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We describe a new species of plethodontid salamander of the genus Pseudoeurycea from the mountains of central Veracruz, Mexico. This species can be distinguished from its congeners by colo...

We describe a new species of large black Bolitoglossa from the Sierra de las Minas and adjacent regions of Guatemala based on both morphological and molecular data. The new species, specimens of which were formerly assigned to Bolitoglossa meliana (Melanistic Salamander), is distinguished by its dark coloration, numerous teeth, and large size. We present molecular data to support the distinctiveness of this species and estimate its phylogenetic placement within the subgenus Magnadigita. Molecular data do not support a sister taxon relationship between the new species and B. meliana, despite their outward morphological similarity. This species adds to the already high salamander species diversity of the Sierra de las Minas, much of which has been described recently.

Some prey species vary the intensity of antipredator responses according to the perceived level of threat associated with different concentrations of chemical cues related to predation. Here, we examine whether Red-Backed Salamanders (Plethodon cinereus) respond to different concentrations of damage-released cues from the tails of conspecifics in a threat-sensitive manner. We exposed salamanders either to a control or to damage-released cues from conspecifics at one of three different concentrations. We found that salamanders exposed to damage-released cues significantly decreased their activity compared to salamanders exposed to a control. However, the intensity of the responses was not related to the concentration of the cue, suggesting that individuals of P. cinereus do not respond in a threat-sensitive fashion at the concentrations tested.

Almost nothing is known about the biology of the recently described, enigmatic Lao Newt (Laotriton laoensis). Using a combination of field surveys and village interviews, we determined that its geographic range is restricted to a small area in northern Laos. Aquatic adults occur in pools of small, cold, slightly acidic, flowing streams that traverse a variety of vegetation types at elevations above 1,100 m. Mark–recapture estimated that aquatic adults are locally abundant where they occur. Analysis of stomach contents found that the species consumes a wide variety of invertebrates and vertebrates, including large prey, and females eat conspecific eggs. Aquatic courtship takes place during the coldest and driest part of the year, and eggs are laid over a long period between dead leaves that accumulate on the bottom of stream pools. Larvae are aquatic and metamorphose into terrestrial efts. The species is threatened from overharvesting for food, medicine, and especially the international pet trade.

Bolitoglossa nicefori is a terrestrial salamander only known from its original description. We report the discovery of two populations of this species; analyze annual reproductive activity, diet, and microhabitat use of one population; and compare this information with that known for other species of Bolitoglossa. Sexually mature males ranged between 29 and 50 mm standard length and females between 39 and 75 mm. Histological analyses of testes and ducts revealed the presence of sperm through the year, indicating continuous reproductive activity for males. Adult females were captured year-round; however, reproductive females were observed only during the driest months of the year (November to February), indicating that females have a seasonal reproductive activity. The contents of 89 stomachs were analyzed, and perch height, type of vegetation, and substrate type were registered. The diet of this population consists of 13 items, with ants and coleopterans accounting for 87% of food ingested by the...

Effective monitoring of population size is critically important for endemic species with specialized habitat requirements so that timely remedial steps can be taken when declines are detected. We initiated a monitoring study of the endemic plethodontid salamander, Plethodon punctatus, which is generally found in talus habitats over 1000 m in elevation in a narrow range on Shenandoah Mountain on the border of Virginia and West Virginia. We tested congruence of nighttime visual encounter surveys (VES) and mark-recapture estimates of population size. VES was a valid index of the abundances of P. punctatus in the two habitats we surveyed. Sites on the eastern and western sides of Shenandoah Mountain were surveyed, and both methods estimated that population size on the west was approximately twice as high as that on the east. Individuals of this species exhibited a high degree of site fidelity. Cover object searches for species in talus habitats are expected to be of limited value, and we conclude that nighttime visual encounter surveys are most effective for population size monitoring of P. punctatus and other species that live in talus.

A new terrestrial species of Pseudoeurycea is described from the mountains of the Mixteca region of Oaxaca. Compared with other members of the genus, the new species is relatively slender and short-legged. The background color of the dorsum of this species is dark gray with reddish-brown spots. The Mixteca region of Oaxaca has long been poorly explored biologically, and the knowledge of the distribution of its salamanders is scarce. Herein, we present new information regarding salamanders species in this region.

Body size is an important variable used in life-history and sexual selection theory to predict reproductive, behavioral, and ecological traits. Except for the presence of special skin glands in males, sexual dimorphism has not been reported in the Red Hills Salamander (Phaeognathus hubrichti), the basal member of the Desmognathinae. These data provide insight into the evolution of SSD (sexual size dimorphism) in the entire subfamily. We conducted multivariate and univariate tests on eight morphological measurements of 92 preserved P. hubrichti. We also examined specimens for broken tails and U-shaped scars, which may indicate bites from conspecifics. Male salamanders were larger than females in all measurements except tail length and had more scars than either females or juveniles. This species exhibited male-biased SSD in shape and size: males were broad and bulky, females were long and thin. Regression analysis showed differences in resource allocation between male and female salamanders. Differences in life-history strategies and sexual selection in the form of male-male combat may explain these differences in body size.

A new species of Paramesotriton is described from northern Laos. This represents the first record of Caudata from the country. The species is placed in the genus Paramesotriton based on osteological characters. It differs from all other members of the genus in dorsal color pattern and reduced tongue development.

— The only mudpuppy (Caudata: Proteidae) known to occur in the Tennessee Valley of the Interior Highlands and Southern Appalachians is the Common Mudpuppy (Necturus maculosus). Necturus maculosus is not known to co-occur with any other congeners. Here, we report evidence that an additional Necturus occurs in the Hiwassee River, a tributary of the Tennessee River, in eastern Tennessee. Some specimens from the Hiwassee River are clearly identified as N. maculosus, but others resemble the Neuse River Waterdog (N. lewisi), known from only the Tar-Neuse river system draining to the Atlantic Ocean on the opposite side of the Appalachian Mountains. Concordance between color pattern, mitochondrial DNA, and four nuclear loci demonstrate that these two co-occurring forms represent distinct lineages rather than color variants within a single, panmictic population. A few mismatched genotypes (7 of 32 individuals in total) suggest rare hybridization and backcrossing. Phylogenetic analyses indicate that the ne...

A new species of Oedipina (subgenus Oedopinola) is described from a cloud forest locality in northwestern Honduras. The new species appears most closely related to Oedopinola gephyra, a cloud forest species from north-central Honduras. It differs from O. gephyra in having a larger size, a jet-black coloration, a row of white flecks ventrolaterally on the body in life, and less interdigital webbing on the hind limbs.

The morphology of the bony palate in larval and metamorphosed Epicrionops bicolor Boulenger and E. petersi Taylor (Rhinatrematidae) was studied to assess the extent of palatal change at metamorphosis. In larvae the maxilla is short; it abuts the dorsolateral process of the palatine at mid-choanal level. The pterygoid is long and straight; it runs anteromedially, close to the lateral edge of the parasphenoid. At metamorphosis the maxilla fuses with the palatine. The maxillary part of the maxillopalatine expands dorsally and caudally, surrounding the orbit and lacrimal ducts and completing the lateral border of the subtemporal fenestra. The anterior part of the pterygoid shifts laterally, broadening the interpterygoid vacuity, and separates from the posterior part of the pterygoid. The quadrate develops a rostrally directed quadratojugal process, overlapped by the maxilla and squamosal. A review of the literature shows that a similar pattern of palatal metamorphosis (except for the division of the pterygoid) is seen in other caecilian genera with free-living larvae: Ichthyophis (Ichthyophiidae), Grandisonia (Caeciliaidae), and probably Uraeotyphlus (Uraeotyphlidae) and Sylvacaecilia (Caeciliaidae). This implies that the shared pattern is plesiomorphic for caecilians. Features of palatal metamorphosis shared among caecilians, salamanders, and frogs support the hypothesis of lissamphibian monophyly. rnal of Herpetol gy, Vol. 30, No. 1, pp. 27-39, 1996 yright 19 6 Society for the Study of Amphibians and Reptiles latal Metamorphosis n Basal Caecilians phibia: Gymnophiona) as Evidence for sa phibian Monophyl Extant amphibians comprise three distinct, monophyletic groups: frogs (Anura), salamanders (Caudata), and caecilians (Gymnophiona) (Duellman and Trueb, 1986; Milner, 1988; Cannatella and Hillis, 1993). The relationship of each group to the others, and of these to fossil taxa, has been much debated. However, an emerging consensus-based on both morphological and molecular evidence-supports the hypothesis first explicitly proposed by Parsons and Williams (1962, 1963; see also Gadow, 1901; 1 Present Address: Department of Molecular and Cellular Biology, University of Arizona, Tucson, Arizona 85721, USA. tant amphibians compris three distinct, onophyletic groups: frogs (Anura), salamaners (Caud ta), and caecilians (Gym ophiona) uellman and Trueb, 1986; Milner, 1988; Canatella and Hillis, 1993). The relationship of ch group to the others, and of the e to fossil Parker, 1956; Szarski, 1962): that the extant groups are a monophyletic unit, the Lissamphibia (e.g., Milner, 1988, 1993; Bolt, 1991; Trueb and Cloutier, 1991; Cannatella and Hillis, 1993; Hedges and Maxson, 1993). Since basal members of all three modern groups have a biphasic life cycle with a discrete metamorphosis from the larval to the juvenile form (Duellman and Trueb, 1986), we might expect to find common character transformations retained in the metamorphosis of each group. Such shared character transformations could serve as additional lissamphibian synapomorphies (cf. de Queiroz, 1985). Both frogs and salamanders undergo profound changes in skull structure at metamorphosis, especially in the palate and the hyoer, 1956; Szarski, 1962): hat the extant ps are a monophyletic unit, the Lissami ia (e.g., Milner, 1988, 1993; Bolt, 1991; Trueb 27 This content downloaded from 157.55.39.32 on Mon, 10 Oct 2016 04:24:55 UTC All use subject to http://about.jstor.org/terms

Movements and growth in a population of the terrestrial plethodontid salamander Ensatina eschscholtzii platensis were investigated using mark-recapture methods over a period of 1946 days at a mid-elevation site in the central Sierra Nevada of California. The study site included two plots of equal size, totaling 3.0 ha, in an old-growth pine-fir-incense cedar forest. Our sample included 925 captures, and 14% of the animals were recaptured at least once. Males were more active than females; recapture rates were significantly higher for males than for females, despite a 1:1 sex ratio. Variance in distance traveled was significantly greater for males than for females, and most long-range movements were by males, but mean distance traveled did not differ significantly between sexes. The maximum distance moved for males and females was 120.4 m and 60.6 m, respectively. These movement differences between sexes may explain differences in patterns of mtDNA and allozyme variation within the Ensatina species complex. We suggest that terrestrial plethodontids routinely travel distances > 10 m and caution against calculating home-range sizes from studies conducted only on plots of small size.

A new species of salamander of the genus Bolitoglossa is described from a cloud forest locality in northwestern Honduras. Its combination of strongly sexually dimorphic color patterns, exten- sively webbed feet that lack subdigital pads, and well-developed terminal phalangeal elements with most terminal phalanges expanded distally easily distinguish the new species from all other species of the Bolitoglossa beta assemblage. The new species cannot be confidently placed into any of the recognized species groups of Bolitoglossa, but may be most closely related to B. cuchumatana and B. helmrichi. RESUMEN.-Se describe una especie nueva de salamandra del genero Bolitoglossa de una localidad de bosque nublado en el noroeste de Honduras. Se distingue facilmente de los otros miembros del Bolitoglossa beta por la combinacion de patrones de color fuertemente dimorficos sexualmente, membranas indigitales extensas que faltan cojinetes subdigitales y elementos terminales de los falanges bien desarrollodos con la mayoria de los falanges terminales expandidas distalmente. La especie nueva no puede ser puesta dentro de ningun grupo de especies reconocido des Bolitoglossa, pero quizas es mas cercanamente relacionado a B. cuchumatana y B. helmrichi.

The spermathecae of the salamander Ambystoma tigrinum are simple tubuloalveolar exocrine glands in the dorsal and lateral walls of the cloaca. In laboratory-raised individuals, the glands are slender cords of epithelial cells at 1 yr and production of 160-400 nm apical, electron-dense secretory granules occurs at 1.5 yrs. More extensive synthesis of secretory products (granules 250-450 nm) is evident at sexual maturity, which can occur at 2 yrs. The glycoprotein secretion is synthesized prior to the breeding season and compacted into the electron-dense granules. Secretory material contained in the granules dissociates and is transferred to vacuoles during the breeding season. The vacuoles and their contents are exocytosed into the lumen during oviposition, perhaps serving to flush sperm from the spermathecae. Oviposition follows within two days of mating in A. tigrinum, and this condition is hypothesized as the plesiomorphic state for duration of sperm storage in the spermathecae of salamanders. . , i i il s

tanus), and red efts (Notophthalmus viridescens). Herpetologica 35:289-303. BRAME, A. H., JR., M. C. LONG, AND A. A. CHIRI. 1973. Defensive display of the desert slender salamander, Batrachoseps aridus. Herpeton 8:1-3. BRODIE, E. D., JR. 1970. Western salamanders of the genus Plethodon: systematics and geographic variation. Herpetologica 26:468-516. 1977. Salamander antipredator postures. Copeia 1977:523-535. 1983. Antipredator adaptations of salamanders: evolution and convergence among terrestrial species. In N. S. Margaris, M. Arianoutsou-Faraggitaki, and R. J. Reiter (eds.), Plant, Animal and Microbial Adaptations to Terrestrial Environment, pp. 109-133. Plenum Publ. Corp., New York. , J. A. JOHNSON, AND C. K. DODD, JR. 1970. Immobility as a defensive behavior in salamanders. Herpetologica 30:79-85. BRUNO, S. 1973. Anfibi d'Italia: Caudata. Natura, Soc. It. Sc. Nat. 64:209-450.

MUSHINSKY, H. R., J. J. HEBRARD, AND M. G. WALLEY. 1980. The role of temperature on the behavioral and ecological associations of sympatric water snakes. Copeia 1980:744-754. OTis, D. L., K. P. BURNHAM, G. C. WHITE, AND D. R. ANDERSON. 1978. Statistical inference from capture data on closed animal populations. Wildl. Monogr. 62. 135 pp. PARKER, W. S., AND M. V. PLUMMER. 1987. Population ecology. In R. A. Seigel, J. T. Collins, and S. S. Novak (eds.), Snakes: Ecology and Evolutionary Biology, pp. 253-301. MacMillan Publ. Co., New York.

Many plethodontid salamanders establish feeding territories. We studied interactions of two species in different genera to investigate the roles of species identity, residency, and relative body size in gaining or maintaining control of a cover object. Species identity had the largest effect, with Desmognathus ochrophaeus proving more successful than Plethodon cinereus as a resident and as an intruder. Residency also influenced the outcome of trials: residents were more likely to maintain control of their cover objects than to be displaced. The relative body size of the salamanders in a trial had a small, marginally nonsignificant effect on the outcome. Species identity may be important in determining spatial relations of salamanders in the field. Foraging by plethodontid salamanders is re- stricted by dryness of the habitat. When the forest is wet, plethodontids forage widely and even climb plants in search of prey (Jaeger, 1978). When the forest becomes drier, activity

A study was conducted in 1991 on the population dynamics of the Peaks of Otter salamander, Plethodon hubrichti, a salamander with an extremely limited distribution. From a 10 m x 10 m site in central Virginia, a total of 250 P. hubrichti were collected with 42 recaptures. The population size for P. hubrichti was estimated to be 450 salamanders. There was a slightly clumpedalee y distribution of salamanders throughout the site. The overall growth rate for young of the year was 0.10 mm/d and 0.08 to 0.11 mm/d for older animals. Surface activity occurred primarily between 2100 and 2300 h in the spring, 2200 and 2400 h in the summer, and 2000 and 2300 h in the fall. There was a positive linear relationship between the percentage of surface-active salamanders and the number of days between rains. Salamanders were often observed on vegetation; the percentage of salamanders on vegetation increased in relation to un- derstory denseness. The median area for home ranges was 0.6 m2; with this limited movement, the potential for fragmentation of the population was discussed. An alternative study plot design was considered based upon the clumped distribution and limited movements. The use of three 4 m x 4 m plots instead of one 10 m x 10 m plot reduced the amount of field time by 20% while still obtaining similar information.

The spotted salamander, Ambystoma maculatum, has three types of egg masses; white, clear, and intermediate. White and clear forms are widely-distributed and often sympatric, whereas intermediate forms may be scarce or absent in local populations. Opacity of the egg mass depends on the concentration of hydrophobic protein crystals (1-3 μm in length, 15,400 kD in molecular size) in the outer jelly layers of white and intermediate egg masses. A water soluble protein (14,400 kD) replaces the hydrophobic crystals entirely in outer jelly layers of clear egg masses and partly in intermediate egg masses, and is found in the inner jelly of all three egg mass types. A mitochondrial DNA sequence analysis supports the hypothesis that these variants represent a simple polymorphism of a single gene, rather than the presence of a cryptic species complex

Adult red-spotted newts, Notophthalmus viridescens, tend to be site tenacious within ponds and philopatric to their native ponds. We conducted laboratory experiments to determine the ability of these newts to distinguish between familiar and unfamiliar individuals from the same pond and between individuals from the same versus foreign ponds. We also observed social behavior among newts in a montane pond. In laboratory chambers, newts spent significantly more time moving away from familiar versus unfamiliar individuals from the same pond. They also exhibited significantly more "avoidance" or "escape" behavior when paired with individuals from a foreign pond than when paired with individuals from the same pond. These results demonstrate that red-spotted newts identify individuals with which they are closely associated and distinguish individuals from a foreign pond.

Four species of small, typically red-backed salamanders (Plethodon cinereus, P. dorsalis, P. serratus, P. websteri) occur in the southeastern United States and were formerly included in P. cinereus (Bishop, 1943). Aspects of the life history of P. cinereus, P. serratus, and P. websteri have been studied (Sayler, 1966; Werner, 1971; Nagel, 1977; Lotter, 1978; Semlitsch and West, 1983; Camp, 1988; Taylor et al., 1990). The reproductive biology of southern populations of these species is similar. Sexual maturity is usually attained in the third year of life at a size of about 30-35 mm snout-vent length (SVL); surface activity occurs from fall to spring; oviposition is annual, and each mature female broods 3-15 eggs, presumably during summer in subterranean refuges (Nagel, 1977; Semlitsch and West, 1983; Camp, 1988; Taylor et al., 1990). The purpose of this study was to compare the reproductive biology of P. dorsalis in Arkansas with its morphologically similar congeners. Salamanders were sampled monthly from January to December 1988 by turning logs and rocks in Newton County, Arkansas. However, in June, July, and August, when salamanders were scarce on the surface, all but three individuals were collected in a cave.

Most researchers who have studied effects of low pH on amphibians that breed in temporary ponds measured direct lethal effects on single species. We summarize here results of a study conducted over 4 years in central Pennsylvania that illustrate the potential ecological importance of direct and indirect effects of low pH at lethal and sublethal levels. We argue for use of a multilevel approach (field sampling and experimentation in the laboratory, simulated ponds, and natural ponds) to study such effects on embryonic and larval amphibians of temporary ponds. A greater proportion of our study ponds was lower in pH than any reported previously in North America. Ponds were soft and often had high (Al,t,,,). Median seasonal pHs of 9 ponds were negatively associated with total amounts of rainfall over 4 years, but were not associated with H+ deposition during that same period. The pHs of many ponds declined as total rainfall increased. Embryonic mortality of the salamander Ambystoma jeffersonianum was high in ponds below pH 4.5. Less pH-associated embryonic mortality was observed among the more tolerant salamander, Ambystoma maculatum, and frog, Rana sylvatica. Laboratory-determined LC,, values for hatching of R. sylvatica, A. maculatum, and A. jeffersonianum were 4.10, 4.31, and 4.51, respectively, and predicted embryonic mortalities in ponds with good accuracy. More larval R. sylvatica survived at pH 4.1 in the laboratory when initially contained with A. jeffersonianum than at pH >6. This was due to reduced survival of, and subsequent reduced predation by, A. jeffersonianum at pH 4.1. In the same experiment, survival of A. maculatum was not different between pHs as they were eaten by A. jeffersonianum at pH >6 and suffered pH-induced mortality at pH 4.1. More larval A. jeffersonianum survived at pH 4.1 over 7 days when larval R. sylvatica and A. maculatum were available as prey, than when no prey were available. Larval A. jeffersonianum did not survive to metamorphosis in pH 4.2, and survived in low numbers in pH >6, in simulated ponds. Ambystoma maculatum metamorphosed less often, took longer to metamor- phose, and weighed less at metamorphosis in simulated ponds of pH 4.2 versus those of pH >6. Repro- duction of the newt Notophthalmus viridescens in simulated ponds was significantly lower, and adults were trapped more often exiting ponds, in pH 4.2 than in pH >6. Presence of adult N. viridescens in simulated ponds resulted in significantly lower survival of R. sylvatica at metamorphosis; survival was not significantly affected by pH 4.2. Our results in simulated ponds were likely due to a combination of lethal and sublethal effects that impinged either directly on individual larvae, or indirectly via the food web. These results suggest that studies focused only on direct lethal effects of low pH could substantially underestimate total impacts of low pH on amphibians of temporary ponds. Thus, in addition to the pronounced direct lethal effects we observed on A. jeffersonianum, the chronic and intermittent fluctu- ations in pHs in our study ponds that were associated with total rainfall could conceivably have induced less obvious effects throughout the food webs.

We monitored the demographics of the salamander Ambystoma tigrinum nebulosum as part of a community-wide study on the effects of acidification in sub-alpine (elevation 3600 m) ponds in central Colorado. A decline in A. t. nebulosum at this site from 1982 to 1988 has been hypothesized to result from embryonic mortality during a pulse of acidity that accompanies snowmelt in spring. Since 1988 we have monitored salamander population size, reproduction, and recruitment, and compared survival and individual growth rates among ponds which differ five-fold in acid-neutralizing capacity (ANC)

Allozyme variation was examined in Hynobius sadoensis from Sado Island in the Japan Sea and H. nigrescens from Honshu, Japan. Hynobius sadoensis cannot be distinguished genetically from populations of H. nigrescens, and is thus synonymized with H. nigrescens. A complex pattern of genetic differentiation was found among mainland populations of H. nigrescens. The Sado salamander, Hynobius sadoensis, was described by Sato (1940) from Sado Island in the Japan Sea, off the coast of Niigata in the Chubu District of mainland Honshu. Sato (1940) noted that this species was most similar to H. nigrescens Stejneger, 1907, but differed from it in several characteristics of adult external and internal morphology, and the egg sac envelope. Hynobius nigrescens is the most widely distributed of Japanese Hynobius species; it occurs throughout the Tohoku District and parts of the Kanto and Chubu districts of northeastern Honshu (Iwasawa, 1981). Nakamura and U6no (1963) considered these two species conspecific and synonymized H. sadoensis with H. nigrescens, ado salamander, Hynobius adoensis, was ibed by Sato (1940) from Sado Island in apan Sea, of the coast of Niigata in the District of mainland Honshu. Sato (1940) that this species was most similar to H. escens Stejneger, 1907, but differ d from it 5 Present Address: Tochigi Prefectural Museum, Utsunomiya, Tochigi 320, Japan. without recognizing subspecies. Later, Okada (1970) treated H. sadoensis as a distinct subspecies of H. nigrescens. Both of these taxonomic treatments were relatively superficial, and few subsequent studies have addressed this problem (Iwasawa, 1981; Nambu, 1991). Small salamanders of the genus Hynobius, including H. sadoensis and H. nigrescens, are notoriously difficult to identify based on morphology alone. Recent studies of Japanese Hynobius demonstrate that electrophoresis is particularly useful for clarifying the taxonomic status of populations and systematic relationships of these salamanders (Matsui and Miyazaki, 1984; Matsui, 1987). This study provides electrophoretic evidence that Hynobius sadoensis should be included in the synonymy of H. ni

Relationships among North American plethodontid salamanders have been the subject of numerous molecular studies. In particular, immunological investigations of albumin evolution using the quantitative technique of micro-complement fixation have provided insights into relationships between the genera Ensatina and Plethodon, as well as within Plethodon. However, examination of relationships within eastern Plethodon has been hindered due to lack of antisera from a representative of each species group. An antiserum to albumin from P. wehrlei (wehrlei group) was produced and compared to all available Plethodon antigens. With this antiserum, all currently recognized species groups of eastern Plethodon (glutinosus, cinereus, welleri, and wehrlei) are now represented in a reciprocal matrix of im- munological distance values. Phylogenetic trees derived from these data indicate that the welleri and glutinosus groups are sister taxa, and cluster with the wehrlei group; the cinereus group is the most distant within the eastern Plethodon. This topology conflicts with that based on electrophoretic and morphological data, which suggest that the glutinosus and wehrlei groups are sister taxa. A close examination of the albumin immunological data shows that the rate of albumin evolution within eastern Plethodon has been variable, and therefore these data may be less reliable for phylogenetic reconstruction in this taxon.

RAYMOND, L. R., AND L. M. HARDY. 1990. Demography of a population of Ambystoma talpoideum (Caudata: Ambystomatidae) in northwestern Louisiana. Herpetologica 46:371-382. SEMLITSCH, R. D. 1983. Structure and dynamics of two breeding populations of the eastern tiger salamander, Ambystoma tigrinum. Copeia 1983:608-616. SERVAGE, D. L. 1979. Biosystematics of the Ambystoma jeffersonianum complex in Ontario. Unpubl. MSc Thesis, Univ. Guelph, Guelph, Ontario. 116 pp. STOREZ, R. A. 1969. Observations on the courtship of Ambystoma laterale. J. Herpetol. 3:87-95. UZZELL, T. M. 1964. Relations of the diploid and triplold species of the Ambystoma jeffersonianum complex (Amphibia, Caudata). Copeia 1964:257300. ersonianum complex in Indiana. Herp tologica . 1969. Notes on spermatophore production by salamanders of the Ambystoma jeffersonianum c mplex. Copeia 1969:602-612. WELLER, W. F. 1980. Migration of the salamanders Ambystoma jeffersonianum (Green) and A. platineum Cope to and from a spring breeding pond, and the growth, development and metamorphosis of their young. Unpubl. M.Sc. Thesis, Univ. Toronto, Toronto, Ontario. 242 pp. WILBUR, H. M. 1971. The ecological relationship of the salamander Ambystoma laterale to its all-female, gynogenetic associate. Evolution 25:168-179. ZAR, J. H. 1984. Biostatistical Analysis, 2nd ed. Prentice-Hall, Englewood Cliffs, New Jersey. 718 pp. ZEYL, C. W., AND L. A. LOWCOCK. 1989. A morphometric analysis of hybrid salamanders (genus Ambystoma) and their progenitors on Kelleys Island in Lake Erie. Can. J. Zool. 67:2376-2383.

The syntopic occurrence of two or more species of salamander larvae may be characterized by interspecific differences in size and ontogeny (Worthington, 1968; Hassinger et al., 1970; Keen, 1975; Taylor et al., 1988). The relative timing of breeding seasons and rates of embryo development, larval development, and growth can play important roles in determining whether the period of coexistence of syntopic larvae is dominated by minimal interaction, potential competition, or interspecific predation (e.g., Worthington, 1968; Wilbur, 1972; Stenhouse, 1985). Throughout much of their overlapping ranges, spotted salamanders (Ambystoma maculatum) are often preceded in the onset of breeding by one week or less by blue-spotted salamanders of the A. lateralejeffersonianum complex (e.g., Mohr, 1930; Bishop, 1941; Cook, 1967; Wilson, 1976). Despite a widespread association, published information on larval ecology where these species co-occur is limited. Previous studies have discussed nocturnal spatial relationships (Anderson and Graham, 1967), thermal preferentia (Stauffer et al., 1983), and aspects of larval development (Mohr, 1930; Bishop, 1941). Experimental studies using pond-based enclosures have explored competitive (Wilbur, 1972; Cortwright, 1988) and predatory relationships (Cortwright, 1988). In this report I compare body growth, ontogenetic development, and diet of syntopic larvae of A. maculatum and the A. lateralejeffersonianum complex (A. jeffersonianum and A. laterale-2 jeffersonianum; Nyman et al., 1988) at two New Jersey ponds in 1977. The study ponds, both in Sussex County, New Jersey, USA, were a vernal-autumnal, limestone sinkhole pond (at maximum ca. 0.75 ha and 2 m deep) near Springdale (Anderson and Martino 1966; their "Springdale A"), and a vernal-autumnal, woodland pond (at maximum ca. 0.05 ha and 1.5 m deep) in Culvers Gap (Nyman, 1987). The Springdale pond is only partly shaded by trees along its shoreline and develops extensive, submergent aquatic vegetation (macrophytes and algal mats) in some years, whereas the Culvers Gap pond is shaded much of the day by the surrounding woods and neither macrophytes nor algal mats develop. I determined the onset of breeding by A. maculatum and A. laterale-jeffersonianum complex salamanders at Springdale pond by monitoring salamander movements on a nearby road and capturing salamanders with pitfalls along a 9 m fence placed across a natural migrational corridor (J. D. Anderson, pers. comm.). In the pond I captured salamanders with minnow traps on the pond bottom in an area where salamanders were known to enter the pond (Nyman et al., 1988). By these methods I confirmed that the breeding periods of these salamanders were sequential, but overlapping. Ambystoma laterale-jeffersonianum complex salamanders began to arrive at the pond first and males preceded females of each species; however, even the first large migration (13 March 1977), comprised mostly of A. laterale-jeffersonianum complex females (Nyman et al., 1988) and male A. maculatum, contained a small number of female A. maculatum.

Data on the nesting biology of marbled salamanders (Ambystoma opacum) were obtained on 292 nests from 11 temporary pools in central North Carolina. About 6% of the nests were communal and contained eggs from two or three females. Each of the remaining nests contained a single clutch of eggs. Females placed nests primarily at moderate to high elevations in dried pond beds and rarely nested in pond bottoms. Embryonic survival was dependent on elevation of nests in some ponds, but not in others. About 25% of nests were abandoned within 2-4 weeks after oviposition. Mean egg viability and clutch size were significantly lower in unattended nests than in attended nests. These results are consistent with the view that brooding and nest site selection along elevational gradients are adaptations for terrestrial breeding in Ambystoma opacum.

Comparaison avec le comportement reproducteur d'une autre sous-espece : Notophtalmus viridescens viridescens.

FRASER, D. F. 1976. Empirical evaluation of the hypothesis of food competition in salamanders of the genus Plethodon. Ecology 57:459-471. HAIRSTON, N. G., SR. 1987. Community ecology and salamander guilds. Cambridge University Press, Cambridge. 230 pp. HORNE, E. A., AND R. G. JAEGER. 1988. Territorial pheromones of female red-backed salamanders. Ethology 78:143-152. JAEGER, R. G. 1972. Food as a limited resource in competition between two species of terrestrial salamanders. Ecology 53:535-546. . 1978. Plant climbing by salamanders: periodic availability of plant-dwelling prey. Copeia 1978:686-691.

BURNS, T. A. 1970. Temperature of Yarrow's spiny lizard Sceloporus jarrovi at high altitudes. Herpetologica 26:9-16. CHRISTIAN, K. C. R. TRACY, AND W. P. PORTER. 1983. Seasonal shifts in body temperature and use of microhabitats by Galapagos land iguanas (Conolophus pallidus). Ecology 64:463-468. CROWLEY, S. R. 1985. Thermal sensitivity of sprintrunning in the lizard Sceloporus undulatus: support for a conservative view of thermal physiology. Oecologia (Berl.) 66:219-225. HERTZ, P. E., AND E. NEVO. 1981. Thermal biology of four Israeli agamid lizards in early summer. Israel J. Zool. 30:190-210. HUEY, R. B. 1982. Temperature, physiology, and the ecology of reptiles. In C. Gans and F. H. Pough (eds.), Biology of the Reptilia, Vol. 12, pp. 25-91. Academic Press, London. , AND E. R. PIANKA. 1977. Seasonal variation in thermoregulatory behavior and body temperature of diurnal Kalahari lizards. Ecology 58:10661075.

In the ponds of some regions of France, Triturus cristatus and Triturus marmoratus coexist and natural hybrids can be found. In the course of our study examining evolutionary strategies of newts, the method of individual age determination (skeletochronology) was applied. Cross-sections of femurs and phalanges were stained with Ehrlich hematoxylin. Hematoxylinophilic lines considered to be Lines of Arrested Growth (LAG) were observed and carefully counted. Our results on longevity, median age, sexual maturity and growth rate concur with the observations on life span of the two species in allopatric situations. The general morphology of the hybrids, age at sexual maturity and median life span were intermediate between those of the parental species. In Mayenne (western France), the ranges of two closely related species of newts, Triturus cristatus (Laurenti, 1768) and T. marmoratus (La- treille, 1800) overlap (Thorn, 1968; Rafinski and

Population interactions among three species of salamanders in the genus Desmog- nathus (D. quadramaculatus, D. monticola, and D. ochrophaeus) were studied in the laboratory using an artificial stream system. These salamanders occur sympatrically and are known to partition the habitat along an aquatic-terrestrial gradient. The presence of D. quadramaculatus decreased the activity of D. monticola, but the reverse was not true. D. monticola negatively affected the spatial distribution of D. ochrophaeus. D. ochrophaeus depressed the level of activity of similarly sized D. monticola only. Larger species may represent a predatory threat to smaller species while com- petitive effects may be present among the smaller individuals of each species.

In central Kentucky, adults of Ambystoma texanum become active on the ground surface in early October. Migrations to breeding streams begin in late October and continue through March. Breeding begins in late December or early January, peaks in mid-March, and ends in mid- April. Breeding occurs more or less independently of rainfall and temperature conditions. Males tend to appear on the ground surface and in breeding streams before females. The sex ratio of migrating and breeding adults was 2.2 and 1.5, respectively. The number of mature ova in gravid females was positively correlated with SVL and averaged 262 ? 61. Egg masses averaged 122 ? 97 eggs and were most numerous in shallow, rocky pools.

Verification de l'hypothese de la differentiation ecologique des morphotypes en relation avec la temperature et de l'association de cette differentiation ecologique avec une mortalite et une predation specifiques du morphotype

ABSTRAcr.-Ambystoma maculatum and Rana sylvatica larvae from three ponds in Marquette County, Michigan were raised in the laboratory at pH's 3, 4, 5 and the pond pH. At pH 3, all larvae died within 12 h after exposure. A. maculatum larvae reared at pH 4 and 5 had significantly slower rates of development and growth than larvae reared at pH > 5. The development of R. sylvatica larvae was not significantly different in any of the pH treatments. A survey of 55 county amphibian breeding ponds revealed that 42% of the ponds had a pH < 5.5 and thus, may be inhibiting development and growth of A. maculatum but not R. sylvatica. Based on alkalinity data, 57% of 42 ponds in 1984 were classified as sensitive or highly sensitive to further acidification by acid precipitation. Between May-July 1984, 20% of the A. maculatum breeding ponds dried prior to metamorphosis, producing a possible synergistic mortality effect from desiccation and low pH.

This experiment showed that the courtship behavior of Desmognathus aeneus includes a prolonged biting phase, during which the male bites the female tenaciously. This unusual behavior has been described in only one other plethodontid, Desmognathus wrighti. In most other respects, the courtship behavior of D. aeneus is similar to that of its congeners. Morphological characters associated with biting, as well as the functional and evolutionary significance of biting, are evaluated. The courtship behavior preceding and including insemination in salamanders is elaborate. Among plethodontid salaman- ders there is relatively little diversity in courtship behavior, but numerous studies have documented courtship behavior in plethodontids (Noble and Brady, 1930; Or- gan, 1961a; Salthe, 1967; Arnold, 1972). Sal- amanders have no intromittent organ. In- stead, the male transfers sperm by means of a spermatophore, a gelatinous base sup- porting a sperm mass. After the male courts with the female for a period lasting up to several hours, he deposits a spermatophore on the substrate. He leads the female over