Search

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.

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.

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

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.

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.

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.

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.

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

tions of the vessels of the human skin to cold. Heart 15:177-208. 1941. Observations on some normal and injurious effects of cold upon the skin and underlying tissues. I. Reactions to cold and injury of normal skin. Brit. Med. J. 20:795-7. MICHEEL, F. AND F. JUNG. 1936. Zur Kenntnis der schlangengifte. Ztschr. f. Physiol. Chem. 239:217-30. ORNDOFF, B. H. 1915. Practical considerations of snake poisoning. Med. Brief. 43:231-8. PARRISH, H. M. 1957. The poisonous snake bite problem in Florida. Quart. J. Florida Acad. Sci. 20:185-204. RUSSELL, F. E. 1960a. Rattlesnake bites in Southern California. Am. J. Med. Sci. 239:1-10. 1960b. Snake venom poisoning in Southern California. Calif. Med. 93:347-50. SLOTTA, C. H. AND G. SZYSZKA. 1937. Estudos chimicos sobre os venenos ophidicos: I. Determinacao de sua toxicidade em camondon