collected 14 March, 1977 at locality 5. F. USNM , 37.2 mm SVL, collected 24 October, 1977 at locality 5. GH = M. geniohyoideus, IM = M. intermandibularis posterior, MG = mental glands, and SC = dermal stratum compactum. stratum compactum between layers of the M. intermandibularis posterior ventrally and the M. geniohyoideus dorsally. Such a mental hedonic gland was also found in all 126 male E. bislineata examined from other parts of its range including types or topotypes of all subspecies. As shown by Weichert (1945), the mental hedonic gland of E. bislineata is most highly developed during the breeding season. However, it is easily recognizable by gross dissection at other times of the year.

6 SECONDARY SEXUAL CHARACTERS OF EURYCEA 249 FIGURE 6. Transverse sections through the lower jaw posterior to the mandibular symphysis in male E. junaluska. Scale in lower right hand corner = 250 um. A. MVZ , 39.0 mm SVL, collected 6 May, 1974 at locality 7. B. DMS 2932, 43.8 mm SVL, collected 15 March, 1977 at locality 2. Abbreviationsame as Fig. 5. The mental hedonic gland of E junaluska (Fig. 6) does not differ at the light microscopy level from that of E. bislineata. Examination of E. aquatica also revealed a mental hedonic gland similar to that of E. bislineata. All but one specimen of morph A lack any trace of a mental hedonic gland cluster (Fig. 7). Ordinary mucous glands are the only type of gland in the lower jaw. The one exception, AMNH from locality 8, has a small cluster of 9 glands at the anterior apex of the lower jaw ventral to the mandible (Fig. 4B). The gland cluster is.75 by.85 mm and totally encased in the dermis ventral to the throat musculature. Premaxillary Teeth.-October samples of male E. b. wilderae from the 16 localities possess 1-8 (mode 4) premaxillary teeth piercing the upper lip anterior to the apex of the lower jaw (Fig. 4A). Such elongate premaxillary teeth are absent in October samples of morph A. A male E junaluska from October has 3 premaxillary teeth protruding like those of E. b. wilderae. Temporal Musculature.-During the breeding season, morph A has a relatively wider head posterior to the eyes than male E. b. wilderae. This difference in head width is related to anatomy of the M. levator mandibulae anterio profundus and M.. m. externus (Fig. 8). In male E b. wilderae the M. I. m. anterior profundus is relatively small, and muscle fibers extend dorsally only to the medial border of the eyelids (Fig. 8A). When the skin is removed from the dorsum of the head, the posterior frontals and anterior parietals are clearly visible in male E. b. wilderae. In morph A muscle fibers of the M.. m. anterior profundus extend to the medial sutures of the posterior frontals and anterior parietals, and the muscles are relatively thick (Fig. 8B). The M. I. m. extemus is similar in position in male E. b. wilderae and morph A, but it is relatively thicker in morph A during the breeding season. The sexually active male E. b. bislineata examined from Ohio, Kentucky and West Virginia possess temporal musculature similar to morph A. These E. b. bislineata all possess mental hedonic glands and relatively enlarged premaxillary teeth but lack cirri. The one male E. junaluska dissected also has temporal musculature like morph A. Other Characters.-Samples are too small and over too limited of a size range to fully analyze traditional metric and meristic characters. However, initial examination of data for some characters shows trends that warrant further investigation. Data for E. junaluska are from Sever et al. (1976). Tail length is generally about 50% of the total length in E. junaluska while it is usually 55-60% of the total length in morph A and male E. b. wilderae. In E junaluska and male E b. wilderae there are 0-2 costal grooves between toes of the adpressed limbs while in morph A there is most commonly 2-3. All male E b. wilderae (N = 39) and morph A (N = 41) examined have 7-22 vomerine teeth with means for various samples ranging from while Sever et al. (1976:28) reported that E junaluska has vomerine teeth, x = 20.6.

7 250 DAVID M. SEVER I0 FIGURE 7. Transverse sections through the lower jaw posterior to the mandibular symphysis in morph A. Scale in lower right hand comer = 250 jim. A. MVZ , 40.2 mm SVL, collected 23 October, 1977 at locality 1. B. USNM , 39.9 mm SVL, collected 17 March, 1977 at locality 9. C. USNM , 39.9 mm SVL, collected 23 October, 1977 at locality 2. D. USNM , 42.6 mm SVL, collected 23 July, 1975 at locality 9. E. AMNH 99654, 39.2 mm SVL, collected 28 October, 1977 at locality 8. F. USNM , 40.4 mm SVL, collected October, 1977 at locality 9. Abbreviations same as Fig. 5, except that DG = dermal mucous glands. Samples of morph A from localities 1 and 8 at 1600 m elevation have a mode of 15 costal grooves while samples from lower altitudes usually have 14 costal grooves. Most male E. b. wilderae examined have 14 costal grooves, but large samples were not examined from altitudes greater than 1350 m. E. junaluska has been collected only below 700 m. elevation, and all specimens examined possess 14 costal grooves (Sever et al., 1976:28).

8 SECONDARY SEXUAL CHARACTERS OF EURYCEA 251 Coloration cannot distinguish male E. b. wilderae and morph A. Both are usually some shade of orange, yellow or brown with black dorsolateral stripes passing from the eyes onto the tail. There is an extreme amount of variation in both forms in concentration and distribution of melanophores. Females associated with either morph are similar in coloration to males. E junaluska is easily distinguished by color pattern from both male E. b. wilderae and morph A. E. junaluska has a brownish mottled dorsum and lacks dorsolateral stripes. Habitat and Distribution.-There are no gross differences in habitat between male E. b. wilderae and morph A. Both are most common during the breeding season and are found under rocks, logs and leaves in and along seepages, springs, and small and large streams. Where they are sympatric, one form usually seems more common than the other. On 23 October 1977, morph A was more common than male E. b. wilderae at locality 1 (1600 m elevation), but male E. b. wilderae was more common at locality 2 ( m). Altitude does not seem to be critical however. Morph A is the only one of the two forms I have collected at localities 8 (1600 m), 9 ( m) and 10 (670 m) whereas male E. b. wilderae is the only one I have seen at locality 3 (670 m) and locality 11 ( m). Both are found at locality 12 ( m) with morph A being most common. At locality 4 ( m), only male E. b. wilderae was collected in March, 1977 and only morph A in October, E. junaluska is restricted to base-level streams below 700 m elevation. E. junaluska is always uncommon when sympatric with male E. b. wilderae or morph A. The only locale where E. junaluska is common is along the Cheoah River in Graham Co., North Carolina (locality 7), and neither male E. b. wilderae nor morph A have been collected there. Females.-Females are commonly collected with both male E b. wilderae and morph A, but I have found no morphological character that consistently assigns females to one form or the other. Despite locality, most females possess temporal musculature similar to male E. b. wilderae. The jaw adductors of females are not hypertrophieduring the breeding season, and fibers of right and left M. I. m. anterior profundus usually do not meet in the dorsal midline. Females generally have 2-4 costal grooves between toes of the adpressed limbs which is similar to morph A. Most females collected in March and October are swollen with eggs, and this may bias this character. Male and female E. junaluska are similar in coloration and overall body proportions. Coloration is the most obvious difference between female E. junaluska and females associated with male E. b. wilderae and morph A, but available data, as for males, also indicates significant differences between female E. junaluska and females of the other forms at least in relative tail length and number of vomerine teeth. LMAP DISCUSSION LMAS Morph A can be distinguished from male E. b. wilderae in the southern Appalachians by lack of mental hedonic glands, relatively elon- A B gate premaxillary teeth and cirri and by pre- FIGURE 8. Gross appearance of male temporal musculature. sence of strongly developed temporal mus- A. E b. wilderae (USNM , 37.1 mm SVL) collected 27 culature and 2-3 versus 0-2 costal grooves October, 1977 at locality 11. B. Morph A (USNM , between toes of the adpressed limbs. Morph A 38.5 mm SVL) collected October, 1977 at locality 9. can be distinguished from male E. junaluska LMAP = M. levator mandibulae anterior profundus, LMAS = again m by l lackg h c g, r- M. I. m. a. superficialis, and LME = M. I. m. exterus. again by lacking mental hedonic glands, relatively enlarged premaxillary teeth and cirri and by number of costal grooves between toes of the adpressed limbs; furthermore, the two forms differ in coloration and probably by relative tail length and mean number of vomerine teeth. In its original description, E. junaluska was compared to samples from Graham Co., North Carolina assigned to E. bislineata (Sever et al., 1976). Most males of these specimens are referrable to morph A.

9 252 DAVID M. SEVER E. junaluska differs from male E. b. wilderae by lesser development of cirri, presence of strongly developed temporal musculature, coloration and apparently in relative tail length and number of vomerine teeth. The taxonomic status of morph A is uncertain. Mental hedonic glands were present in all male specimens assignable to E. bislineata from other parts of its range including types or topotypes of E b. bislineata, E. b. cirrigera and E. b. wilderae. Mental hedonic glands similar to that of E. bislineata were found in male E. junaluska and male E. aquatica. Mental hedonic glands are also known from all other non-neotenic species of Eurycea (Sever, 1976) except possibly E neotenes, individuals of which were recently found to complete metamorphosis (Sweet, 1977). To my knowledge, E neotenes and the neotenic Eurycea have not been examined for mental hedonic glands. Within the rest of the Hemidactyliini, only Pseudotriton and Gyrinophilus are known to lack mental hedonic glands (Sever, 1976:229), but Stereochilus and neotenic Typhlomolge and Haideotriton have not been examined for such glands. The mental hedonic glands apparently play a "priming" role in courtship (Arnold, 1977:159), and their absence would seem highly significant. Both Gyrinophilus and Pseudotriton are considered primitive genera (Wake, 1966:62), and the courtship pattern of Pseudotriton was described by Organ and Organ (1968:222) as almost schematic in its simplicity. My results support Arnold's (1977: ) findings that the presence of relatively enlarged premaxillary teeth is correlated with possession of a mental hedonic gland that secretes at the anterior apex of the lower jaw. The projecting teeth scrap the female's skin during courtship allowing secretions of the male's mental hedonic glands to enter the superficial circulation of the female (Arnold, 1977:156). Cirri occur in populations assigned to E. bislineata from the Coastal Plain, Piedmont and southern Appalachian Highlands (Sever, 1972:318). Elsewhere males of the species may have short blunt protuberances at the base of the nasolabial groove, but elongate cirri are lacking. My findings suggest that temporal muscle hypertrophy in male E. bislineata occurs only in those populations that lack cirri. Morph A either represents an extreme case of polymorphism in male E. b. wilderae, or morph A is a separate species. I believe the evidence so far leads to the latter conclusion. Further work is in progress. ACKNOWLEDGMENTS This study was supported by a grant from the Johnson Fund of the American Philosophical Society. I thank A. L. Braswell, E. E. Williams, G. R. Zug and R. G. Zweifel for loan of specimens in their care. I thank J. H. Caruso and C. D. Sullivan for supplying me with some specimens from Pennsylvania and Tennessee respectively. I thank R. C. Bruce for aid in the field during October, APPENDIX I Specimens from the 16 localities shown in Fig. 1 are listed below except for the type series of E. junaluska which are listed by Sever et al. (1976:26): Locality 1: E. b. wilderae (USNM ); morph A (MVZ ). Locality 2: E. b. wilderae (USNM ); morph A (USNM ); E junaluska (DMS 2932). Locality 3: E b. wilderae (USNM ) Locality 4: E. b. wilderae (USNM ); morph A (USNM ). Locality 5: E. b. wilderae (USNM ); morph A (USNM ). Locality 6: morph A (USNM ). Locality 7: E. junaluska (DMS 3367). Locality 8: morph A (AMNH ). Locality 9: morph A (USNM ); E junaluska (DMS 2341). Locality 10: morph A (MCZ ). Locality 11: E. b. wilderae (USNM ). Locality 12: E. b. wilderae (NCSM 14497, USNM ); morph A (NCSM 14130, , , , , , ). Locality 13: morph A (NCSM 14487, 14504). Locality 14: E. b. wilderae (NCSM , , , , , ); morph A (NCSM 14492, 14818, ). Locality 15: E. b. wilderae (NCSM 14491, USNM ). Locality 16: morph A (NCSM ).

10 SECONDARY SEXUAL CHARACTERS OF EURYCEA 253 APPENDIX II Specimens examined from localities shown in Fig. 3: Eurycea aquatica. Alabama: 3.2 km W Bessemer, Jefferson Co. (USNM ). E bislineata. CANADA. Quebec: Stoneham Prov. Pk., 32 km N Quebec (DMS 2675). USA. Alabama: 19 km E Abbeville, Henry Co. (DMS 1315). Florida: 2.4 km SE Munson, Santa Rosa Co. (DMS ). Indiana: Shades St. Pk., Montgomery Co. (DMS ); McCormick's Crk. St. Pk., Owens Co. (DMS , ). Kentucky: Carter Caves St. Pk., Carter Co. (DMS ). Louisiana: Talisheek, St. Tammany Par. (DMS 781); 4.8 km NWN Robert, Tangipahoa Par. (DMS 1654). Mississippi: Rocky Springs, Franklin Co. (DMS 873). New Jersey: Clouster, Bergen Co. (AMNH ); Fort Lee, Bergen Co. (USNM 82428, 82432, ); Englewood, Bergen Co. (USNM , 93643); Leonia, Mercer Co. (AMNH ); Hightstown, Mercer Co. (USNM ). North Carolina: Bearpen Gap, Jackson Co. (DMS ); Highlands, Macon Co. (DMS , ); Wilmington, New Hanover Co. (NCSM ); Hoffman, Richmond Co. (DMS 2226); 5.6 km N New Salem, Union Co. (DMS ). Ohio: Dyssart Woods, Belmont Co. (DMS ); Conkle's Hollow, Hocking Co. (DMS ); Darwin, Meigs Co. (DMS ). Pennsylvania: Lake Warren, Bucks Co. (DMS 3390). Tennessee: Genesis Rd., Cumberland/Morgan Co. line (DMS ); 21.7 km S Waverly, Humphrey's Co. (DMS 611). West Virginia: Copper's Rock St. Pk., Monongalia Co. (DMS , ). LITERATURE CITED Arnold, S. J The evolution of courtship behavior in new world salamanders with some comments on old world salamanders. In The reproductive biology of amphibians. D. H. Taylor and S. I. Guttman, eds. Plenum Press, New York. 475 pp. Humason, G. L Animal tissue techniques. W. H. Freeman and Co., San Francisco. 641 pp. Noble, G. K The biology of the Amphibia. McGraw-Hill Book Co., New York. 577 pp. Organ, J. A. and D. J. Organ Courtship behavior of the red salamander, Pseudotriton ruber. Copeia 1968: Sever, D. M Geographic variation and taxonomy of Eurycea bislineata (Caudata: Plethodontidae) in the upper Ohio River valley. Herpetologica 28: Morphology of the mental hedonic gland clusters of plethodontid salamanders (Amphibia, Urodela, Plethodontidae). J. Herp. 10: , Dundee, H. A. and C. D. Sullivan A new Eurycea (Amphibia: Plethodontidae) from southwestern North Carolina. Herpetologica 32: Sweet, S. S Natural metamorphosis in Eurycea neotenes, and the generic allocation of the Texas Eurycea (Amphibia: Plethodontidae). Herpetologica 33: Wake, D. B Comparative osteology and evolution of the lungless salamanders, family Plethodontidae. Mem. S. California Acad. Sci. 4: Weichert, C. K Seasonal variation in the mental gland and reproductive organs of the male Eurycea bislineata. Copeia 1945: Accepted 5 Apr 1979 Copyright 1979 Society for the Study of Amphibians and Reptiles 0