Behavioral thermoregulation and the role of melatonin in a nocturnal snake

Daily and seasonal variations in hormone levels influence the complex interactions between behavior and physiology. Ectothermic animals possess the unique ability behaviorally to adjust body temperature (T(b)) to control physiological rate processes. Thus, a hormone may indirectly influence a physiological rate by directly influencing the behaviors that adjust or control that rate process. Although many hormonal influences on behavioral regulation of T(b) remain uninvestigated, melatonin (MEL) generally is considered a hormone that decreases mean preferred T(b). Many ectotherms demonstrate the selection of lower T(b)'s in response to increased MEL concentrations. Here, we examined the influence of MEL on the behavioral regulation of T(b) in the nocturnal African house snake Lamprophis fuliginosus. A series of experiments with two injection regimes of MEL had no significant effect on the mean preferred T(b) of L. fuliginosus. In addition, mean preferred T(b)'s during the photophase did not differ significantly from those during scotophase. Our findings suggest that L. fuliginosus does not respond to elevated concentrations of either endogenous or exogenous MEL. To verify that the African house snake is nocturnal, we investigated activity patterns of L. fuliginosus throughout the photoperiod. The activity period of L. fuliginosus occurs in the scotophase of the photoperiod, a pattern consistent with that of nocturnal species. This suggests that nocturnal organisms such as L. fuliginosus may not respond to MEL in the same manner as many diurnal species. Our results support the hypothesis that some animals, particularly nocturnal species, may have developed alternative responses to increased plasma concentrations of MEL.     

A developmental staging series for the African house snake, Boaedon (Lamprophis) fuliginosus

Embryonic staging series are important tools in the study of morphological evolution as they establish a common standard for future studies. In this study, we describe the in ovo embryological development of the African house snake (Boaedon fuliginosus), a non-venomous, egg-laying species within the superfamily Elapoidea. We develop our staging series based on external morphology of the embryo including the head, eye, facial prominences, pharyngeal slits, heart, scales, and endolymphatic ducts. An analysis of embryonic growth in length and mass is presented, as well as preliminary data on craniofacial skeletal development. Our results indicate that B. fuliginosus embryos are well into organogenesis but lack well-defined facial prominences at the time of oviposition. Mandibular and maxillary processes extend rostrally within 8 days (stage 3), corresponding to the first appearance of Meckel's cartilages. Overall, the development of the craniofacial skeleton in B. fuliginosus appears similar to that of other snake species with intramembraneous bones (e.g., dentary and compound bones) ossifying before most of the endochondral bones, the first of which to ossify are the quadrate and the otic capsule. Our staging series is the first to describe the post-ovipositional development of a non-venomous elapoid based on external morphology. This species is an extremely tractable captive that can produce large clutches of eggs every 45 days throughout the year. As such, B. fuliginosus should be a good model for evolutionary developmental biologists focusing on the craniofacial skeleton, loss of limbs, generational teeth, and venom delivery systems.     

Molecular systematics of the African snake family Lamprophiidae Fitzinger, 1843 (Serpentes: Elapoidea), with particular focus on the genera Lamprophis Fitzinger 1843 and Mehelya Csiki 1903

The snake family Lamprophiidae Fitzinger (Serpentes: Elapoidea) is a putatively Late Eocene radiation of nocturnal snakes endemic to the African continent. It incorporates many of the most characteristic and prolific of Africa's non-venomous snake species, including the widespread type genus Lamprophis Fitzinger, 1843 (house snakes). We used approximately 2500 bases of mitochondrial and nuclear DNA sequence data from 28 (41%) of the approximately 68 recognised lamprophiid species in nine of the eleven genera to investigate phylogenetic structure in the family and to inform taxonomy at the generic level. Cytochrome b, ND4 and tRNA gene sequences (mitochondrial) and c-mos sequences (nuclear) were analysed using Maximum Likelihood, Bayesian Inference and Maximum Parsimony methods. The genus Mehelya Csiki, 1903 was paraphyletic with respect to Gonionotophis Boulenger, 1893. To address this, the concept of Gonionotophis is expanded to include all current Mehelya species. The genus Lamprophis emerged polyphyletic: the enigmatic Lamprophis swazicus was sister to Hormonotus modestus from West Africa, and not closely related to its nominal congeners. It is moved to a new monotypic genus (Inyoka gen. nov.). The remaining Lamprophis species occur in three early-diverging lineages. (1) Lamprophis virgatus and the widely distributed Lamprophis fuliginosus species complex (which also includes Lamprophis lineatus and Lamprophis olivaceus) formed a clade for which the generic name Boaedon Duméril, Bibron & Duméril, 1854 is resurrected. (2) The water snakes (Lycodonomorphus) were nested within Lamprophis (sensu lato), sister to Lamprophis inornatus. We transfer this species to the genus Lycodonomorphus Fitzinger, 1843. (3) We restrict Lamprophis (sensu strictissimo) to a small clade of four species endemic to southern Africa: the type species of Lamprophis Fitzinger, 1843 (Lamprophis aurora) plus Lamprophis fiskii, Lamprophis fuscus and Lamprophis guttatus.     

On the taxonomy of Littorina africana (Mollusca: Gastropoda)

L. africana and L. knysnaensis are regarded as two morphs of a single species which exhibits a genetic cline along the south-eastern coast of southern Africa. The dark brown morph knysnaensis dominates the western, cooler end of the cline and is replaced by the pale blue morph africana at the warmer end of the cline. These conclusions are based on evidence from the latitudinal distributions, the complete range of intermediate forms regarding shell colour and shell morphology and the lack of differences in redular morphology, penial morphology or habitat.     

Effects of feeding on luminal pH and morphology of the gastroesophageal junction of snakes

At the gastroesophageal junction, most vertebrates possess a functional lower esophageal sphincter (LES) which may serve to regulate the passage of liquids and food into the stomach and prevent the reflux of gastric contents into the esophagus. Snakes seemingly lack an LES and consume meals large enough to extend anteriorly from the stomach into the esophagus thereby providing the opportunity for the reflux of gastric juices. To explore whether snakes experience or can prevent gastric reflux, we examined post-feeding changes of luminal pH of the distal esophagus and stomach, the fine scale luminal pH profile at the gastroesophageal junction, and the morphology of the gastroesophageal junction for the Burmese python (Python molurus), the African brown house snake (Lamprophis fuliginosus), and the diamondback water snake (Nerodia rhombifer). For each species fasted, there was no distension of the gastroesophageal junction and only modest changes in luminal pH from the distal esophagus into the stomach. Feeding resulted in marked distension and changes in tissue morphology of the gastroesophageal junction. Simultaneously, there was a significant decrease in luminal pH of the distal esophagus for pythons and house snakes, and for all three species a steep gradient in luminal pH decreasing across a 3-cm span from the distal edge of the esophagus into the proximal edge of the stomach. The moderate acidification of the distalmost portion of the esophagus for pythons and house snakes suggests that there is some anterior movement of gastric juices across the gastroesophageal junction. Given that this modest reflux of gastric fluid is localized to the most distal region of the esophagus, snakes are apparently able to prevent and protect against acid reflux in the absence of a functional LES.     

A taxonomic reappraisal of the Daphnia hyalina complex (Crustacea: Cladocera): an experimental and ecological approach

The paper is a taxonomic study of the limnetic Daphnia populations of the English Lake District, the populations upon which the most popular taxonomic scheme is based (Johnson, 1952).
Johnson treats these populations as three varieties of the species D. hyalina, namely D. hyalina s. str., D.h. var. galeata and D.h. var. lacustris .
Careful examination of the populations of Esthwaite reveals that what has previously been regarded as a population of a single variety of D. hyalina. includes three distinct forms. Experimental work confirms that these are genetically distinct. One of these forms corresponds to D. hyalina var. galeata , whilst the other two are designated the "toothed" and "round-headed" forms.
The first part of the paper is concerned with the taxonomic status of var. galeata . Brooks (1957) and some later workers treat this as a separate species (D. galeata) on morphological grounds. This is fully supported by the present work. Differences between var. galeata and co-existing forms in incidence of sexual reproduction, in vertical distribution and in seasonal abundance and average brood-size provide further support for the separation of D. galeata.
The second part of the paper is concerned with D. hyalina s.str. which, on the basis of morphology, ecology and reproductive behaviour, is shown to be at least as distinct from other members of the complex as is D. galeata . It is argued that other forms should therefore to be removed from D. hyalina . The taxonomic status and affinities of the remaining "toothed" and "round-headed" forms are discussed.     

Evolutionary relationships and biogeography of Biomphalaria (Gastropoda: Planorbidae) with implications regarding its role as host of the human bloodfluke, Schistosoma mansoni.

The wide geographic distribution of Schistosoma mansoni, a digenetic trematode and parasite of humans, is determined by the occurrence of its intermediate hosts, freshwater snails of the genus Biomphalaria (Preston 1910). We present phylogenetic analyses of 23 species of Biomphalaria, 16 Neotropical and seven African, including the most important schistosome hosts, using partial mitochondrial ribosomal 16S and complete nuclear ribosomal ITS1 and ITS2 nucleotide sequences. A dramatically better resolution was obtained by combining the data sets as opposed to analyzing each separately, indicating that there is additive congruent signal in each data set. Neotropical species are basal, and all African species are derived, suggesting an American origin for the genus. We confirm that a proto-Biomphalaria glabrata gave rise to all African species through a trans-Atlantic colonization of Africa. In addition, genetic distances among African species are smaller compared with those among Neotropical species, indicating a more recent origin. There are two species-rich clades, one African with B. glabrata as its base, and the other Neotropical. Within the African clade, a wide-ranging tropical savannah species, B. pfeifferi, and a Nilotic species complex, have both colonized Rift Valley lakes and produced endemic lacustrine forms. Within the Neotropical clade, two newly acquired natural hosts for S. mansoni (B. straminea and B. tenagophila) are not the closest relatives of each other, suggesting two separate acquisition events. Basal to these two species-rich clades are several Neotropical lineages with large genetic distances between them, indicating multiple lineages within the genus. Interesting patterns occur regarding schistosome susceptibility: (1) the most susceptible hosts belong to a single clade, comprising B. glabrata and the African species, (2) several susceptible Neotropical species are sister groups to apparently refractory species, and (3) some basal lineages are susceptible. These patterns suggest the existence of both inherent susceptibility and resistance, but also underscore the ability of S. mansoni to adapt to and acquire previously unsusceptible species as hosts. Biomphalaria schrammi appears to be distantly related to other Biomphalaria as well as to Helisoma, and may represent a separate or intermediate lineage.     

On the variability of fire-bellied toads in the Far East

We studied external morphology and molecular genetics of two forms of the oriental fire-bellied toad, described as separate subspecies Bombina orientalis silvatica Korotkov, 1972, and Bombina orientalis practicola Korotkov, 1972. We have found significant differences between them in body size, coloration and some morphometric parameters but not in genetics. Taking into account their geographical separation, one can consider them as forest and meadow ecological morphs but not different taxa. Therefore, the names Bombina orientalis practicola Korotkov, 1972, and B. orientalis silvatica Korotkov, 1972, should be considered as junior synonyms of the name Bombina orientalis (Boulenger, 1980). We suppose that these morphs have been isolated from each other recently, and their morphological differences at genetic similarity are related to high plasticity of the species, when ecologically determined morphological differentiation precedes genetic differentiation.     

Phylogeography and speciation of colour morphs in the colonial ascidian Pseudodistoma crucigaster

Variation in pigmentation is common in marine invertebrates, although few studies have shown the existence of genetic differentiation of chromatic forms in these organisms. We studied the genetic structure of a colonial ascidian with populations of different colour morphs in the northwestern Mediterranean. A fragment of the c oxidase subunit 1 (COI) mitochondrial gene was sequenced in seven populations of Pseudodistoma crucigaster belonging to three different colour morphs (orange, yellow and grey). Maximum likelihood analyses showed two well-supported clades separating the orange morph from the yellow-grey morphotypes. Genetic divergence between these clades was 2.12%, and gamma(ST) values between populations of the two clades were high (average 0.936), pointing to genetic isolation. Nested clade and coalescence analyses suggest that a past fragmentation event may explain the phylogeographical origin of these two clades. Non-neutral mtDNA evolution is observed in our data when comparing the two clades, showing a significant excess of nonsynonymous polymorphism within the yellow-grey morphotype using the McDonald-Kreitman test, which is interpreted as further support of reproductive isolation. We conclude that the two clades might represent separate species. We compare the population genetic differentiation found with that estimated for other colonial and solitary ascidian species, and relate it to larval dispersal capabilities and other life-history traits.     

Evidence of genetic and reproductive isolation between two morphs of subtropical-dominant coral Acropora solitaryensis in the non-reef region of Japan

We reveal the existence of a cryptic species of Acropora solitaryensis ( Veron and Wallace, 1984 ), a dominant species in high-latitude coral communities. Although some morphs, such as arborescent table (AR), solid plate (PL), and intermediate (IM) forms, had been known in this species, it was unclear whether these are reproductively isolated from one another. Here, potential reproductive exchange between two representative morphs, AR and PL, were examined using genetic and reproductive methods. Molecular phylogenetic analyses using both mitochondrial and nuclear molecular markers (mitochondrial control region and mini-collagen intron, respectively) indicated that AR is clearly distinct from PL, suggesting that gene flow between the morphs is absent. In cross-fertilization experiments, gametic compatibility between AR and PL was extremely low, suggesting prezygotic isolation of these morphs. These results strongly suggest that AR and IM forms are variations of A. solitaryensis, whereas PL form may be an undescribed species. In addition, AR was closely related genetically to A. pruinosa, which is a high-latitude species with arborescent form, and AR and A. pruinosa were able to hybridize, although with lower fertilities than observed in intra-specific crosses. The two species are thus likely to have speciated not in tropical regions, but in non-reef regions due to habitat segregation.     

COLOR PATTERN POLYMORPHISM IN THE LAKE ERIE WATER SNAKE,NERODIA SIPEDON INSULARUM

Populations of the water snake, Nerodia sipedon, on islands in western Lake Erie are polymorphic for color pattern. These populations include banded, intermediate, and unbanded morphs while surrounding mainland populations consist solely of the banded morph. The hypothesis that this polymorphism is maintained by strong selection and migration pressures is widely accepted. Unbanded morphs are apparently more cryptic along island shorelines while banded morphs are more cryptic on the mainland. Migration of banded morphs from the mainland explains their persistence in island populations. Data collected in a capture-mark-recapture program on six islands provide no evidence of differential selection among morphs; morph frequencies do not differ among age classes, between once-captured and multiply-captured snakes, or between scarred and unscarred snakes. Furthermore, herring gulls, the most common snake predators in the island area, appear to detect banded and unbanded model snakes with equal ease. High site fidelity of water snakes and the distribution of morphs among islands suggest that migration from the mainland is not common. However, islands close to each other are similar in morph frequency, and water snakes have colonized islands elsewhere in the Great Lakes, indicating that some migration does occur. Recently, the frequency of banded morphs has increased in island populations while adult population sizes have declined. This increase in banded morphs is interpreted as reflecting an increased impact of migration from the mainland into these reduced populations. One scenario for the evolution and maintenance of this polymorphism is that selection was important in establishing unbanded morphs in island populations as they became isolated from the mainland. As populations declined to their present size, the impact of migration from the mainland increased and is now swamping the effect of selection. Further declines in island population size may result in fixation of the banded morph.     

Amphibians and squamates from the middle Eocene of Namibia,with comments on pre-Miocene anurans from Africa

Three middle Eocene localities (Silica North, Silica South, Black Crow) recently discovered in Namibia have produced terrestrial faunas that rank among the few known from the period of insulation of Africa (Aptian-early Miocene). Collectively, the three localities have yielded anuran amphibians (one pipid frog, the earliest assemblage [three taxa] of ranoid frogs in Africa, one indeterminate family) and squamate reptiles (an amphisbaenian ‘lizard’, a snake that likely represents a colubroid, and two indeterminate ‘lizards’). These Eocene faunas suggest that ranoids, colubroids and African pipids are autochthonous to Africa. However, whereas pipids are vicariants inherited from West Gondwana, ranoids and colubroids (if really autochthonous) originated in Africa from unknown stems. Silica North and Silica South correspond to aquatic environments, permanent fresh water being present in the first locality; the environment of Black Crow was drier.     

Risk of predation as a promoting factor of species divergence in threespine sticklebacks (Gasterosteus aculeatus L.)

Icelandic freshwater systems are geologically young and contain only six species of freshwater fish. As these species colonized Icelandic fresh waters they were presented with a diversity of unique, uncontested habitats and food resources, promoting the evolution of new behaviour strategies crucial to the formation of new morphs and speciation. To determine the likelihood that predation threat could affect the antipredator behaviour and possibly the sympatric divergence of prey populations, we analysed antipredator behaviour of seven groups of Icelandic threespine sticklebacks ( Gasterosteus aculeatus ): two marine groups, one group from a lake without piscine predators, and two polymorphic lake populations, each with two groups occupying unique habitats. Shoaling cohesion, school formation and duration, and vigilance in predator inspection/avoidance behaviour varied greatly among groups. The differences appeared to be related to the risk of predation as well as to opportunities and constraints set by the different habitats. Antipredator behaviour was especially pronounced and differed extensively in two polymorphic forms from the lake Thingvallavatn, where predation risk is very high. By keeping the two morphs separate in their respective habitats, high predation risk may be a contributing factor in promoting the habitat-specific divergence of G. aculeatus seen in the lake. This suggests that in situations where refuge habitats are spatially separated, the risk of predation may contribute to the evolution of separate sympatric forms of small fish such as G. aculeatus .  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 189–203.     

A comparison ofGalago moholi in South Africa withGalago zanzibaricus in Kenya

Two populations of small bushbabies that frequently have been regarded as subspecies of Galago senegalensisare compared and contrasted. The animals in South Africa differ from those in coastal forests of Kenya in aspects of their morphology, locomotion, calls, reproduction, diet, social organization, and karyotype. The two forms are clearly separate species, namely G. moholiand G. zanzibaricus.     

The Taxonomic Status and Intraspecific Differentiation of the Black Sea Horse Mackerel, <Emphasis Type="Italic">Trachurus mediterraneus ponticus</Emphasis> (Aleev, 1956) (Carangidae)

The taxonomic status of the horse mackerel, Trachurus mediterraneus ponticus (Aleev, 1956), which inhabits the Black Sea, has been clarified based on the analysis of variation in the nucleotide sequence of the mtDNA cytochrome b (Cyt-b) locus. A comparison of this species with the horse mackerel from the Mediterranean basin and the World Ocean has shown that the Black Sea horse mackerel is neither an independent species nor a subspecies and can only be qualified as the species Trachurus mediterraneus (Steindachner, 1868) distributed throughout the Mediterranean Sea basin. A hypothesis has been made about the possible cause of the origin and existence of two forms, “large” and “small”, of the horse mackerel. By using the technique of calculating ontogenetic trajectories, it has been found that these are two morpho-ecological forms, whose differentiation is most likely determined by the type of feeding.     

Postovipositional development of the sand snake Psammophis sibilans (Serpentes:Lamprophiidae) in comparison with other snake species

Characterizing and comparing developmental progress across different species helps to interpret how different or similar body forms evolved. We present an embryonic table for the oviparous African Sand Snake Psammophis sibilans from the Lamprophidae family, describing its postovipositional in ovo development. Psammophis is a good model of a genus that is widely distributed in Africa and Asia and includes 22 species. We describe ten embryonic stages based on the development of externally visible morphological characteristics such as; pharyngeal arches, facial processes, eyes, scales, body pigmentation and body colour pattern development. This study discusses the development of this snake and compares it with that of the closely related brown house snake Lamprophis fulliginosus (Lamprophidae) and the medically important venomous cobras Naja haje haje and Naja kaouthia from the sister lineage Elapidae. The distantly related basal snake Python sebae, which displays different morphology and behaviour, was chosen for deeper insight into the evolution of body structures within the snake clade. We found interspecific differences in the relative stage of development of embryonic structures at the time of oviposition and during postovipositional embryonic development. One of the outcomes of this study is that embryonic structures such as the pharyngeal processes, eye pigmentation and scales are interspecifically conservative in regard to timing of morphodifferentiation, while body pigmentation, colour and colour pattern are interspecifically plastic in their temporospatial development.     

The evolution of phally polymorphism in Bulinus truncatus (Gastropoda,Planorbidae): the cost of male function analysed through life-history traits and sex allocation

In the hermaphrodite freshwater snail Bulinus truncatus, two sexual morphs, euphallic (regular hermaphrodites) and aphallic individuals without a male copulatory organ, co-occur at various ratios in natural populations. Both aphallic and euphallic individuals can reproduce by selfing, but when outcrossing aphallic individuals can only play the female role. A comparison of life-history traits and sex allocation in these two forms provides the opportunity to investigate the evolution and maintenance of sexual polymorphisms. This study was performed to test whether a reallocation of resources from the lost male function to the female function occurs in aphallic snails at the level of both sex organs (sex allocation) and life-history traits. In a first experiment we compared life-history traits over a whole life-cycle under selfing between the two sexual morphs. In a second experiment, the sex organs were weighed to test for a difference in sex allocation between the two morphs. No difference in resource allocation to female function between the two morphs was observed in either experiment. This is in contrast to patterns frequently observed in sexually polymorphic plants, and in a previous study performed on aphally in the same snail species. We discuss the genetic and physiological hypotheses that could explain these results, and their consequences for the evolution and maintenance of phally polymorphism in B. truncatus.     

Polymorphism and speciation in Arctic charr

The Arctic charr Salvelinus alpinus exhibits 1–4 sympatric morphs in postglacial lakes, of which one or two are epibenthic zoobenthos feeders, one is a limnetic planktivore and one is a piscivorous form. In addition, northern rivers support partly migratory populations with anadromous and freshwater resident fish. The morphs vary in their coloration, morphology, life history, behaviour and genetic characteristics. The morphs usually differentiate according to their ontogenetic stage at maturity, which parallels paedomorphism in amphibians. The young usually start as epibenthic zoobenthivores, but may become pelagic at a certain size according to the predation risk experienced at that time. From a length of > 20–25 cm, charr start to become piscivorous. The proportion of piscivorous fish increases with increasing body size. In partly anadromous populations, fish that mature before smolting become freshwater resident, the others anadromous. In some rivers, the morphs occupy separate niches (epibenthic and limnetic), from emergence onwards. The morphs exhibit different degrees of reproductive isolation that vary from a high degree of interbreeding to complete isolation. Usually, they spawn within morph (assortative mating), but alternative male mating behaviour (sneaking, fighting) may occur in stream‐spawning populations and at great depths in lakes. Morphologically specialized morphs appear to feed more effectively than intermediate forms, and selection according to feeding mode, site fidelity and associated assortative mating are prerequisites for the evolution of the different morphs. Charr morphs develop into stable feeding niches under conditions of intense intraspecific competition when there is little competition with other species. Sympatric morphs exhibit different degrees of speciation, but similar morphs in different systems are not individual species because of (1) their polyphyletic origin, (2) the supporting systems are often young, transient environments making the future situation for the populations uncertain, and (3) the genetic differentiation among morphs is low. Sympatric morphs may interbreed and produce fertile hybrids. Nevertheless, sympatric charr morphs should be managed as separate species. Changes in the natural conditions or human impacts to which the morphs are adapted will have a strong influence on the persistence and survival of each different morph.