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This paper explores the author’s clinical work with a young boy between the ages of four and eleven. The author postulates that there are, in addition to “defenses of the self” that develop in a pa...

The fungal pathogen Ophidiomyces ophiodiicola, the causative agent of snake fungal disease, has been implicated in declines of North American snake populations since 2006 and the geographic range of this pathogen is still not fully known. In Tennessee, US, O. ophiodiicola has been detected since 2012, but large portions of the state have not been surveyed for this pathogen. Our primary objective was to monitor the prevalence of O. ophiodiicola in the Interior Plateau ecoregion of Tennessee by swabbing all snakes that were encountered during road cruising survey efforts in 2017 and 2018. Eleven snakes of four species, copperhead (Agkistrodon contortrix), common water snake (Nerodia sipedon), black kingsnake (Lampropeltis nigra), and smooth earthsnake (Virginia valeriae), tested positive for the presence of O. ophiodiicola. Overall, 9.2% (11/120) of snakes sampled tested positive for the presence of O. ophiodiiola, and we further observed a seasonal trend in detections with summer months having the greatest frequency of detections. Our results extend the known geographic range of O. ophiodiicola in Tennessee by adding four previously unconfirmed O. ophiodiicola-positive counties. Further sampling will need to be conducted across west Tennessee because this is the most data-deficient region of the state. Our results offer additional evidence of the presence of this pathogen in Tennessee and will help researchers further understand the geographic distribution and host range.

Snakes can move through almost any terrain. Similarly, snake robots hold the promise as a versatile platform to traverse complex environments such as earthquake rubble. Unlike snake locomotion on flat surfaces which is inherently stable, when snakes traverse complex terrain by deforming their body out of plane, it becomes challenging to maintain stability. Here, we review our recent progress in understanding how snakes and snake robots traverse large, smooth obstacles such as boulders and felled trees that lack “anchor points” for gripping or bracing. First, we discovered that the generalist variable kingsnake combines lateral oscillation and cantilevering. Regardless of step height and surface friction, the overall gait is preserved. Next, to quantify static stability of the snake, we developed a method to interpolate continuous body in three dimensions (3D) (both position and orientation) between discrete tracked markers. By analyzing the base of support using the interpolated continuous body 3-D kinematics, we discovered that the snake maintained perfect stability during traversal, even on the most challenging low friction, high step. Finally, we applied this gait to a snake robot and systematically tested its performance traversing large steps with variable heights to further understand stability principles. The robot rapidly and stably traversed steps nearly as high as a third of its body length. As step height increased, the robot rolled more frequently to the extent of flipping over, reducing traversal probability. The absence of such failure in the snake with a compliant body inspired us to add body compliance to the robot. With better surface contact, the compliant body robot suffered less roll instability and traversed high steps at higher probability, without sacrificing traversal speed. Our robot traversed large step-like obstacles more rapidly than most previous snake robots, approaching that of the animal. The combination of lateral oscillation and body compliance to form a large, reliable base of support may be useful for snakes and snake robots to traverse diverse 3-D environments with large, smooth obstacles.

A 10-year-old, intact, captive-born female Mexican black kingsnake ( Lampropeltis getula nigrita ) was presented with a 1-week history of caudal coelomic distention. On physical examination a 5cm in length, soft nonmobile mass was identified in the caudal third of the coelomic cavity, 4cm cranially to the cloaca. Survey radiographic images revealed an increased soft tissue opacity within the affected area with variable sized, irregular mineralized structures. An intracloacal contrast study showed contrast retention in the cloaca. The retention of contrast within the cloaca was compatible with the presence of a mass compressing the rectum. An ultrasound evaluation confirmed a 2 × 3cm, oval, well-defined mass with an echogenic capsule. Computed tomographic imaging revealed a heterogeneous mass with some mineral areas with irregular shape and size. The mass was excised and the tissue submitted for histopathology and culture. The histopathological and microbiological diagnosis of the mass was purulent interstitial nephritis with presence of multiple histiocytic granulomas that contained intralesional bacteria ( Morganella morganni ). The animal responded well to the surgical removal of the mass and subsequent medical treatment.

Early amniotic vertebrates evolved large-yolked eggs that permitted production of well-developed, terrestrial hatchlings. This reproductive pattern required new mechanisms for cellularizing the yolk and mobilizing it for embryonic use. In birds, cells that line the yolk sac cavity phagocytose and digest the yolk material, a pattern that is commonly assumed to be universal among oviparous amniotes. However, recent evidence challenges the assumption that all squamate reptiles conform to the avian developmental pattern. In this paper, scanning electron microscopy and histology were used to study mechanisms of yolk processing in two colubrid snakes, the kingsnake Lampropeltis getula and the milksnake L. triangulum. Endodermal cells from the yolk sac splanchnopleure proliferate massively as they invade the yolk sac cavity, forming elaborate chains of interlinked cells. These cells grow in size as they phagocytose yolk material. Subsequently, vitelline capillaries invade the masses of yolk-laden cells and become coated with the endodermal cells, forming an elaborate meshwork of cell-coated strands. The close association of cells, yolk, and blood vessels allows yolk material to be cellularized, digested, and transported for embryonic use. The overall pattern is like that of the corn snake Pantherophis guttatus, but contrasts markedly with that of birds. Given recent evidence that this developmental pattern may also occur in certain lizards, we postulate that it is ancestral for squamates. Studies of lizards, crocodilians, and turtles are needed to clarify the evolutionary history of this pattern and its implications for the evolution of the amniotic (terrestrial) vertebrate egg.

Many snakes live in deserts, forests, and river valleys and traverse challenging 3-D terrain like rocks, felled trees, and rubble, with obstacles as large as themselves and variable surface properties. By contrast, apart from branch cantilevering, burrowing, swimming, and gliding, laboratory studies of snake locomotion focused on that on simple flat surfaces. Here, to begin to understand snake locomotion in complex 3-D terrain, we study how the variable kingsnake, a terrestrial generalist, traversed a large step of variable surface friction and step height (up to 30% snout-vent length). The snake traversed by partitioning its body into three sections with distinct functions. Body sections below and above the step oscillated laterally on horizontal surfaces for propulsion, while the body section in between cantilevered in a vertical plane to bridge the large height increase. As the animal progressed, these three sections traveled down its body, conforming overall body shape to the step. In addition, the snake adjusted the partitioned gait in response to increase in step height and decrease in surface friction, at the cost of reduced speed. As surface friction decreased, body movement below and above the step changed from a continuous lateral undulation with little slip to an intermittent oscillatory movement with much slip, and initial head lift-off became closer to the step. Given these adjustments, body partitioning allowed the snake to be always stable, even when initially cantilevering but before reaching the surface above. Such a partitioned gait may be generally useful for diverse, complex 3-D terrain.

Prescribed fire is an essential tool for the restoration and maintenance of the longleaf pine (Pinus palustris) ecosystem. This type of management benefits wildlife species that are frequently associated with open canopy pine forests in the Southeastern U.S., including upland snakes. But few studies have examined the long term effects of prescribed fire (i.e., fire frequency over time) on snake occurrences at local and landscape scales. Likewise, comprehensive data are lacking on the response of snakes to individual prescribed burns (i.e., how long after a fire does a snake occupy a location). To investigate snake responses to these two factors, we used radio-telemetry data for five species of snakes, including three species of open-canopied forest specialists and two species that occur more broadly in upland forests (i.e., generalists), on a large site managed with prescribed fire. We hypothesized that open-canopied forest specialists, eastern diamondback rattlesnake (Crotalus adamanteus), Florida pine snake (Pituophis melanoleucus), and eastern coachwhip (Coluber flagellum) would select more frequently burned locations over less frequently burned locations and would occupy locations sooner after a fire than generalist species, eastern kingsnake (Lampropeltis getula) and gray ratsnake (Pantherophis allegheniensis). We used spatial data collected from previous snake radio-telemetry studies conducted over a nine year period (2003–2012) and annual burn data from 1994 to 2012. We used compositional analysis to evaluate snake use (selection) versus availability of locations based on fire frequency at different spatial scales (home range and landscape). As predicted, open-canopied forest specialists selected locations with more frequent fire (≤2.5 y return interval) significantly more often than locations with less frequent fire (≥3.3 y return interval; F7, 18 = 9.06, p

A 22-yr-old, female gray-banded kingsnake ( Lampropeltis alterna) was examined for stomatitis and hard ventral scales. On examination, the ventral scales palpated as rigid and brittle. A blood sample yielded marked hypovitaminosis D (11 nmol/L). Radiographs revealed a generalized bone radiopacity outlining the ventral scales; scale biopsies were obtained for histologic examination and presented linear, superficial, and midzonal foci of well-differentiated dermal bone and bone marrow. The stomatitis was successfully treated, but the general condition of the snake declined, and it was euthanized. The most significant finding in the postmortem examination was abnormally pale kidneys. Histologic examination revealed diffuse renal gout and diffuse osseous metaplasia in the dermis of all of the ventral scales. Generalized osseous metaplasia of the ventral scales in snakes has not, to our knowledge, been reported; the cause in this report was not identified, but given the extensiveness of the lesion, it is likely that the cause was multifactorial.

Within many biomes, the cause of phylogeographic structure remains unknown even across regions throughout North America, including within the biodiverse Chihuahuan Desert. For example, little is known about population structure or the timing of diversification of Chihuahuan endemics. This is due largely to the lack of population genomic studies within this region. We generated ultra-conserved element data for the gray-banded kingsnake (Lampropeltis alterna) to investigate lineage divergence and historical demography across the Chihuahuan Desert. We found three unique lineages corresponding to the Trans-Pecos and Mapimian biogeographic regions of the Chihuahuan Desert, and a distinct population in the Sierra Madre Occidental. Using several mutation rates to calibrate the timing of divergence among these lineages, we show that lineage divergence likely occurred during the Pleistocene, which indicates that careful consideration needs to be used when applying mutation rates to ultra-conserved elements. We suggest that biogeographic provinces within the Chihuahuan Desert may have served as allopatric refugia during climatic fluctuations of the Quaternary. This work serves as an important template for further testing biogeographic hypotheses within the region.

Coral snakes and their mimics often have brightly colored banded patterns, generally associated with warning coloration or mimicry. However, such color patterns have also been hypothesized to aid snakes in escaping predators through a “flicker-fusion” effect. According to this hypothesis, banded color patterns confuse potential predators when a snake transitions from resting to moving because its bands blur together to form a different color. To produce this motion blur, a moving snake’s bands must transition faster than the critical flicker-fusion rate at which a predator’s photoreceptors can refresh. It is unknown if coral snakes or their mimics meet this requirement. We tested this hypothesis by measuring the movement speed and color patterns of two coral snake mimics, Lampropeltis triangulum campbelli and L. elapsoides, and comparing the frequency of color transitions to the photoreceptor activity of the avian eye. We found that snakes often produced a motion blur, but moving snakes created a blurring effect more often in darker conditions, such as sunrise, sunset, and nighttime when these snakes are often active. Thus, at least two species of coral snake mimics are capable of achieving flicker-fiision, indicating that their color patterns may confer an additional defense aside from mimicry.

It is not clear if rodents express inducible defenses in response to reptilian predators such as snakes. We tested the hypothesis that adult house mice Mus musculus decrease aspects of their reproductive effort upon 1 hour of exposure every 48 hours for a 25-day period to the fecal material and shed skins of a euryphagous ophidian predator, the kingsnake Lampropeltis getula, that had been fed mice. We found no significant differences in the total number of offspring born, the number of pups per litter, and the mean weight of pups in litters born to male and female mice that were exposed to predator cues and those mice that were not exposed to such cues. The lack of an inducible response may be associated with the low cost of an effective defense, or the lack of an effective defense against a generalist snake predator.

Many rodents exhibit inducible defenses when exposed to chemical cues from mammalian predators. These re- sponses may include delays in sexual maturation, smaller adult body size and decreases in litter size and pup weight. We exposed the hybrid juvenile offspring of field-caught and lab-descended house mice Mus musculus to the chemical cues of mouse-fed or chick-fed kingsnakes, Lampropeltis getula, for 20 days after weaning, to examine the effects of ophidian predator cues on prey development. We hypothesized that these cues would elicit inducible defenses such as alteration of growth rates, and/or the timing of reproductive development in mice. Once mature, the reproductive effort of the mice might also be impacted by producing smaller litter sizes or lighter pups or not reproducing at all. We found no effect of kingsnake cues on any of the measures. These findings support the hypothesis that inducible defenses may have evolved as a strategy to deal with specific predators (Current Zoology 58 (6): 797-804, 2012). Keywords Inducible defenses, House mice, Kingsnakes, Chemical cues, Growth and development

We used radio-telemetry to identify diel activity patterns in the Eastern Kingsnake, Lampropeltis getula getula in Southwestern Georgia. We monitored tagged snakes at four time periods within a 24-hour cycle (0800–1359 h, 1400–1959 h, 2000–0159 h, and 0200–0759 h) in spring, summer, and fall (March–November), which comprise the annual activity period for Eastern Kingsnakes in this region. We used a logistic regression approach to identify specific environmental factors that best predicted Eastern Kingsnake movement. We found Eastern Kingsnakes were primarily active during the day (χ2 = 34.10, P < 0.001, n = 452) in the spring (χ2 = 6.25, P < 0.04, n = 474) and activity was best predicted by air temperature, soil temperature, relative humidity, and light intensity. Our observations differed from anecdotal descriptions of nocturnal activity and seasonal shifts in diel activity patterns in Eastern Kingsnakes.

Studies of speciation and taxon delimitation are usually decoupled. Combining these methods provides a stronger theoretical ground for recognizing new taxa and understanding processes of speciation. Using coalescent methods, we examine speciation, post-speciation population demographics, and taxon delimitation in the Arizona Mountain Kingsnake (Lampropeltis pyromelana), a species restricted to high elevations in southwestern United States and northern Mexico (SW). These methods provide a solid foundation for understanding how biogeographic barriers operate at the regional scale in the SW. Bayesian species delimitation methods, using three loci from samples of L. pyromelana taken throughout their range, show strong support for the existence of two species that are separated by low elevation habitats found between the Colorado Plateau/ Mogollon Rim and the Sierra Madre Occidental. Our results suggest an allopatric mode of speciation given the near absence of gene flow over time, which resulted in two lineages of unequal population sizes. Speciation likely occurred prior to the Pleistocene, during the aridification of the SW and/or the uplift of the Colorado Plateau, and while these species occupy similar high-elevation niches, they are isolated by xeric conditions found in the intervening low deserts. Furthermore, post-speciation demographics suggest that populations of both lineages were not negatively impacted by climate change throughout the Pleistocene. Finally, our results suggest that at least for this group, where divergence is old and gene flow is low, Bayesian species delimitation performs well.

This paper reports information on size and growth of snakes in two populations of Lampropeltis nigra (Black Kingsnake) over 20 years of study and provides a comparative analysis that builds on the work of Jenkins et al. (2001). During a 7-year study (1990–1996) at the Anderson County Wildlife Sanctuary (ACWS) and a 13-year study (1997–2009) at the University of Tennessee Forestry Resources Research and Education Center (FES) in Oak Ridge, TN, we captured 265 individual Black Kingsnakes a total of 556 times. The size of Black Kingsnakes in these two populations are the smallest reported for this species, with mean (± SD) snout-to-vent length (SVL) of 66.9 ± 24.5 cm (maximum = 112 cm) at ACWS and 55.8 ± 16.8 cm (maximum = 87 cm) at FES. At FES, the mass-SVL relationship is represented by an exponential equation (mass [g] = 0.0004 SVL [cm]-2,98) similar to ACWS (mass [g] = 0.0005 SVL [cm]2,95). Across both sites, juvenile kingsnakes grew 1.1 cm/mo faster than adult individuals. There was a significa...

The Common Kingsnake, Lampropeltis getula, is thought to be experiencing population declines in the southeastern portion of its geographic range. However, limited information exists regarding the natural history and habitat requirements of the species in this region. We conducted a radio-telemetry study to investigate habitat selection in L. getula at multiple scales in southwestern Georgia. At the home range scale, L. getula did not show habitat selection. However, at the study-area scale, L. getula selected for natural pine and hardwood forest types over other available habitats. At the microhabitat scale, L. getula were found in locations with more coarse woody debris and woody vegetation than random sites. Lampropeltis getula primarily used small mammal (Peromyscus spp.) burrows and stumpholes as below ground refuge sites. We demonstrate that habitat selection of L. getula is complex and occurs at multiple scales.

3 Corresponding author Abstract We present a systematic revision of the Lampropeltis getula group, based on a recent range-wide phylogeographic analysis. We define our theoretical and operational concepts of species delimitation, and provide diagnoses based on mitochondrial DNA evidence, ecological niche modeling, morphology, and historical precedence. We find support for the recognition of five distinct species, which bear the name of the nominate subspecies found primarily within the range of each phylogeographic lineage: the Eastern lineage (Lampropeltis getula, Eastern Kingsnake), the Mississippi lineage (L. nigra, Black Kingsnake), the Central lineage (L. holbrooki, Speckled Kingsnake), the Desert lineage (L. splendida, Desert Kingsnake), and the Western lineage (L. californiae, California Kingsnake). Interestingly, all of these taxa had originally been described as distinct species and recognized as such for up to 101 years (in the case of L. californiae) before being demoted to subspecies. We discuss the impact that increasingly detailed genetic information from phylogeographic analyses may have on traditional taxonomy.

Niche conservatism and niche divergence are both important ecological mechanisms associated with promoting allopatric speciation across geographical barriers. However, the potential for variable responses in widely distributed organisms has not been fully investigated. For allopatric sister lineages, three patterns for the interaction of ecological niche preference and geographical barriers are possible: (i) niche conservatism at a physical barrier; (ii) niche divergence at a physical barrier; and (iii) niche divergence in the absence of a physical barrier. We test for the presence of these patterns in a transcontinentally distributed snake species, the common kingsnake (Lampropeltis getula), to determine the relative frequency of niche conservatism or divergence in a single species complex inhabiting multiple distinct ecoregions. We infer the phylogeographic structure of the kingsnake using a range-wide data set sampled for the mitochondrial gene cytochrome b. We use coalescent simulation methods to test for the presence of structured lineage formation vs. fragmentation of a widespread ancestor. Finally, we use statistical techniques for creating and evaluating ecological niche models to test for conservatism of ecological niche preferences. Significant geographical structure is present in the kingsnake, for which coalescent tests indicate structured population division. Surprisingly, we find evidence for all three patterns of conservatism and divergence. This suggests that ecological niche preferences may be labile on recent phylogenetic timescales, and that lineage formation in widespread species can result from an interaction between inertial tendencies of niche conservatism and natural selection on populations in ecologically divergent habitats.

The habitat use and activity range of Lampropeltis getula getula (Eastern Kingsnake) in the New Jersey Pine Barrens were studied from 1996–1998. Five male and four non-gravid female Eastern Kingsnakes were routinely radiotracked during daylight hours during one or two active seasons. Habitat and climatic conditions at snake locations were characterized using 9 climatic and 14 structural habitat features. Multivariate statistical comparisons with randomly selected locations indicated that Eastern Kingsnakes use available habitat in a non-random fashion with respect to microhabitat features (Wilks' lambda = 0.511; df = 28, 1066; P < 0.01). Eastern Kingsnakes preferred sites with thick leaf litter and dense shrub-layer foliage. They used a broad range of macrohabitats that spanned both wetland and pine-dominated upland areas. Moist areas were used for hibernation. Snakes exhibited a largely fossorial lifestyle, spending a great proportion of their time concealed under the cover of soil and/or leaf l...

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...

We report the first observation of depredation on a nest of the eastern meadowlark (Sturnella magna) by a speckled kingsnake (Lampropeltis getula holbrooki). Depredation events probably are opportunistic, but they may have an impact on local populations of eastern meadowlarks.

Morphology of kingsnakes, Lampropeltis getula, is described and analyzed in the Apalachicola region of the Florida panhandle. Populations inhabiting the eastern Apalachicola Lowlands, a distinct biotic province, are different from the surrounding populations in having fewer and wider light body crossbands, distinct ontogenetic interband lightening, unique ventral patterns, and the presence of non-banded (striped and patternless) individuals. We conclude that the name L. g. goini as well as the hypothesis that Apalachicola L. getula are relict populations of intergrades between L. g. getula and L. g. floridana are invalid. We believe the polymorphic eastern Apalachicola Lowlands populations are most closely related to L. g. getula, and evolved in isolation on a barrier island or the coastal strand of a peninsula during one of the many higher stands of sea in the Pleistocene.

The unusual morphological adaptations associated with specialization on a diet of eggs have been used as evidence to justify familial or subfamilial status for the egg-eating snakes, including the African genus Dasypeltis. We evaluate the phylogenetic relationships of Dasypeltis scabra by comparing albumin evolution in Dasypeltis and available representative colubrine taxa using the quantitative immunological technique of micro-complement fixation. We find that Dasypeltis scabra is included among a large colubrine assemblage containing racers (Coluber, Masticophis, Spalerosophis, and Ptyas), the ratsnake Elaphe, and the kingsnake Lampropeltis. Further, these data suggest that Dasypeltis is genetically most similar to some members of a racer lineage and arose from an ancestral colubrine stock as early as ten million years ago.

Young corn snakes,Elaphe guttata, were tested for responses to chemicals from heterospecific snakes. Corn snakes exhibited more tongue-flicks to swabs freshly rubbed against the skin of an ophiophagous kingsnake,Lampropeltis getulus, than to blank swabs. Responses toL. getulus and a nonophiophagous western plains garter snake,Thamnophis radix haydeni, did not differ significantly. Corn snakes exhibited more tongue-flicks to swabs treated with chloroform extracts of the shed skins ofL. getulus; an ophiophagous eastern coachwhip,Masticophis flagellum; and a nonophiophagous gray ratsnake,Elaphe obsoleta, than to blank swabs, but they did not discriminate between ophiophagous and nonophiophagous species in every case. Corn snakes, when offered shelters containing bedding from the home cages of a nonophiophagous water snake,Nerodia erythrogaster, an occasionally ophiophagous water moccasin,Agkistrodon pisdvorus; orL. getulus and untreated bedding, failed to reside under snake-scented shelters at a rate significantly different from that expected by chance. The responses of corn snakes are compared with those reported for other snakes presented with heterospecific snake chemicals.

In this study we investigated how ophiophagous snakes are able to ingest prey snakes that equal or exceed their own length. We used X-ray video, standard video, dissection, and still X-rays to document the process of ophiophagy in kingsnakes (Lampropeltis getula) feeding on corn snakes (Elaphe guttata). Most kingsnakes readily accepted the prey snakes, subdued them by constriction, and swallowed them head first. In agreement with previous observations of ophiophagy, we found that the predator snake forces the vertebral column of the prey snake to bend into waves. These waves shorten the prey's body axis and allow it to fit inside the gastrointestinal (GI) tract and body cavity of the predator. Dissection of a kingsnake immediately following ingestion revealed extensive longitudinal stretching of the anterior portion of the GI tract (oesophagus and stomach), and no visible incursion of the prey into the intestine. X-ray video of ingestion showed that the primary mechanism of prey transport was the pterygoid walk, with some contribution from concertina-like compression and extension cycles of the predator's vertebral column in two out of three observations. Complete digestion was observed in only one individual, as others regurgitated before digestion was finished. X-ray stills taken every 4 days following ingestion revealed that the corn snakes were about half digested within the first 4 days, and digestion was complete within 15 days.

In March 2000, an approximately 30-yr-old, male coastal mountain kingsnake (Lampropeltis zonata multifasciata) presented with disequilibrium and unresponsiveness to stimuli that ultimately lead to euthanasia. Histologically, there were foci of gliosis primarily within the caudal cerebrum, brainstem, and cervical spinal cord. Several glial cells and endothelial cells contained magenta, intranuclear inclusion bodies. Electron microscopy of the inclusions revealed paracrystalline arrays of 79–82 nm, viral-like particles. DNA in situ hybridization of sections of formalin-fixed brain using a mixture of two digoxigenin-end-labeled, adenovirus specific, oligonucleotide probes at low and high stringency was positive for adenovirus.

Common Kingsnakes (formerly known collectively as Lampropeltis getula) are experiencing localized declines throughout the southeastern United States. Because there have been limited studies to determine how snakes regulate prey populations, and because Kingsnake declines may result in ecosystem impacts, we evaluated the hypothesis that Kingsnakes regulate the abundance of one of their prey, the venomous Copperhead (Agkistrodon contortrix). We generated a database of captures of the two species across the southeastern United States and, while controlling for large-scale habitat preferences, identified a negative relationship between the relative abundance of Kingsnakes and the relative abundance of Copperheads. Our results are correlative but consistent with the hypothesis that Copperhead populations experience a release from predation pressure where Kingsnake abundances are low. We suggest that Kingsnake declines, which are occurring for unknown reasons, are having ecological effects in affected ecosystems. We further highlight the potential role that snakes play in influencing the population dynamics of their prey.

Nest predation involving a California kingsnake (Lampropeltis getula californiae) and two fledglings of least Bell's vireo (Vireo bellii pusillus) was observed along the San Luis Rey River in northern San Diego County, California. The kingsnake was discovered consuming a fledgling when the adults were observed mobbing the snake. The kingsnake then proceeded to systematically search for the second fledgling by traveling out and back from the nest multiple times over the next 15 min, each time in a new direction, traveling ca. ≤6–10 m before returning to the nest. The snake then found the original roosting branch of the surviving fledgling and proceeded quickly toward the fledgling that had moved ca. 15 m away. The kingsnake traveled this distance in ca. 2 min and killed and consumed the last fledgling. While California kingsnakes have been reported to feed on birds and bird eggs, this systematic searching behavior has not been described previously.

Most reptiles and mammals, with the exceptions of crocodilians, aquatic mammals and some primates, have a functional vomeronasal organ that detects and perceives semi-volatile chemicals in the environment. This organ is used in detection of prey and is also important for recognition of conspecifics and potential predators. We tested eight species of North American pit vipers for behavioural responses to an ophiophagous (snake-eating) predator, the common kingsnake, Lampropeltis getula. Kingsnakes have a substance in their skin that is recognized by crotalines, which react with a series of defensive responses including, but not limited to, avoidance, fleeing, body bridging and head hiding. The vomeronasal duct of the pit vipers was sutured closed to determine the role of this organ in detection of kingsnakes. Pit vipers with intact and sutured vomeronasal ducts were tested in a neutral cage with a kingsnake and monitored for behavioural responses. Results demonstrated that the vomeronasal organ is important in the recognition of kingsnakes by pit vipers and raises doubts that any other sense plays a major role in this behaviour. Copyright 1999 The Association for the Study of Animal Behaviour.

Summary Cases of diphtheritic necrotizing gastritis in a Rosy Boa (Lichanura trivirgata) and of necrotizing tracheitis in a double-headed Kingsnake (Lampropeltis hondurensis), both associated with Salmonella arizonae, are described. An immuno-peroxidase technique indicated that S. arizonae played a role in the causation of the lesions. In addition, the study showed the value of the techique for the detection of S. arizonae in the tissues of infected snakes.

When confronted by an ophiophagous (snake-eating) kingsnake, venomous snakes of the subfamily Crotalinae exhibit a suite of defensive responses including head hiding, thrashing, and an unusual response termed 'body bridging'. Other responses observed, such as biting and 'freezing', are more general in nature and can occur in a variety of contexts. Various crotalines of differing sizes were tested for their responses to kingsnakes (Lampropeltis getulus). Responses of individuals were recorded for up to 18 months. The results indicate that, if habituation can be overcome by periodically allowing a kingsnake to confront but not harm the crotaline, the response is dependent on the size of the crotaline, in that smaller specimens (0.9 m) respond readily, while larger snakes (1.0 m) tend not to respond. The size of the kingsnake apparently does not have an effect on the crotaline response. These data appear to resolve apparent conflicts in the literature regarding whether certain species respond to ophidian ophiophages. In addition, hexane extracts of kingsnake skin were fractionated using an alumina column. The various fractions obtained were tested to determine which elicited the defensive response. Activity was found only in the most non-polar fraction. Preliminary analysis by gas chromatography/mass spectrometry indicated that this fraction contained straight and branched, saturated and polyunsaturated long-chain hydrocarbons.

We examined the role of tongue extrusions for the detection of chemicals in the iguanid lizard Sceloporus jarrovi , prior to the breeding season. Most extrusions resulted in the tongue touching the substrate and are referred to as tongue-touches. A total of 2819 tongue-touches were observed for four age/sex classes of lizards. We could not demonstrate that the tongue is involved in the detection of a natural predator, the Arizona kingsnake ( Lampropeltis pyromelana ), and it was clear that it was not used to find food. It did, however, seem to be involved in the exploration of temporally and spatially unfamiliar areas. The possibility that the tongue is involved in the detection of conspecifics is discussed for this study, as well as for complementary studies. Possible marking attempts in the form of chin wipes, pelvic rubs, and defaecations were examined. The latter two possibilities were not important at this time of year. Chin wiping may result in the depositing of chemical markers, although no external chin glands were found and the possibility is still uncertain.

A total of 21 new taxa of New World pit vipers (Serpentes: Crotalinae) responded by elevating the middle portion of the body in a defensive posture (body bridge) when exposed to the skin substances of certain colubrid snakes (Colubridae). Newborn snakes from two of the three species tested gave the response. Several new species of colubrid snakes also are documented as capable of eliciting a response, and it is suggested that the termophiophage defensive response be used to denote body bridging and associated defensive behaviors instead of the restrictive “kingsnake defense posture.” Most of the snakes which elicit the response in crotaline snakes are known to feed on lizards and/or snakes. There is no apparent correlation between the stimulus snakes' ability to elicit a response in the crotaline snakes and sympatry with the crotaline snakes.

A female snake chosen on the basis of her previously determined blood protein genotype was mated on successive days to 2 males similarly selected. Electrophoretic analysis of 6 young showed that each male had sired 3 off-spring, thereby providing an unequivocal demonstration of multiple insemination.

Naive garter snakes, Thamnophis (Colubridae), were tested for responses to skin chemicals from ophiophagous snakes. Thamnophis elegans emitted a significantly greater number of tongue flicks to swabs rubbed against the skin of the kingsnake Lampropeltis getulus, an ophiophagous species, than to those from some generally nonophiophagous snakes or to blank swabs. Thamnophis sirtalis emitted more tongue flicks to swabs from the black racer Coluber constrictor, another ophiophage, than to control-snake or blank swabs. Olfactometric studies show that T. sirtalis is sensitive to airborne chemical cues from Lampropeltis, emitting more tongue flicks to the scent of this snake than to the scent of the rat snake Elaphe obsoleta or to plain air. The results from this study suggest that some of the same snake species recognized as predators by crotaline snakes are recognized as such by Thamnophis.

The eggs and hatchlings of the mole kingsnake, Lampropeltis calligaster rhombomaculata, are described. Various aspects of reproduction are reported, including incubation time, copulation, and courtship behavior in which the female played the active role. The mole kingsnake, Lampropeltis calligaster rhombomaculata, is a medium-sized form occupying a variety of habitats from Baltimore, Maryland, south through the Florida panhandle and west into Mississippi and Tennessee (Conant, 1975). This subspecies has been known for over 100 years, yet very little has been reported concerning reproduction, probably because of its secretive nature and infrequent collection. Brimley (1943) noted that the eggs of this subspecies were much like those of other kingsnakes and remarked on their adherent nature, but he failed to give details concerning clutch and egg size. Grogan and Prince (1971) reported on a group of nine apparent hatchlings plowed up in Maryland on 27 August, and cited one additional reproductive record also from Maryland (Howden, 1946). Subsequent accounts have not provided reproductive data, although hatchlings and juveniles have been described and photographed (Conant, 1975; Mount, 1975). A moderate amount of data are available on the nominate form, L. c. calligaster, from much of its range (see Fitch, 1970, 1979; Wright and Wright, 1957, for a review). Nevertheless, this appears to be the first account describing copulation, egg-laying and hatching in L. c. rhombomaculata. Certain aspects of reproduction observed by us differ from previous accounts on colubrids and are noted. This content downloaded from 157.55.39.35 on Fri, 02 Sep 2016 05:54:46 UTC All use subject to http://about.jstor.org/terms VOLUME 83, NUMBER 2 67 Fig. 1. Adult specimens of Lampropeltis calligaster rhombomaculata. MATERIALS AND METHODS Two Georgia specimens from Clayton County were available (Fig. 1): a male (total length 894 mm, weight 260 g) received as an adult on 1 July 1976, and a captive-hatched (summer, 1975) female (total length 617 mm, weight 84.1 g after oviposition) received 4 April 1976. The snakes were housed individually in various 1 or 2 ft fiberglass enclosures (Hulsey, 1973) both on and off exhibit at the Fort Worth Zoological Park, and were fed on laboratory mice. Temperatures varied from approximately 20'C in winter to 300C in summer. Artificial lighting was provided by 30-watt Vita-lite? fluorescent tubes above the cages from 0800-1730 h daily. Clear skylights provided a natural photoperiod for the Fort Worth, Texas area. The female was isolated prior to oviposition and provided with a nesting box filled with damp sphagnum moss. Eggs were measured with a vernier caliper, weighed on a triple-beam balance, and incubated in a vermiculite medium (Tryon, 1975) at 23-320C. Adults and neonates were measured for snout-vent length and total length using the squeeze-box technique (Quinn and Jones, 1974). Copulation was recorded on 35 mm color transparencies. This content downloaded from 157.55.39.35 on Fri, 02 Sep 2016 05:54:46 UTC All use subject to http://about.jstor.org/terms 68 TRANSACTIONS OF THE KANSAS ACADEMY OF SCIENCES

Eimeria lampropeltis sp. n., Caryospora lampropeltis sp. n., and Cryptosporidium lampropeltis sp. n. are described from the host Lampropeltis c. calligaster, and Eimeria zamenis is redescribed from this host and from L. getulus holbrooki. Oocysts of E. lampropeltis are ellipsoidal with a smooth outer wall measuring 29 to 38 by 14 to 20 t, mean 30.4 by 16.3. E. zamenis oocysts are ellipsoidal with a smooth outer wall measuring 28 to 33 by 14 to 19 u, mean 30.2 by 16.2. Oocysts of Caryospora lampropeltis are round with a finely pitted outer wall measuring 20 to 25 /t in diameter, mean 23.3. Each oocyst contains one thick-walled sporocyst with a prominent Stieda body and eight bananashaped sporozoites. Cryptosporidium lampropeltis oocysts are ovoid with a thin, smooth outer wall measuring 10 to 12 by 8 to 9 Iu, mean 10.9 by 8.1. Each oocyst contains four sporozoites which measure 6 to 8 by 1 to 2 IL, mean 7.0 by 1.9. Two kingsnakes, Lampropeltis c. calligaster, the prairie kingsnake, and L. getulus holbrooki, the speckled kingsnake, were captured in Jackson County, Illinois, in April 1965, and September 1966, respectively. Shortly after capture, they were transported to Fort Collins, Colorado, and kept in cages. Fecal examinations made in 1966 and 1967 showed the presence of oocysts of four species of coccidia. One of these, Eimeria zamenis, has not been adequately described (Phisalix, 1921; Roudabush, 1937), and the other three appear to be new

some that succumbed) quivered and twitched recurrently through the final 10 to 15 minutes of the chilling; and all of five others, when chilled 4 minutes at 10, continued to react to smart tapping on their container and spontaneously resumed breathing' in 10 minutes while kept at 4 to 50C. Divergent results, viz. prolonged primary chill coma, obtained with Leptocottus armatus by Doudoroff (1945), may perhaps be accounted for, equally in accord with the suggestion offered above, by acclimatization to warmer water (which his brief footnote does not disaffirm). Facilities were generously provided at the Lerner Marine Laboratory, a field station of the Department of Fishes and Aquatic Biology of The American Museum of Natural History. I am grateful to the staff for assistance and to all concerned for the hospitality shown me; I wish to thank especially the director, Dr. C. M. Breder, Jr.

HEATWOLE, H., E. CAMERON AND G. J. W. WEBB. 1971. Studies on anuran water balance-II. Desiccation in the Australian frog, Notaden bennetti. Herpetologica 27:365-378. HEUSSER, H. 1969. Die Lebenweise der Erdkrbte, Bufo bufo (L.); das Orientierungsproblem. Rev. suisse Zool. 76:443-518. HUBBS, C., AND F. D. MARTIN. 1967. Bufo valliceps breeding in artificial pools. Southwest. Natur. 12:105-106.