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Records of Pleistocene reptiles are relatively well known in different countries. However, reptile fossil records of Pleistocene from Latin America are scarce and limited to presence data associated with records of megafauna remains. Taphonomic studies in Pleistocene reptiles are focused on finding out the origin of the paleontological remains; that is, if the remains found at a particular place correspond to organisms that lived and died there or if they were carried out and deposited there by water currents or predators. Pleistocene reptiles have been used as a proxy for qualitative paleoclimatic reconstructions of localities. In this study, we describe the fossil record of squamate reptiles from the Pleistocene of San Jose Buenavista (Puebla, Mexico), and the taphonomic relationship between them and the associated megafauna, as well as reconstruct the paleoclimate of the paleontological site in Central Mexico where the remains were found. We present the first Mexican Pleistocene record of Anolis and Storeria, the third fossil record of the alligator lizard Barisia sp. in Mexico, and the first North American Pleistocene record for the rattlesnake Crotalus triseriatus. The rattlesnake (C. triseriatus) was found inside the mandible of the American mastodon (Mammut americanum). This finding suggests that the mastodon remains were used as burrow by the rattlesnake during the Pleistocene. Paleoclimate was reconstructed using the Mutual Ecogeographic Range method, which uses the distribution and climate niche of the nearest extant representatives of the identified fossil taxa. A mean annual temperature of 13.91 ± 1.54 °C and an annual precipitation of 964.04 ± 316.82 mm were inferred for the locality. This suggests that the assemblage of fossil species found inhabited the locality during a glacial period.

When discovered by humans, Dekay’s Brownsnake (Storeria dekayi) exhibits a range of defensive behaviours that are likely used to cope with a variety of potential predators. After being handled, a brownsnake at Rondeau Provincial Park, Ontario, Canada, was observed to coil the anterior portion of its body and to sway the coil from side to side as it attempted to flee. Swaying has rarely been documented in Dekay’s Brownsnake and may be a tactic to distract or confuse a visually oriented predator.

We investigated diet breadth and diet overlap in three sympatric snakes of similar body size: Dekay’s Brownsnakes (Storeria dekayi), Red-bellied Snakes (S. occipitomaculata), and sub-adult Common Gartersnakes (Thamnophis sirtalis), by examining recently consumed prey (n = 388) collected from wild-caught snakes (n = 263) in northern Illinois. Storeria occipitomaculata were dietary specialists, feeding nearly exclusively on slugs. Storeria dekayi fed predominately on slugs but also consumed snails and earthworms. Sub-adult T. sirtalis fed predominately on earthworms but also consumed frogs and small mammals. Diet overlap was extensive between Storeria species but relatively low between Storeria and Thamnophis. It is noteworthy that the two most abundant prey types, slugs and earthworms, are non-native. These non-native prey occur in high numbers, which may ameliorate competitive interactions and influence grassland snake abundance and persistence regionally.

Current and historic records of species occurrences in particular localities enable researchers to monitor changes in species distribution and abundance. While current fauna can be documented with sufficient effort, missing historical records represent data that can never again be obtained. It is with this importance in mind that we report for the first time records of eight reptile species in Montgomery County, Indiana, USA, that were documented in 1964, though no vouchered specimens persist. These species include: Sternotherus odoratus, Terrapene carolina carolina, Chrysemys picta marginata, Apalone spinifera spinifera, Thamnophis sauritus sauritus, Storeria dekayi wrightorum, Lampropeltis calligaster, Lampropeltis triangulum. We also provide a list of the 23 currently documented reptile species that occur (or have occurred recently) in Montgomery County.

Systematics at the species level is still marked by theoretical and empirical tensions amongst the desires to identify geographical lineages, delimit species, and estimate their relationships. These goals are often confounded because each relies, at least to some extent, on the others being known. However, recently developed methods can simultaneously address all three. Furthermore, next-generation genomic sequencing allows us to generate large-scale molecular data sets to examine variation within species at a fine scale. Finally, a renaissance in morphological species validation allows us to integrate historical species definitions with coalescent models for species delimitation. Here, we investigate the applicability of these methods in an empirical case, in the Nearctic snake genus Storeria. Integrating trait data into species delimitation reduces the number of species delimited from molecular data alone. Whereas molecular data support eight distinct species-level lineages, including morphological data reduces this to four. The taxa Storeria dekayi, Storeria occipitomaculata, Storeria storerioides, and Storeria victa are considered distinct, monotypic species, with no subspecies recognized. We highlight the need for careful assessment of species delimitation, combining both computational genetic methods as well as traditional character-based descriptions. It is now possible to identify phylogeographical lineages, delimit species using molecular and morphological data, and estimate their relationships in a single coherent set of analyses. Moving forward, this will allow for more rapid and objective assessments of cryptic diversity at the species level.

The defensive behaviors of free-ranging Dekay’s Brownsnakes, Storeria dekayi, were studied at a site in Erie County, Pennsylvania, USA. Twenty-nine unique sequences of defensive behavior were documented. A total of 50 individual snakes (26 males and 24 females) provided 88 observations during the initial phase, of which 78% (n = 69) were of snakes that remained in place. Snakes were tapped with the investigator’s hand to elicit defensive behaviors during the contact phase. Snakes were more than twice as likely to attempt to flee during the contact phase (46%) than during the initial phase (22%). During the contact phase, mean surface body temperatures were significantly higher in snakes attempting to flee (22.3 ± 1.3 °C) than those that remained in place (16.1 ± 2.2 °C). The most frequently observed response during the contact phase was dorso-ventral flattening of the head and body (n = 42). During capture, most snakes (94%) smeared their cloacal contents on themselves and the investigator’s hand.

ResumenSe presentan los registros de una especie de anfibio (Ambystoma flavipiperatum) y 3 de reptiles (Plestiodon dugesii, Plestiodon lynxe y Storeria storerioides), que utilizan las madrigueras de la tuza Pappogeomys bulleri en el bosque de pino-encino del Área Natural Protegida Sierra de Quila, Jalisco.AbstractOne species of amphibian (Ambystoma flavipiperatum) and 3 of reptile were recorded (Plestiodon dugesii, Plestiodon lynxe y Storeria storerioides) using the burrow system of gopher Pappogeomys bulleri. These records were obtained in the temperate forest at the Natural Protected Area Sierra de Quila, Jalisco.

A population of Dekay’s Brownsnake, Storeria dekayi was studied using mark-recapture techniques in Erie County, Pennsylvania, USA during the spring, summer and autumn of 2012. Morphometric data were similar to that reported for the species, with adult females averaging larger and more massive than adult males. However, sexual dimorphism in snout-vent length (SVL) and total length (TL) was not significantly different (P>0.05) in juveniles, although relative tail length (tl/TL) was dimorphic. Relative tail length in both juveniles and adults was greater in males (tl/TL = 0.22-0.27) than females (tl/TL = 0.18-0.23). Storeria dekayi were active from 21 March through 22 October, and displayed a bimodal activity pattern, with peaks in April and August. Using the Schnabel and the Schumacher-Eschmeyer methods, population size was estimated to be 122 ± 19 and 130 ± 35 individuals, respectively. Density was estimated to be 244 and 260 snakes/ha, and biomass 1.60 and 1.71 kg/ha. Additional data regarding population structure, mortality, diet, reproduction, body temperature, movements and site fidelity are also presented.

We ascertained the assemblage structures of snakes occurring in a mixed habitat matrix of natural and disturbed habitats during 2008–2011 at Letterkenny Army Depot (LEAD), a 7200 ha U.S. Army base in south-central Pennsylvania, to understand the patterns of species abundance as they related to habitat type of managed lands. We detected eight species in 12 sites comprising natural and disturbed habitats of wetlands, forest, and thicket and open fields. The Common Gartersnake (Thamnophis sirtalis) occurred in the most sites, the Red-bellied Snake (Storeria occipitomaculata) was the rarest species in the study. Two to six species occupied each site and were distributed unevenly. Dynamics of assemblages could be explained in part by habitat and also by the presence of the North American Racer (Coluber constrictor). All species for which data were available exhibited a unimodal pattern to their seasonal activity (mostly May and June); however, seasonal activity peaks differed between sexes. Sex ratios varied among species but were consistently female–biased in the Common Gartersnake and Ring-necked Snake (Diadophis punctatus) in Pennsylvania and surrounding areas. As elsewhere in Pennsylvania and the Northeast, body sizes of adults were larger for species syntopic with the North American Racer than for species not syntopic with this potential predator. We found a degree of predictability with respect to snake assemblage dynamics among habitats at LEAD, which in turn can prove useful in resource management of this large and protected human-impacted system.

Storeria dekayi anomala Dugès, 1888 e The type from “Orizaba (Mexique)” (Dugès, 1888 e) was redescribed and photographed by Smith & Necker (1943: 210, plate 6, fig. 10). They also reassigned the type locality to Orizaba, Veracruz. Unfortunately, the type has apparently been lost; we were unable to find it after checking all specimens belonging to the genus Storeria at the MADUG. Current valid name: Storeria dekayi anomala Dugès, 1888., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 48, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Snake fungal disease (SFD), caused by the fungus Ophidiomyces ophiodiicola, is an emerging threat to wild snake populations in the US. Data regarding its distribution, prevalence, and population-level impacts are sparse, and more information is needed to better manage SFD in the wild. In this study, we captured 38 wild snakes of five species in Connecticut in the summers of 2015 and 2017. Skin lesions were biopsied and evaluated histologically for fungal dermatitis. At least one individual from each species was positive for SFD, and 48% of snakes sampled in 2015 and 39% of snakes sampled in 2017 were positive for SFD. A Dekay's brownsnake (Storeria dekayi dekayi) with SFD lesions, captured in the summer of 2017, extended the host range of the disease. Thus, SFD was present in wild Connecticut snakes in 2015 and 2017, which demonstrated a wide-spread distribution throughout the state.

A new species of coccidian (Protista: Apicomplexa: Eimeriidae) collected from the faeces of a midland brown snake Storeria dekayi wrightorum Trapido (Ophidia: Colubridae) in Arkansas, USA, is described. Oocysts of Isospora holbrooki n. sp. are subspherical to ovoidal with a smooth, colourless, bi-layered wall, measure on average 27.1 × 24.0 µm, and have a length/width (L/W) ratio of 1.1; both micropyle and oocyst residuum are absent, but a polar granule is present. Sporocysts are ovoidal, 14.8 × 10.0 µm on average (L/W 1.5); the Stieda body is nipple-like, the sub-Stieda body is ellipsoidal and the sporocyst residuum is composed of coarse granules in a cluster. Sporozoites have a spheroidal anterior refractile body, a subspheroidal posterior refractile body, and one centrally-located nucleus. This is the first description of an isosporan from the snake genus Storeria Baird & Girard as well as the largest oocysts and sporocysts of any previous snake isosporan to date.

Understanding habitat selection is a fundamental goal of ecology and is vitally important for effective species conservation. Dekay's Brownsnake (Storeria dekayi; hereafter “Brownsnake”) is one of the most widespread snake species in eastern North America, and its association with cover objects is generally well known, but details regarding choice of refuges are largely unknown. We investigated Dekay's Brownsnake microhabitat use during annual surveys at Point Pelee National Park, Ontario, Canada from 1990 to 2010. We systematically checked all woody debris in stabilized dune habitat for snakes and noted the size and state of decay of woody debris that was used. Nearly all of our observations were of gravid females under woody debris. Greater use of large, moderately decayed woody debris relative to the range of available debris provided strong evidence of selection. Although most snakes observed were solitary, we found a significant number of aggregations. Female Brownsnakes appear to migrate to...

Roads cause direct mortality of animals, but less is known about how this mortality translates into changes in populations. This is particularly true for snakes, which have been subject to little research at the population level compared to other taxa. We studied the effects of proximity to a heavily traveled road on snake populations in three old-field sites at Cicero Swamp Wildlife Management Area in Cicero, New York. We conducted 26 surveys of snakes from June to August 2006 on a regular grid of cover boards at different distances from the road and nearest forest edge and measured the microclimate at each cover board. Adult Common Gartersnakes, Thamnophis sirtalis, and Northern Brown Snakes, Storeria dekayi dekayi, occurred independently of distance from the road but were more abundant farther from the forest edge. The lack of a “road effect” was surprising, given that both species of snakes have been shown to be prone to road-mortality in previous studies and could be caused by either snakes ...

Snakes are capable of moving in one of several ways depending on the substrate and medium being traversed. The locomotor velocities of neonate Brown Snakes (Storeria dekayi) during two modes of terrestrial locomotion (lateral undulation and concertina) and swimming were assessed at 10, 20, and 30 C. At all three temperatures, the fastest velocities were recorded during swimming while the slowest were recorded during concertina locomotion. Velocities of all three modes increased significantly with temperature but Q10 values differed greatly among locomotor modes indicating that the influence of temperature on velocity was mode dependent. Body length of neonates had no significant influence on velocities attained via any locomotor mode at any temperature. Neonate snakes displayed similar, but slightly lower, body length–relative velocities of terrestrial undulation and swimming compared to other snake species tested at the same temperatures. Previous studies have suggested that a trade-off exists b...

In many areas, small fossorial snakes are among the most abundant vertebrates present; yet, the ecology of these species remains poorly understood. Between 1999 and 2002 we collected 210 small fossorial snakes representing five species in a small area of northern Mecklenburg and southern Iredell Counties, North Carolina. The eastern worm snake (Carphophis amoenus; n = 116) was the most frequently captured species in this region, with fewer numbers of ringneck snakes (Diadophis punctatus; n = 44), brown snakes (Storeria dekayi; n = 24), redbellied snakes (S. occipitomaculata; n = 20), and smooth earth snakes (Vir- ginia valeriae; n = 6). The three most abundant species exhibited significant sexual dimorphism, with females being larger and having shorter relative tail lengths than males. Carphophis amoenus were more abundant in dry upland forest than D. punctatus, which were most prevalent in moist, lowland forest. Snake activity was weakly correlated with environmental conditions. A peak in activity of male C. amoenus, D. punctatus, and S. occipitomaculata during September suggests fall breeding seasons for these species.

Summary 1 For gape-limited predators such as snakes, it should be possible to predict the relationship between maximum prey size and body size from the relationship between maximum prey size and gape size and between gape size and body size. Such predictions were generated for Water Snakes, Nerodia sipedon L., using a data subset and then tested with a larger data set. 2 Gape size was computed based on jaw length and width and cyclical regression was used to identify prey of maximum size for snakes of a given gape or mass. 3 Predicted and observed maximum prey cross-section–snake mass allometry were in good agreement. Predicted maximum prey mass–snake mass allometry somewhat exceeded observed allometry which did not differ from 1. 4 Observed minimum prey size–snake size allometry was significantly greater than 0, indicating that larger snakes drop small prey from their diets. 5 Gape size–body size allometry in two other natricine snakes (Thamnophis sirtalis, Storeria dekayi) suggest that patterns of ontogenetic change in prey size should differ among species in predictable ways. 6 Sex differences in gape size–snake size allometry suggest that sex differences in maximum prey size should increase with increasing snake size, even when linear measures of head dimensions do not.

Patterns of population subdivision and gene flow were quantified using allozymes in three sympatric species of natricine snakes from the island region of western Lake Erie. Species share similar population history (post-Pleistocene range expansion followed by subdivision into island and mainland demes) and life-history characteristics. However, species differ in anticipated rates of gene flow because of differences in body size and degree to which they use aquatic habitats. Significant population subdivision was present in all three species, and isolation by distance was evident in water snakes (Nerodia sipedon) and, to a lesser degree, in brown snakes (Storeria dekayi) but not in garter snakes (Thamnophis sirtalis). As predicted, water snakes, the largest and most aquatic of the three species, showed significantly less population structure (lower values of FST) than did the smaller and more terrestrial garter snakes and brown snakes. Furthermore, variance in locus-specific estimates of FST was h...

For a comparative study of facial grooming behavior, we selected five species of colubrid snakes (Thamnophis sirtalis, T. butleri, T. melanogaster, Storeria dekayi, and Cyclophiops semicarinatus) based upon their phylogenetic relationships and natural diets. In an initial descriptive study, head-rubbing behavior reliably followed ingestion of earthworms in all five species. Three forms of head rubbing were identified, including two forms of labial rubbing and one form of rostral area rubbing. The latter occurs during foraging as well as grooming. Two additional controlled experiments were then performed. One experiment assessed the effects of multiple feedings with either normal or immobilized earthworms. The second tested the effects of allowing snakes to contact surface secretions of earthworms and purified mucin without physically contacting the worm. The snakes showed a significantly greater number of head rubs when consuming active earthworms than when consuming immobilized worms, and this increased over the five trials. The snakes also head rubbed more frequently in response to earthworm mucus than to either earthworm surface wash or diluted mucin. The physical properties of earthworm mucus (adhesiveness) thus seem to stimulate head-rubbing behavior. Differences in head rubbing among the species are discussed in relation to natural diet and phylogenetic relatedness. Furthermore, mouth gaping often occurred with head rubbing and can be distinguished from yawning. Tactile and kinesthetic cues associated with this gaping behavior may aid in regulating the occurrence of head rubbing. Neonate snakes head rubbed in a species-typical manner.

Sex differences in body size and head dimensions are widespread in adult snakes, but because data are scarce for neonates, it is unclear whether differences are present from birth or arise post-natally. Here we analyse patterns of sexual dimorphism in neonates and adults of four species of natricine snakes, Nerodia sipedon, Storeria dekayi, Thamnophis radix, and T. sirtalis. Two measures of body size (snout–vent length, mass), four measures of head morphology (head length, head width, jaw length, and interocular distance), and tail length were obtained from wild-caught adults and from offspring born to wild-caught females. Among neonates, significant sexual dimorphism was found in body size for S. dekayi and T. sirtalis, in head dimensions for S. dekayi, T. radix, and T. sirtalis, and in tail length for all four species. Among adults, significant sexual dimorphism was found in body size, head dimensions, and tail length for all four species. The degree of sexual dimorphism in body size among adults greatly exceeded that among neonates. In contrast, the degree of sexual dimorphism in head dimensions was similar between neonates and adults. The presence of significant sexual dimorphism among neonates suggests that hypotheses regarding the evolutionary significance of sexual dimorphism in snakes should consider newborns as well as adults.

Current studies on fetal membranes of reptiles are providing insight into three major historical transformations: evolution of the amniote egg, evolution of viviparity, and evolution of placentotrophy. Squamates (lizards and snakes) are ideal for such studies because their fetal membranes sustain embryos in oviparous species and contribute to placentas in viviparous species. Ultrastructure of the fetal membranes in oviparous corn snakes (Pituophis guttatus) shows that the chorioallantois is specialized for gas exchange and the omphalopleure, for water absorption. Transmission and scanning electron microscopic studies of viviparous thamnophine snakes (Thamnophis, Storeria) have revealed morphological specializations for gas exchange and absorption in the intra-uterine environment that represent modifications of features found in oviparous species. Thus, fetal membranes in oviparous species show morphological differentiation for distinct functions that have been recruited and enhanced under viviparous conditions. The ultimate in specialization of fetal membranes is found in viviparous skinks of South America (Mabuya) and Africa (Trachylepis, Eumecia), in which placentotrophy accounts for nearly all of the nutrients for development. Ongoing research on these lizards has revealed morphological specializations of the chorioallantoic placenta through which nutrient transfer is accomplished. In addition, African Trachylepis show an invasive form of implantation, in which uterine epithelium is replaced by invading chorionic cells. Ongoing analysis of these lizards shows how integration of multiple lines of evidence can provide insight into the evolution of developmental and reproductive specializations once thought to be confined to eutherian mammals.

HAHN, W. E., AND D. W. TINKLE. 1965. Fat body cycling and experimental evidence for its adaptive significance to ovarian follicle development in the lizard Uta stansburiana. J. Exp. Zool. 158:79-86. JAYNE, B. C., AND A. F BENNETT. 1990. Scaling of speed and endurance in garter snakes: a comparison of cross-sectional and longitudinal allometries. J. Zool. (Lond.). 220:257-277. KING, R. B. 1993. Determinants of offspring number and size in the brown snake, Storeria dekayi. J. Herpetol. 27:175-185. NAULLEAU, G., AND X. BONNET. 1996. Body condition threshold for breeding in a viviparous snake. Oecologia 107:301-306. NILSON, G. 1981. Ovarian cycle and reproductive dynamics in the female adder, Vipera berus (Reptilia, Viperidae). Amphib.-Reptilia 2:63-82. SEIGEL, R. A., AND H. S. FITCH. 1985. Annual variation in reproduction in snakes in a fluctuating environment. J. Anim. Ecol. 54:497-505. SEIGEL, R. A., AND N. B. FORD. 1987. Reproductive ecology. In R. A. Seigel, J. T. Collins, and S. S. Novak (eds.), Snakes, Ecology and Evolutionary Biology, pp. 210-252. McGraw-Hill, New York. . 1991. Phenotypic plasticity in the reproductive characteristics of an oviparous snake, Elaphe guttata: implications for life history studies. Herpetologica 47:301-307. SOKAL, R. R., AND F. J. ROHLF. 1981. Biometry. W. H. Freeman and Co., New York. WEATHERHEAD, P. J., G. P. BROWN, M. R. PROSSER, AND K. J. KISSNER. 1999. Factors affecting neonate size variation in northern water snakes, Nerodia sipedon. J. Herpetol. 33:577-589. WHITTIER, J. M., AND D. CREWS. 1990. Body mass and reproduction in female red-sided garter snakes (Thamnophis sirtalis parietalis). Herpetologica 46: 219-226.

Sex, family, and microgeographic variation in body size, head dimensions, tail length, and scalation were assessed from 273 wild-caught brown snakes (Storeria dekayi) from seven island and mainland sites near Lake Erie and from 145 neonates born to 25 wild-caught females. Significant differences between males and females were present both in wild-caught snakes and in neonates, with females exceeding males in snout-vent length and number of ventral scales and males exceeding females in tail length, head dimensions, and number of subcaudal scales. Previous analyses have typically focused on the effect that sex differences might have on adult snakes, e.g, in foraging or reproduction. The presence of sex differences among neonates raises the possibility that these differences may be of ecological and evolutionary significance in younger snakes as well. Significant differences among families were found in neonates for all characters except number of labial scales, and significant heritability (estimated from offspring-dam regression) was found for tail length, head length, and numbers of ventral, subcaudal, and temporal scales. Heritable variation in scalation is well known, but this is the first study to document heritable variation in snake morphology. This result is important because heritable variation is an implicit assumption of hypotheses for the evolution of sex, population, and species differences in morphology. Significant differences among sites were found for adult snout-vent length, head dimensions, number of subcaudal scales, and number of temporal scales. In addition, significant phenotypic correlations (e.g, among head dimensions, between tail length and number of subcaudal scales, between snout-vent length and number of ventral scales) and genetic correlations (e.g, between tail length and number of subcaudal scales, between head length and number of ventral scales) were found between pairs of traits. The presence of these correlations suggests that groups of traits may be influenced by the same genetic or ontogenetic processes and may exhibit patterns of correlated evolution. rnal of Herpetol gy, Vol. 31, No. 3, pp. 335-346, 1997 right 19 7 Society for the Study of Amphibians and Reptiles iation in Brown Snake (Storeria dekayi) Morph logy and Scalation: , Family, and Microgeographic Differences Differences in morphology, behavior, or physiology between sexes, among age classes, or among localities may influence ecological relationships and reflect evolutionary trends. In snakes, sex differences in body size and relative head size may result in reduced diet overlap between males and females (Shine, 1991a; Houston and Shine, 1993) and geographic differences in body size and relative head size may reflect adaptation to local prey characteristics (Forsman and Lindell, 1993). Such hypotheses can be strengthened by an understanding of the genetic and ontogenetic processes giving rise to observed patterns of variation. For example, natural selection will result in sex or geographic differences only if traits show heritable variation. In addition, if multiple traits are genetically or ontogenetically correlated (e.g., if they are influenced by the same genes or by the same ontogenetic processes), then ecological or evolution335 This content downloaded from 157.55.39.4 on Fri, 09 Sep 2016 04:13:43 UTC All use subject to http://about.jstor.org/terms

Determinants of reproductive variables in the brown snake, Storeria dekayi, are examined using path analysis, a method which reveals both direct effects (e.g., of female size on offspring number) and indirect effects (e.g., of female size on offspring size through offspring number). In S. dekayi, there is a clear trade-off between offspring number and offspring size. Part of this trade-off is mediated through the timing of parturition suggesting that placental nourishment contributes to offspring development. However, placental nourishment is not sufficient to result in an uncoupling of offspring number and size. Another clear result is that female size and female condition at ovulation (a size-corrected measure of mass) both influence offspring number. As a result, female size and female condition have indirect effects on offspring size such that overall, larger females and females in better condition produce smaller offspring. Larger females also produce offspring in poorer condition. Additional reproductive mass (mass lost by females at parturition beyond that accounted for by offspring) is determined directly by offspring number suggesting it represents a per-offspring allocation of resources. Captivity has marked effects on repro- duction; females held captive longer produced offspring which were smaller and in poorer condition. Comparisons among species reveals that female size consistently has positive effects on offspring number and offspring size in viviparous snakes but that the effect of offspring number on offspring size varies among species.

We found 20 species of snakes in the Long Pine Key region of Everglades National Park. From 1984-86, data were obtained on 1782 snakes. Peak activity occurred during the wet season from May-Oct. We analyzed activity of the nine most common species in detail. The overall pattern was bimodal; captures peaked in early summer and fall. Coluber constrictor, Diadophis punctatus, Opheodrys aestivus, and Thamnophis sauritus were most common in early summer (MayJuly). Sistrurus miliarius was most common in fall (Oct.). Elaphe guttata, E. obsoleta, Storeria dekayi, and T. sirtalis had two or more peaks of activity. Species samples were broken down into adult males, females, and non-adults (juveniles plus young-of-the-year [YOY]). Males were more common in the samples of C. constrictor, E. guttata, E. obsoleta, and Sistrurus miliarius. Females were more common in D. punctatus, 0. aestivus, Storeria dekayi, and T. sirtalis. Both sexes were equally common in T. sauritus. Juveniles and YOY were especially common in the fall in E. obsoleta, Storeria dekayi, T. sirtalis, and Sistrurus miliarius, and contributed significantly to a second peak in numbers. Individuals of many species were encountered at varying frequencies over all 12 mo in a year. Since activity patterns and/or relative abundances for the subgroups were different in most species, we conclude that activity curves for snake species that do not include data on sex and age may be of limited value for study of phenology and reproductive ecology. Data on hatching and birth dates are presented, and when taken with activity data reveal only moderate alteration of typical temperate zone patterns of reproduction and activity in this subtropical region.

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The helminth parasites of 141 snakes were collected. The nematode Rhabdias fuscovenosa was found in all snake species except Nerodia sipedon, whereas Cosmocercoides dukae was restricted to Storeria dekayi. The trematodes Zeugorchis aequatus and Lechriorchis primus occurred only in Thamnophis sirtalis, whereas Dasymetra nicolli infected only N. sipedon. Pneumatophilus variabilis primarily parasitized N. sipedon but was occasionally found in T. sirtalis. Mesocercariae of Alaria sp. were found in all species except S. dekayi. The cestodes Ophiotaenia sp. and Cylindrotaenia sp. were found in N. sipedon and T. sirtalis, respectively.

species, in eastern Mexico, chiefly in the area of Volc.in San Martin of southern Veracruz. The actual length of time available at the latter site was but two days, but even this brief stay verified the zoogeographic significance of the region previously suggested by announcement of the extraordinary and unique toad Bufo cavifrons Firschein (1950). One new salamander was secured there, a snake new to Mexico (Tantilla schistosa), a series of Scincella requiring allocation of caudaequinae as a subspecies of S. silvicolum, and a number of rarities and useful locality records in addition to a second example of Bufo carifrons. Specimens secured en route to and from the volcano include a series of tadpoles presumably representing Hyla miolympanum (not heretofore described), and records of Storeria dekayi temporalineata from the states of San Luis Potosi and Tamaulipas.

A female DeKay's snake, 348 mm. long, gave birth to 16 young on August 4, 1943. One of the young was dead at birth. The remaining 15 were separated from the female and placed in a glass-sided aquarium 50 cm. by 25 cm., the bottom of which was covered with 3 or 4 cm. of damp sand. One-half of the sanded area was covered with moss; the other half contained a small water dish, a food dish, and for hiding places a potsherd and a piece of bark. The water was changed weekly, and fresh food was offered daily except as noted elsewhere. The total weight of the brood and the length of the snakes were taken at frequent intervals until observations were terminated on October 31.

Examination of specimens of Storeria from coastal areas of Louisiana and Texas indicates the existence of a distinctive population inhabiting the coastal marshes. On the basis of color pattern and scutellation this population is sharply distinct from populations of Storeria dekayi on adjacent higher ground and appears closely allied to Storeria of Mexico, Guatemala and Honduras. The few specimens suggesting possible interbreeding with contiguous high ground populations come from areas of ecological disturbance and appear to represent occasional hybridization, rather than an indication of intergradation. Specimens believed by Trapido to represent intergrades between Storeria dekayi temporalineata and S. d. texana or S. d. wrightorum were derived from the distinctive marsh population, and the southern and northern groups of subspecies placed by Trapido in the species Storeria dekayi thus appear to be reproductively isolated. Storeria tropica Cope is therefore revived for the southern group, with four subspecies; Storeria tropica tropica, Storeria tropica anomala, Storeria tropica temporalineata, and Storeria tropica limnetes, subsp. nova, the population of the Louisiana and Texas coastal marshes. In his monograph of the snakes of the genus Storeria, Trapido (1944) recognized six subspecies of Storeria dekayi, distributed from the northeastern United States to Guatemala and Honduras. Three of these subspecies, S. d. texana, S. d. wrightorum, and S. d. dekayi occur north of the Rio Grande and replace each other from west to east, between the Great Plains and the Atlantic Coast. As defined by Trapido, the easternmost subspecies, dekayi, is found from the Atlantic Seaboard to the western side of the Appalachians. Its northern limit is the limit of the species distribution in Quebec and it intergrades with S. d. wrightorum over a broad area extending from the Carolinas around the southern Appalachians and northward to Illinois. Subspecies wrightorum extends from this zone of intergradation westward across the Mississippi valley and intergrades with S. d. texana along a line from Iowa to western Louisiana. Subspecies texana is found between this line and the eastern edge of the Great Plains and extends southward to the gulf coast of Texas and into Mexico as far south as Hidalgo. The three re.maining subspecies which 'Trapido recognized are found in Mexico, Guatemala and Honduras. They are described as replacing each other southward along the gulf coastal plain, with S. d. tem poralineata extending from the Rio Grande south into Veracruz, where S. d. anomala occurs. The latter is said to be characteristic of an area about the towns of Orizaba and Jalapa. It presumably intergrades with S. d. tropica, which inhabits the Atlantic drainages of Guatemala and Honduras.

SAINT GIRONS, J. 1962. Presence de receptacles seminaux chez les cameleons. Beaufortia 9: 165-172. TRAPIDO, H. 1940. Mating time and sperm viability in Storeria. Copeia 1940:107-109. TINKLE, D. W. 1961. Population structure and reproduction in the lizard Uta stansburiana stejnegeri. Amer. Midi. Nat. 66:206-234. , D. MCGREGOR, AND S. DANA. 1962. Home range ecology of Uta stansburiana stejnegeri. Ecology 43:223-229.

TRAPIDO, HAROLD 1940 Mating time and sperm viability in Storeria. Ibid., 1940: 107-109. WOOD, F. D. 1933 Mating of the prairie rattlesnake Crotalus confluentus confluentus Say. Ibid., 1933: 84-87. WOODWARD, S. F. 1933 A few notes on the persistence of active spermatozoa in the African night adder, Causus rhombeatus. Proc. Zool. Soc. London, 1933: 189-190. WUNDERER, H. 1910 Beitrige zur Biologie und Entwicklungsgeschichte des Alpensalamanders (Salamandra atra Laur.). Zool. Jahrb., (Syst.), 28: 23-80.