Evolution of coloration, urotomy and coral snake mimicry in the snake genus Scaphiodontophis (Serpentes: Colubridae)

The Neotropical hinged-tooth, coral snake mimics of the genus Scaphiodontophis are characterized by extremely long and disproportionately thick tails that are extremely fragile. Both the coloration and tail structure are putative antipredator devices. While all examples have components of the coloration that match those of the venomous coral snakes (family Elapidae), the range of variation is extreme, leading to controversy on the status of various populations, including nine named taxa. Individual, ontogenetic and geographic variation in scutellation and head, body and tail coloration were analysed to evaluate population status and possible evolutionary trends based on a sample of 183 examples from Mexico, Central America and Colombia. Variation in subcaudal counts show population differences (higher in Mexico and upper Central America) but are not congruent with geographic variation in coloration. Generally snakes from north of Nicaragua and from central and eastern Panama have a pattern of dyads (black-light-black bands separating red bands), those from Atlantic slope Nicaragua to western Panama a pattern of monads (light-black-light bands separating the red ones) and those from Colombia have both pattern types on the same snake. The dyads and/or monads may be present the length of the body and tail, restricted to the anterior part of the body or on the entire body or on the anterior part of the body and on the tail. Two or more of these variants may occur at a single geographic locality or only a single one may be present. Head and nuchal colour patterns (Z, A, V and Du) are relatively consistent geographically. The Adantic slope Guatemala, Belize and Honduras population have the A pattern, those of Nicaragua, Costa Rica and western Panama the V pattern, and those in Colombia a Du pattern. Other populations have the Z coloration. Intermediate conditions in coloration of the body and tail and head and neck are found at localities intermediate between the main pattern types, indicating intergradation among adjacent populations. Consequently, we regard these snakes as representative of a single species, Scaphiodontophis annulatus Dumeril and Bibron and the eight other names applied to various populations and individuals as synonyms. Analysis of colour pattern leads us to the conclusion that the tricolour pattern evolved from a uniform one through a lineate-spotted condition (usually present on the non-tricolour portions of the snake) through a bicolour red and black pattern to the dyadal condition. The monadal pattern in turn was derived from the dyadal one. The data further indicates that tricolour components first appeared anteriorly and progressively expanded posteriorly. The evolutionary sequence for the head and nuchal pattern appears to be A → Z → V → Du S. annulatus has a series of jaw and tooth specializations designed for rapid processing of hard-bodied prey found during diurnal foraging in the leaf-litter. Urotomy in this species involves intervertebral tail-breakage (pseudoautotomy) without regeneration. Evidence is presented supporting the long-tail multiple break hypothesis as applicable to Scaphiodontophis and other snakes with similar tail morphology (specialized pseudoautotomy). This is in contrast to snakes with similar tail morphology (specialized pseudoautotomy). This is in contrast to Coniophanes and other snakes with a high incidence of urotomy having long but unspecialized tails (unspecialized pseudoautotomy) without multiple breaks over time. All Scaphiodontophis colour patterns have a general resemblance to that of venomous coral snakes and offer protection from generalizing predators having innate or other triggered responses to coral snake colours. The aposematic effect is enhanced by tail thrashing and head twitching behaviours. The characteristic foraging pose of S. annulatus, which tends to expose the head and anterior body, makes even the incomplete tricolour pattern effective as an antipredator defence. No evidence supports the idea that tail thrashing or the incomplete tricolour pattern directs the predator attacks to the tail to expedite pseudoautotomy. Coral snake mimicry and specialized pseudoautotomy are shown not to be co-evolved and pseudautotomy seems to have evolved long before mimetic coloration in this genus.     

Understanding the formation of ancient intertropical disjunct distributions using Asian and Neotropical hinged-teeth snakes (Sibynophis and Scaphiodontophis: Serpentes: Colubridae)

Numerous taxa show ancient intertropical disjunct distributions. Many can be explained by well-known processes of historical vicariance, such as the breakup of Gondwanaland. Others, such as Asian–Neotropical divergences are not as well understood. To clarify the phylogenetic position and understand biogeographic and temporal origins of the geographically disjunct and morphologically unique genera of hinged-teeth snakes, Scaphiodontophis (n = 1) and Sibynophis (n = 9; Colubridae), we inferred a time-calibrated phylogeny with additional 107 taxa representing the superfamily Colubroidea using four genes (c-mos, cyt-b, ND2, RAG-1; 3085 bp). We used this tree to estimate ancestral areas for the group. The results show that Scaphiodontophis is sister to Sibynophis, both originated in the late Eocene/Oligocene in Asia and likely dispersed through Beringia to the New World, but unlike other snake groups left no extant species in temperate North America. Current recognition of Scaphiodontophiinae renders Colubrinae paraphyletic, and we resurrect the previously named subfamily Sibynophiinae to encompass both genera and use the tribes Sibynophiini (Sibynophis) and Scaphiodontophiini (Scaphiodontophis) to highlight the geographically distinct areas occupied by these taxa. These results suggest that intercontinental dispersal with extinction in intermediate areas can explain puzzling patterns of ancient intertropical disjunct distributions.     

Complete mitochondrial genome of the black-headed snake Sibynophis collaris (Squamata, Serpentes, Colubridae)

The black-headed snake Sibynophis collaris (Reptilia, Squamata, Colubridae) is a least concern species in the world. Two universal and two specific polymerase chain reaction (PCR) primers were used for long PCRs to amplify the whole mitochondrial genome of S. collaris. The products were subjected to do sequencing reactions. The complete genome is 17,163 bp in size, containing 37 genes coding for 13 proteins, 2 rRNAs, 22 tRNAs, and 2 control regions (CRI and CRII). The results could play an important role in the preservation of genetic resources for helping conservation of the endangered species.     

Hepatozoon species (Apicomplexa: Hepatozoidae) of the corn snake,Elaphe guttata (Serpentes: Colubridae) and the pigmy rattlesnake,Sistrurus miliarius barbouri (Serpentes: Viperidae) in south Florida

Hepatozoon guttata n. sp. is described from the corn snake (Elaphe guttata guttata) from south Florida. Gamonts average 14.6 x 4.6 (13-17 x 3.5-6) and are not recurved, with nucleus situated in the second quarter of the gamont. Erythrocyte cytoplasm rarely appears partially dehemoglobinized. The infected cells are usually distorted and are longer than the uninfected erythrocytes but do not differ in width; nuclei are smaller in length and width than those of uninfected cells. Sporogony in Aedes aegypti occurs within the head and the thorax but rarely in the abdomen. Oocysts are spherical to usually ovoid, 87.2 x 75.9 (45-155 x 40-152.5) and contain on average 7.1 (2-32) sporocysts. Sporocysts are spherical to ovoid, 34.8 x 31.0 (20-55 x 17.5-47.5), with 45.7 (14-89) sporozoites contained within. The pigmy rattlesnake (Sistrurus miliarius barbouri) in south Florida is parasitized by H. sistruri n. sp. Gamonts average 14.1 x 5.6 (12.6-15.8 x 4.7-6.3) in breadth and are not recurved, with the nucleus always situated in the second quarter of the gamont commonly at the midbody. Erythrocyte cytoplasm is not dehemoglobinized. The infected cells are always distorted and similar in length to the uninfected cells but with greater width and enlarged nuclei. Sporogony in A. aegypti occurs within head, thorax, and abdomen. Oocysts are spherical to usually ovoid, 163.6 x 154.7 (92-245 x 82-240) and contain 27.4 (12-42) sporocysts. Sporocysts are round to usually ovoid, 39.7 x 33.5 (25-50 x 20-50), with 45.7 (19-70) sporozoites.     

A new species of snake of the genus Sibon (Serpentes: Colubridae) of the Caribbean slope of Costa Rica

Sibon lamari, a new species of colubrid snake from northeastern Costa Rica is described on the basis of six specimens. The new form differs from the closely allied S. annulatus in color pattern, head size, subcaudal count, and number of labials bordering eye-orbit. This is the third species of Sibon, together with S. argus and S. longifrenis, possessing a green dorsal ground color in Costa Rica. Sibon lamari coexists sympatrically with S. annulatus, S. argus. S. longifrenis and S. nebulatus at the locality of Guayacán de Siquirres (Limón Province).     

Sense-organ-like parietal pits found in Psammophiini (Serpentes,Colubridae)

In several psammophine snake species, small sense-organ-like pits have been discovered occurring on the top of the head. These pits show a heretofore non-described structure. In numbers of one to four per head, they are present in Dromophis lineatus, Malpolon monspessulanus, several Psammophis species and Rhamphiophis rubropunctatus, but not in all individuals which is unrelated to sex. The reason for their sporadic occurrence remains obscure. Their equally obscure function is discussed in relation to the presumed functions of already known upper-head 'pits' in snakes. A histological study is in preparation.     

莽山后棱蛇Opisthotropis cheni Zhao, 1999的再描述(Serpentes:Colubridae)

作者于1999年根据湖南宦章莽山标本发表新种莽山后棱蛇Opisthotropis cheni的鉴别特征,现对该蛇加以详细描述。莽山后棱蛇足小形水栖蛇,全长500～600mm,尾长占全长的五分之一左右,身体圆柱形,头较小,与颈区分不明显。鼻间鳞前端窄,鼻孔背侧位;眼小,背侧位;前额鳞单枚,甚宽,其两外侧扩展至头侧;颊鳞长,后端人眶;无眶前鳞,眶后鳞2;颞鳞1＋2（1）;上唇鳞8（3-2-3）为主;下唇鳞10,前4或5枚接前颔片;颔片2对,前对约为后对长的2倍。背鳞通身17行,除最外行其余微棱,腹鳞159～167．肛鳞二分,尾下鳞55～59对。背面暗橄榄褐色,体侧具多数淡黄色横斑,左右横斑彼此交错或在背中线相连;腹面色白或偶有灰点斑。雄性头腹前端鳞片（如颏鳞、颔片和前4～5对下唇鳞）上有多数疣粒,雌性则无。此蛇见于山溪或小河流水中,白昼隐藏于岩洞、石下、沙砾或杂草间,晚上或黄昏外出活动觅食,共采到5号标本,其叶14号均采于晚上20～22点之间,1号采于18点左右;被捕捉的蛇离水后两三天就死掉。此蛇也曾见于农耕地灌溉渠旁的杂草中。 山     

温泉蛇属Thermophis一新种——四川温泉蛇

From 2004 to 2006, several field trips in western Sichuan, China were carried out. A number of Thermophis snakes were collected in Litang County. After detailed morphological studies, we found that these Thermophis specimens were much different from those from Xizang (Tibet). We believed that Litang snakes should be a new species and describe it as below.     

Geographical variation in the genus Dendrelaphis (Serpentes: Colubridae) within the islands of south-eastern Indonesia

The south-eastern islands of Indonesia between Bali in the west and Timor in the east define a region that is at the interface of the Asian and Australasian biogeographic regions. Three subspecies of the colubrid snake Dendrelaphis pictus have previously been recognized from the region: Dendrelaphis pictus pictus (Bali and Lombok); D. p. inornatus (Sumba to Timor); and D. p. intermedius (Sumbawa to Alor). A re-examination of Dendrelaphis pictus in this region uses multivariate statistics to examine the geographic variation in head-shield and body measurements within the species. The findings re-interpret the taxonomy, indicating that D. pictus from Bali and Lombok are specifically different from D. inornatus on the remaining islands on the Banda Arc. The populations of D. inornatus on the islands of Sumbawa, Sumba, Sawu, Flores, and Lembata are subspecifically different from those on Roti, Semau, Timor, Alor, and Wetar.
Several head-shield measures are strongly correlated with longitude within D. inornatus , suggesting environmentally triggered selection gradients. However, for many variables, the two subspecies differ significantly in the direction of the trends which suggests the subspecies respond differently to these environmental gradients. There is evidence that Dendrelaphis individuals may be transported through the islands by anthropogenic agencies.     

Population genetic structure and geographic differentiation in the hot spring snake Thermophis baileyi (Serpentes, Colubridae): indications for glacial refuges in southern-central Tibet

The endangered hot spring snake (Thermophis baileyi) is endemic to the Tibetan Plateau and restricted to a few sites at high altitudes (>3500 m a.s.l.). Its strong preference for habitats with hot springs might be an early adaptation to the cooling climate during the plateau uplift. Some of these thermal sites may have been free of ice during the last glacial maximum (LGM) serving as refuges for the snake. To test this and to examine the population genetic structure data of 12 microsatellites and mitochondrial DNA were obtained from 153 individuals from 12 sites across the plateau. Four of the microsatellite loci were newly developed for this study. The results suggest that T. baileyi has at least two genetically diverse clades in Tibet, which developed during the Pleistocene and expanded after the LGM. Accordingly, the existence of separate glacial refuges on the central plateau can be assumed. Analyses of the genetic variation indicated a high level of geographic differentiation and population structure on a regional as well as on a range-wide scale. The study shows that, apart from the phylogeographic signatures, the diversification of current Thermophis populations is caused by (a) a limited dispersal due to mountain ranges, (b) a strong preference for hot springs and (c) the insular distribution of suitable habitats on the plateau. The present data should be taken into account in necessary monitoring and management plans for T. baileyi.     

Cryptic diversity in a Eurasian water snake (Natrix tessellata, Serpentes: Colubridae): Evidence from mitochondrial sequence data and nuclear ISSR-PCR fingerprinting

The dice snake, Natrix tessellata (Laurenti, 1768), is a suitable study organism to address questions of Eurasian phylogeography due to its wide Palearctic distribution. We analysed complete mitochondrial cytochrome b sequences and nuclear ISSR-PCR fingerprints of more than 300 specimens representing nearly the entire geographic range. Nine major mitochondrial lineages were discovered based on mtDNA sequences. The three most basal lineages comprised populations from Iran, Jordan–Egypt, and Greece, respectively. Other lineages were associated with samples from the Turkish peninsula, the Caucasus, the Aral Sea, and eastern Kazakhstan. A sister-group relationship was found between two lineages from Crete and the European mainland. Assuming an evolutionary rate of 1.35% sequence divergence per million years, among-lineage p-distances of 1.7–8.4% suggest that intraspecific differentiation might date back as far as the Miocene/Pliocene transition 5–6 million years ago. The pattern of genetic differentiation in mitochondrial phylogeny with regard to Asia Minor and the region of the Aral Sea was not congruent with the results of the nuclear ISSR-PCR analyses, and suggests admixing within some mtDNA clades at contact zones. The taxonomic implications of the high intraspecific variation in the dice snake are discussed.     

八种蛇(蛇亚目:游蛇科:颈槽蛇属)的半阴茎形态比较

蛇类半阴茎形态在研究蛇类的形态进化、分类和系统等方面具有重要的作用。对分布于我国境内的颈槽蛇属八种蛇(虎斑颈槽蛇、海南颈槽蛇、颈槽蛇、九龙颈槽蛇、缅甸颈槽蛇、黑纹颈槽蛇、喜山颈槽蛇和红脖颈槽蛇)的半阴茎进行详细描述和比较,并以此探讨部分物种的有效性和推断它们之间的进化及系统关系。     

A new genus and species of opisthogonimid (Digenea) of Liophis anomalus (Serpentes: Colubridae) from Uruguay

During a study on the helminth fauna of Liophis anomalus specimens deposited in the Reptile Collection of the Secci6n Zoología Vertebrados, Facultad de Ciencias, Uruguay, a new genus of Opisthogoniminae was found. Paracotyletrema n. gen. is described from the buccal cavity and esophagus of specimens from Departamento de Flores, Uruguay. This new genus differs from the other 2 Opisthogoniminae, Opisthogonimus and Westella, by having much smaller body size; maximum width at level of posterior end of oral sucker; oral sucker length more than one-fourth of body length; oral sucker and acetabulum separated by a distance of about the diameter of pharynx; cirrus pouch length more than half of body length; and testes in completely coincident zones. The type and only species, Paracotyletrema poncedeleoni n. gen., n. sp., is defined by the following diagnostic characters: mean oral sucker length one-third to one-half of body length; cirrus pouch of about body length, contorted, mainly acetabular; large, round testes in nearly adjacent fields, and an ovary nearly adjacent to right testis. Besides the type locality, Paracotyletrema poncedeleoni has been found in L. anomalus specimens from the Departamentos of Artigas and Treinta y Tres; therefore, it has a widespread distribution in Uruguay.     

Feeding behavior and venom toxicity of coral snake Micrurus nigrocinctus (Serpentes: Elapidae) on its natural prey in captivity

The feeding behavior and venom toxicity of the coral snake Micrurus nigrocinctus (Serpentes: Elapidae) on its natural prey in captivity were investigated. Coral snakes searched for their prey (the colubrid snake Geophis godmani) in the cages. Once their preys were located, coral snakes stroke them with a rapid forward movement, biting predominantly in the anterior region of the body. In order to assess the role of venom in prey restraint and ingestion, a group of coral snakes was 'milked' in order to drastically reduce the venom content in their glands. Significant differences were observed between snakes with venom, i.e., 'nonmilked' snakes, and 'milked' snakes regarding their behavior after the bite. The former remained hold to the prey until paralysis was achieved, whereas the latter, in the absence of paralysis, moved their head towards the head of the prey and bit the skull to achieve prey immobilization by mechanical means. There were no significant differences in the time of ingestion between these two groups of coral snakes. Susceptibility to the lethal effect of coral snake venom greatly differed in four colubrid species; G. godmani showed the highest susceptibility, followed by Geophis brachycephalus, whereas Ninia psephota and Ninia maculata were highly resistant to this venom. In addition, the blood serum of N. maculata, but not that of G. brachycephalus, prolonged the time of death of mice injected with 2 LD(50)s of M. nigrocinctus venom, when venom and blood serum were incubated before testing. Subcutaneous injection of coral snake venom in G. godmani induced neurotoxicity and myotoxicity, without causing hemorrhage and without affecting heart and lungs. It is concluded that (a) M. nigrocinctus venom plays a role in prey immobilization, (b) venom induces neurotoxic and myotoxic effects in colubrid snakes which comprise part of their natural prey, and (c) some colubrid snakes of the genus Ninia present a conspicuous resistance to the toxic action of M. nigrocinctus venom.     

Reproduction in the yellow-bellied sea snake (Pelamis platurus) from Panama: Field and laboratory observations

Little is known of the reproductive biology of the yellow-bellied sea snake (Pelamis platurus), a species widely distributed in the Indo-Pacific and eastern Pacific Oceans. We observed mating, birth, and free-ranging neonates of P. platurus while collecting this snake once a month over 19 months in the Gulf of Chiriquí, Panama. A pair of copulating snakes was netted on the water surface during February. Neonates, which were identified by size, were observed from September to December. Captive females gave birth during September. Neonates born in captivity emerged head- or tailfirst, shed the remnants of the fetal membranes by coiling their body in a circular loop, and then surfaced to breathe. © 1996 Wiley-Liss, Inc.     

New species of Haplometroides (Digenea: Plagiorchiidae) from Phalotris nasutus (Gomes, 1915) (Serpentes, Colubridae)

A new species of Haplometroides (Digenea, Plagiorchiidae) is described from a specimen of Phalotris nasutus (Gomes, 1915) (Serpentes, Colubridae). The host snake was obtained in the municipality of Corumbd, Mato Grosso do Sul State, Brazil. Trematodes were recovered from esophagus, stomach, and small intestine of the host. The main characteristic of the new species is the vitellaria, which is intercecal, cecal, and extracecal in the preacetabular region. A key for identification of the species in Haplometroides is proposed.     

铅色水蛇Enhydris plumbea(Boie, 1827) 的生物学初探(蛇亚目:游蛇科:仰鼻蛇亚科)

根据野外观察和解剖标本, 本文报道蛇亚目、游蛇科、水游蛇亚科中国南方常见优势种铅色水蛇的生物学研究初步结果.1.长度112号雄蛇全长范围286 ～466 mm,151号雌蛇全长范围306～504 mm.海南陵水103号雄蛇最大全长358+54=412 mm,137号雌蛇最大全长443+61=504 mm.2.鳞片的变异根据299号标本观察,数量恒定无变异的鳞片有吻鳞(1), 鼻间鳞 (1), 前额鳞 (2), 额鳞 (1), 顶鳞 (2), 鼻鳞 (每侧1), 颊鳞 (每侧1), 眶上鳞(每侧1), 前颞鳞 (每侧1), 颏鳞 (1), 颔片 (2 对), 背鳞 (19-19-17 行), 肛前鳞 (完整);根据鳞片有变异的67号标本观察, 可出现变异鳞片的变异范围及其频率眶前鳞1(1→2,4.5%),眶后鳞2(2→1,3.0%),后颞鳞2(2→1,1.5%),上唇鳞3-2-3(3-1-4,0.75%),下唇鳞10(9,0.75%;11,2.25%);腹鳞与尾下鳞则绝对有变异腹鳞雄性112号变异幅度124～136 (平均 128.2), 雌性151号变异幅度123～132 (平均 126.1),尾下鳞雄性107号变异幅度36～41 (平均 39.7)对,雌性149号变异幅度31～36 (平均 33.7)对.3.两代之间变异的消失和出现两例母蛇的右侧眶前鳞均变异为2,她们的全部仔蛇21号均正常为1;一例母蛇的下唇鳞为正常的10枚,而她的仔蛇6号中有两号分别有一侧或两侧变异为9.4.垂直分布范围沿海低地到海拔985 m.5.栖息环境各种水域,以静水水域为主,在海南陵水县吊罗山大里乡以灌水后的稻田为主,极少数发现在静水水域或流动缓慢的灌溉渠内.6.摄食和食性每年4月开始摄食,6月为摄食高峰,9月停止;食物以小形蛙类和蝌蚪为主,其次为小鱼.7.繁殖习性二级性征主要反映在雄性尾的比例较长,尾下鳞的数目相应较多;二级性比(出生时的性比)接近11,三级性比在居群较大的情况下也约为11,而在居群较小的情况下统计约为0.51;雌性性成熟期(以开始产仔为依据)全长都在350 mm以上;解剖28号孕雌得知输卵管卵数3～12(平均6.3)枚;卵在母体内发育进程解剖41号成熟雌性得知4月卵尚未发育,5月及6月处于发育中或晚期、产仔或已产出,7月基本都已产毕;统计两窝21仔得知初产出仔蛇全长127～139 mm,母蛇体长大者其仔蛇亦较长;统计同窝9仔得知初生仔蛇重量2.4～2.5 g,平均2.47 g.本文是作者早期未发表的工作,现整理供对蛇类有兴趣的青年同行参考.本研究所用全部标本(包括3号雌蛇及其刚产出的仔蛇) 均保存在中国科学院成都生物研究所两栖爬行动物标本馆.