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Morphological diversity and evolution of egg and clutch structure in amphibians

The first part of this synthesis summarizes the morphology of the jelly layers surrounding an amphibian ovum. We propose a standard terminology and discuss the evolution of jelly layers. The second part reviews the morphological diversity and arrangement of deposited eggs?the ovipositional mode; we recognize 5 morphological classes including 14 modes. We discuss some of the oviductal, ovipositional, and postovipositional events that contribute to these morphologies. We have incorporated data from taxa from throughout the world but recognize that other types will be discovered that may modify understanding of these modes. Finally, we discuss the evolutionary context of the diversity of clutch structure and present a first estimate of its evolution.

Herpetological Monographs

Multiple factors affect a population of Agassiz's desert tortoise ( Gopherus agassizii ) in the Northwestern Mojave Desert

Numerous factors have contributed to declines in populations of the federally threatened Agassiz's Desert Tortoise ( Gopherus agassizii ) and continue to limit recovery. In 2010, we surveyed a low-density population on a military test facility in the northwestern Mojave Desert of California, USA, to evaluate population status and identify potential factors contributing to distribution and low densities. Estimated densities of live tortoises ranged spatially from 1.2/km 2 to 15.1/km 2 . Although only one death of a breeding-age tortoise was recorded for the 4-yr period prior to the survey, remains of 16 juvenile and immature tortoises were found, and most showed signs of predation by Common Ravens ( Corvus corax ) and mammals. Predation may have limited recruitment of young tortoises into the adult size classes. To evaluate the relative importance of different types of impacts to tortoises, we developed predictive models for spatially explicit densities of tortoise sign and live tortoises using topography (i.e., slope), predators (Common Raven, signs of mammalian predators), and anthropogenic impacts (distances from paved road and denuded areas, density of ordnance fragments) as covariates. Models suggest that densities of tortoise sign increased with slope and signs of mammalian predators and decreased with Common Ravens, while also varying based on interaction effects involving these predictors as well as distances from paved roads, denuded areas, and ordnance. Similarly, densities of live tortoises varied by interaction effects among distances to denuded areas and paved roads, density of ordnance fragments, and slope. Thus multiple factors predict the densities and distribution of this population.

California;Nevada

Protection benefits desert tortoise ( Gopherus agassizii ) abundance: the influence of three management strategies on a threatened species

We surveyed an area of ∼260 km 2 in the western Mojave Desert to evaluate relationships between condition of Agassiz's Desert Tortoise populations ( Gopherus agassizii ) and habitat on lands that have experienced three different levels of management and protection. We established 240 1-ha plots using random sampling, with 80 plots on each of the three types of managed lands. We conducted surveys in spring 2011 and collected data on live tortoises, shell-skeletal remains, other signs of tortoises, perennial vegetation, predators, and evidence of human use. Throughout the study area and regardless of management area, tortoise abundance was positively associated with one of the more diverse associations of perennial vegetation. The management area with the longest history of protection, a fence, and legal exclusion of livestock and vehicles had significantly more live tortoises and lower death rates than the other two areas. Tortoise presence and abundance in this protected area had no significant positive or negative associations with predators or human-related impacts. In contrast, the management area with a more recent exclusion of livestock, limited vehicular traffic, and with a recent, partial fence had lower tortoise densities and high death rates. Tortoise abundance here was negatively associated with vehicle tracks and positively associated with mammalian predators and debris from firearms. The management area with the least protection—unfenced, with uncontrolled vehicle use, sheep grazing, and high trash counts—also had low tortoise densities and high death rates. Tortoise abundance was negatively associated with sheep grazing and positively associated with trash and mammalian predator scat. cat.

California

Ecology of the Eastern Kingsnake (Lampropeltis getula) at Rainey Slough, Florida: A vanished Eden

Eastern Kingsnakes ( Lampropeltis getula ) are an important component and predator in herpetofaunal communities, but many Eastern Kingsnake populations have declined precipitously in the last few decades, particularly in the southeastern United States. Here, we describe an intensive capture–mark–recapture study of L. getula conducted during 1974–1978 in a canal bank–Water Hyacinth ( Eichhornia crassipes ) community at Rainey Slough in southern Florida, where annual capture probabilities of adults ranged from 0.662–0.787. Population size and structure, seasonal activity, movements, microhabitat use, behavior, thermal ecology, and predator–prey relationships are described. At this site kingsnakes were susceptible to capture mostly in winter and spring, were diurnal, used rodent ( Sigmodon hispidus ) burrows on canal banks as nocturnal retreats, and emerged from burrows on 13–26% of the sampling days. Overlap of burrow use by both sexes was extensive with no evidence of territoriality. Kingsnakes readily entered the Water Hyacinths to bask, pursue mates, and forage. At Rainey Slough only snakes were detected in the diet of kingsnakes. Concurrent sampling of potential snake prey in the hyacinths and on canal banks revealed 10 species that varied in use of the two sampled habitats and in body size. A range-wide analysis confirmed that in descending order snakes, reptile eggs, and lizards dominate the diet of L. getula in Florida (94.8%) and remain important prey types elsewhere (80.2%). At Rainey Slough the density of six species of semiaquatic snakes in Water Hyacinths averaged 3534 individuals/ha with a mean annual biomass of 135.8 kg/ha, and kingsnake biomass was only 2.2–3.9% of prey snake biomass. We estimated that the kingsnake population consumed 36.82–63.58 kg/yr, or about 10.0–17.2% of the standing crop of snakes in the Water Hyacinth community. Adult male L. getula lost on average 39.3% of their body mass associated with the spring reproductive season, whereas females lost only 3.4% in the same period. Body condition indices for both sexes improved substantially thereafter. In follow-up surveys at Rainey Slough during 2006–2010 no kingsnakes were found. Semiaquatic snake densities in the Water Hyacinths were 77.2% lower (807.4/ha) than in the 1970s and consisted of only three species. Compared to the enigmatic declines and extirpation of L. getula populations elsewhere, at Rainey Slough the primary cause likely was unsustainable mortality from road reconstruction and paving in the winter–spring of 1979 and subsequent roadkill. Other potentially causative agents of extirpation of L. getula in this system are discussed.

Florida

Hierarchical, quantitative biogeographic provinces for all North American turtles and their contribution to the biogeography of turtles and the continent

Our study represents the first attempt to describe biogeographic provinces for North American (México, United States, and Canada) turtles. We analyzed three nested data sets separately: (1) all turtles, (2) freshwater turtles, and (3) aquatic turtles. We georeferenced North American turtle distributions, then we created presence–absence matrices for each of the three data sets. We used watershed unit as biogeographic units. We conducted an unweighted pair-group method with arithmetic mean clustering analysis on each Jaccard index distance matrix from our watershed species matrices to delineate biogeographic provinces. Provinces were then tested for significant differences in species compositions in a global model with the use of a one-way analysis of similarity. We conducted a best subset of environmental variables with maximum (rank) correlation with community dissimilarities that determined the best model of abiotic variables explaining province delineation (i.e., climate, topography, and stream channel). To identify which species contributed the most to province delineations, we conducted an indicator species analysis and a similarity-percentage analysis. There were 16 all-turtle provinces, 15 freshwater provinces, and 13 aquatic provinces. Species compositions delineating the provinces were explained by abiotic variables, including mean annual precipitation, mean precipitation seasonality, and diversity of streams. Province delineations correspond closely with geographical boundaries, many of which have Pleistocene origins. For example, rivers with a history of carrying glacial runoff (e.g., Arkansas, Mississippi) sometimes dissect upland provinces, especially for aquatic and semiaquatic turtles. Compared with freshwater fishes, turtles show greater sensitivity to decreased temperature with restriction of most taxa south of the last permafrost maximum. Turtles also exhibit higher sensitivity to climatic, geomorphic, and tectonic instability, with richness and endemism concentrated along the more stable Gulf of México and Atlantic (south of the last permafrost maximum) coasts. Although distribution data indicate two aquatic turtles are most cold tolerant (i.e., Chrysemys picta , Chelydra serpentina ), aquatic turtles overall show the most restriction to warmer, wetter climates. Sequential addition of semiaquatic and terrestrial turtles into analyses shows, as expected, that these taxa flesh out turtle faunas in climatically harsh (e.g., grasslands) or remote (e.g., California, Sonoran Desert) regions. The turtle assemblages of southwestern versus southeastern North America are distinct. But there is a transition zone across the semiarid plains of the Texas Gulf Coast, High Plains, and Chihuahuan Desert, including a strong boundary congruent with the Cochise Filter-Barrier. This is not a simple subdivision of Neotropical versus Nearctic taxa, as some lineages from both realms span the transition zone.

Herpetological Monographs

Four new species of frogs and one new species of snake from the Chapare region of Bolivia, with notes on other species

We report on a collection of 74 frogs, 11 lots of frog eggs or tadpoles, and two snakes collected from the Chapare Region in the yungas of the eastern Andean Cordillera de Cochabamba, Bolivia. Collecting localities range from approximately 300 m to >3200 m in elevation. The specimens pertain to 23 species, probably 10 of which are undescribed. We describe four new species of frogs, one each in the genera Bufo, Centrolenella, Colostethus, and Hyla, and one new species of snake, genus Dipsas. We place Atelopus rugulosus in synonymy with A. tricolor.

Herpetological Monographs

Ecology of the island night lizard, Xantusia riversiana, on Santa Barbara Island, California

We studied the population biology and ecology of the island night lizard (Xantusia riversiana) on Santa Barbara Island, California from 1981-1988. We trapped all the major habitats on the island, using pitfall traps and Sherman small mammal traps. In addition to permanently marking individuals by toe-clipping, we used specialized marking techniques on some individuals (fluorescent powder and radioactive tags) to study home range, movement and activity patterns. Island night lizards are exclusively diurnal and exceedingly secretive. Though generally cryptic in coloration, their dorsal coloration is highly variable. They occur most commonly in habitats providing dense vegetative cover (thickets of boxthorn and prickly pear) and in certain types of rock habitat. Night lizards are sedentary and have very small home ranges (x̄=17.2m 2 ). They are slow-growing with some individuals living to at least 12 years old. The most common injuries (tail and scale injuries) apparently result from intraspecific fighting. A wide variety of terrestrial spiders and insects make up the majority of their diet, but they also feed on an unusually large amount of plant material for a small lizard. Because of their diverse diet, low metabolic rate and small, overlapping home ranges, they reach greater densities (>3200 lizards/ha in boxthorn) than previously reported for any other ground-dwelling lizard. Favorable habitat has been reduced because of past farming and grazing, but, given their wide distribution on the island, exceptionally high densities and current protection within a National Park, X. riversiana on Santa Barbara Island are not threatened with extinction, as was previously thought.

Herpetological Monographs

A 21-year study of seasonal and interspecific variation of hatchling emergence in a nearctic freshwater turtle community: to overwinter or not to overwinter?

Hatchling emergence patterns were studied in a community of six species of freshwater turtles in Lancaster County, Pennsylvania, U.S.A. including: Chelydra serpentina, Chrysemys picta, Clemmys guttata, Glyptemys insculpta, G. muhlenbergii , and Sternotherus odoratus . Data were collected every year from 1965 to 1985 on estimated date of emergence, carapace length, April – May precipitation, August – September precipitation, annual precipitation, and low temperature and occurrence of precipitation during the 24-hrs prior to the time of each hatchling detection (n = 806). Chelydra serpentina, Ch. picta , and Cl. guttata hatchlings have a facultative delayed emergence strategy. The other species ( G. insculpta, G. muhlenbergii , and S. odoratus ) appear to be obligate early emergers, with the exception of one hatchling G. muhlenbergii that delayed emergence. Early emergence occurred in some species every year except 1973, the year following intense flooding and nest destruction associated with a major hurricane. However, the majority of hatchlings delayed emergence until the year following oviposition. Mean estimated calendar day of emergence varied annually in C. serpentina and Ch. picta . The same variable also varied among species for comparisons of both early and delayed emergence. Chelydra serpentina hatchlings emerged earlier than all other species whether they used an early or delayed strategy. Carapace length of Ch. picta hatchlings varied significantly among years and C. serpentina hatchlings that delayed emergence were significantly larger in carapace length than those that emerged early. Seasonal and previous 24-hr precipitation had varying effects on the number of emerging hatchlings, but August – September precipitation in one year had a strong correlation with the number of hatchlings that delayed emergence until the following spring. The number of hatchlings detected peaked at a previous 24-hour air temperature of about 12°C for both early and late emergence. Small species like G. muhlenbergii and S. odoratus may emerge early to limit potential hatchling competition in diverse communities of freshwater turtles with primarily delayed emergence.

Herpetological Monographs

Agkistrodon piscivorus conanti (Florida cottonmouth) Diet

Agkistrodon piscivorus is a generalist predator that feeds on a variety of prey, including snakes (Gloyd and Conant 1990. Snakes of the Agkistrodon Complex: A Monographic Review. Society for the Study of Amphibians and Reptiles, Oxford, Ohio. 614 pp.; Lillywhite et al. 2002. Herpetol. Rev. 33:259–260; Hill and Beaupre 2008. Copeia 2008:105–114). Cemophora coccinea (Scarletsnake) is not known as one of the 26 species of snakes consumed by A. piscivorus (Ernst and Ernst 2011. Venomous Reptiles of the United States, Canada, and Northern Mexico: Volume 1. Johns Hopkins University Press, Baltimore, Maryland. 193 pp.). On 16 June 2015, at 2210 h, we found a dead-on-road A. piscivorus (total length [TL] = 51.0 cm) in Everglades National Park on Main Park Road, 1.88 km S Pa-hay-okee, Miami-Dade Co., Florida, USA (25.414085°N, 80.78183146°W, WGS84; elev. 3 m). The snake had been killed by a vehicle and some internal organs were exposed. Visible stomach contents included a small (TL ca. 15 cm) C. coccinea. Photographic vouchers of the A. piscivorus (UF-Herpetology 177194) and C. coccinea (UF-Herpetology 177195) were deposited in the Division of Herpetology, Florida Museum of Natural History, University of Florida. Despite the fact that these species are sympatric over large areas of the southeastern United States, this is the first known documented predation of C. coccinea by A. piscivorus.

Herpetological Review