Genetic and seasonal variation in body colour of the Australian death, adder, Acanthophis antarcticus (Squamata: Elapidae)

Two colour morphs, red and grey, are recognized in the death adder, Acanthophis antarcticus , an Australian elapid snake. The inheritance of these morphs appears to be controlled by two alleles at a single autosomal locus, with red dominant to grey. Both morphs show seasonal changes in dorsal reflectance. They show similar reflectance patterns in winter, whereas in summer red animals become less reflectant, grey animals become more reflectant. Populations in northern and eastern Australian contain predominantly red snakes, whereas southern populations consist of mostly grey snakes. Selection for crypsis through substrata colour matchng may be important in maintaining the polymorephism.     

Reproductive mode may determine geographic distributions in Australian venomous snakes (Pseudechis,Elapidae)

Summary Why are viviparous squamate reptiles more common in cold climates, and oviparous ones in warmer areas? The usual explanation is that (1) oviparous squamates cannot reproduce successfully in cold areas because soil temperatures are too low for embryonic development; and (2) viviparous squamates experience lower survivorship or reproductive success than oviparous taxa in warmer areas. These hypotheses suggest that the boundaries of geographic distributions of congeneric oviparous and viviparous squamates should be predictable from data on thermal tolerances of embryos, and estimated temperatures of soils and gravid female reptiles throughout the potential geographic range of the taxon. In large venomous Australian snakes of the genus Pseudechis, distributional boundaries of oviparous and viviparous taxa can be accurately predicted from such data. This predictive ability, if substantiated by studies of other reproductively biomodal squamate taxa, would support the putative role of reproductive mode as a direct determinant of reptilian geographic distributions.     

Snakes across the Strait: trans-Torresian phylogeographic relationships in three genera of Australasian snakes (Serpentes: Elapidae: Acanthophis, Oxyuranus, and Pseudechis)

We analyze the phylogeny of three genera of Australasian elapid snakes (Acanthophis-death adders; Oxyuranus-taipans; Pseudechis-blacksnakes), using parsimony, maximum likelihood, and Bayesian analysis of sequences of the mitochondrial cytochrome b and ND4 genes. In Acanthophis and Pseudechis, we find evidence of multiple trans-Torresian sister-group relationships. Analyses of the timing of cladogenic events suggest crossings of the Torres Strait on several occasions between the late Miocene and the Pleistocene. These results support a hypothesis of repeated land connections between Australia and New Guinea in the late Cenozoic. Additionally, our results reveal undocumented genetic diversity in Acanthophis and Pseudechis, supporting the existence of more species than previously believed, and provide a phylogenetic framework for a reinterpretation of the systematics of these genera. In contrast, our Oxyuranus scutellatus samples from Queensland and two localities in New Guinea share a single haplotype, suggesting very recent (late Pleistocene) genetic exchange between New Guinean and Australian populations.     

Spatial ecology of arboreal snakes (Hoplocephalus stephensii,Elapidae) in an eastern Australian forest

Abstract Stephens' Banded Snakes (Hoplocephalus stephensii Krefft 1869) are large (to 1 m), highly arboreal elapid snakes, restricted to mesic forested areas along the eastern coast of Australia. Radiotelemetric monitoring of 16 individuals at Whian Whian State Forest in north‐eastern New South Wales over 25 months provided the first data on spatial ecology of this threatened taxon. Two major influences on movements by Stephens' Banded Snakes were identified: the distribution of large hollow‐bearing trees, and the avoidance of conspecifics. Radiotracked snakes were sedentary inside tree hollows for extended periods (mean = 8 days) during their active season, interrupted by occasional long (mean = 124 m) nocturnal movements to another shelter tree. Snakes travelled on the ground rather than within the canopy, and thus were potentially exposed to terrestrial predators. Although the home ranges of the radiotracked snakes overlapped substantially (mean = 27%), simultaneous occupancy of ‘shared’ shelter trees was less common than expected by chance. Hence, we conclude that adult Stephens' Banded Snakes generally avoid the presence of conspecifics. Snakes used from five to 30 shelter trees and home ranges of male snakes were larger than those of females (mean = 20.2 vs 5.4 ha). The large spatial scale of these movements, and limited overlap among individuals, means that a viable population of this taxon requires a large area of contiguous forest. This requirement may explain why the species has not persisted in small forest fragments.     

The Morphology, Growth and Reproduction of Unionidae (Bivalvia) in a Fenland Waterway

The shell morphology and population dynamics of the five BritishUnionidae are compared within a sympatric population. Pseudanodontacomplanata is distinguished from Anodonta anatina and A. cygneaby the hinge length–shell length relationship; this morphologicaldistinction may serve as a useful tool in the identificationof this threatened species. The shell length at a given annuluswas remarkably similar for all five species, although the asymptoticlength is reached most quickly in P. complanata and Unio pictorum.P. complanata is relatively short-lived and attains the lowest maximumlength, while A. cygnea lives more than twice as long and attainsalmost double the length of P. complanata. Unio spp. have ashort gravid season over the summer, while Anodonta spp. havea long gravid period, lasting from Autumn through to Spring.Unlike other members of the Anodontinae, P. complanata has ashort breeding season, overlapping with that of the Unio spp. (Received 4 March 1998; accepted 23 April 1998)     

Isolation, properties and amino acid sequence of a long-chain neurotoxin, Acanthophis antarcticus b, from the venom of an Australian snake (the common death adder, Acanthophis antarcticus).

The venom of an Australian elapid snake, the common death adder (Acanthophis antarcticus), was chromatographed on a CM-cellulose CM52 column. One of the neurotoxic components, Acanthophis antarcticus b (toxin Aa b) was isolated in about 9.4% (A280) yield. The complete amino acid sequence of toxin Aa b was elucidated. Toxin Aa b is composed of 73 amino acid residues, with ten half-cystine residues, and has a formula weight of 8135. Toxin Aa b has no histidine or methionine residue in its sequence. The amino acid sequence of toxin Aa b is homologous with those of other neurotoxins with known sequences, although it is novel in having a valine residue at its N-terminus and an arginine residue at position-23, where a lysine residue is found in almost all the so-far-known neurotoxins. Irrespective of the latter replacement, the toxin Aa b is fully active, with an LD50 value (in mice) of 0.13 microgram/g body weight on intramuscular injection.     

Reproduction and Larval Morphology of Five Ophryotrocha Species (Poly cha eta, Dorvilleidae)

Reproduction and larval morphology of Ophryotrocha puerilis, O. labronica, O. nologlandulata, O. hartmanni , and O. bacci are described. All species have been cultivated in the laboratory. The morphology of adults and larvae is uniform, but there is a considerable variation in the shape of the egg masses. The interrelations between members of the genus Ophryotrocha are discussed: Gonochoric species have the same type of egg mass and the same chromosome number. O. puerilis is unique as a protandrous hermaphrodite. Among contemporary hermaphrodites with male and female segments, fusiform egg masses with a rigid surface are correlated with one larval type, gelatinous egg masses with another larval type.     

Functional morphology of the palato‐maxillary apparatus in “Palatine dragging” snakes (Serpentes: Elapidae: Acanthophis,Oxyuranus)

Elapid snakes have previously been divided into two groups (palatine erectors and palatine draggers) based on the morphology and inferred movements of their palatine bone during prey transport (swallowing). We investigated the morphology and the functioning of the feeding apparatus of several palatine draggers (Acanthophis antarcticus, Oxyuranus scutellatus, Pseudechis australis) and compared them to published records of palatine erectors. We found that the palatine in draggers does not move as a straight extension of the pterygoid as originally proposed. The dragger palato‐pterygoid joint flexes laterally with maxillary rotation when the mouth opens and the jaw apparatus is protracted and slightly ventrally during mouth closing. In contrast, in palatine erectors, the palato‐pterygoid joint flexes ventrally during upper jaw protraction. In draggers, the anterior end of the palatine also projects rostrally during protraction, unlike the stability of the anterior end seen in erectors. Palatine draggers differ from palatine erectors in four structural features of the palatine and its relationships to surrounding elements. The function of the palato‐pterygoid bar in both draggers and erectors can be explained by a typical colubroid muscle contraction pattern, which acts on a set of core characters shared among all derived snakes. Although palatine dragging elapids share a fundamental design of the palato‐maxillary apparatus with all higher snakes, they provide yet another demonstration of minor structural modifications producing functional variants. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Thermoregulation in the Red-beillied Blacksnake, Pseudechis porphyriacus (Elapidae)

Cold Pseudechis porphyriacus aid heating by basking, flattening and tilting. When body temperatures are higher, thermoregulation is achieved by shuttling between sun and shade. A warm snake in a cooling environment frequently coils.
The major factors associated with rate of temperature change in the body core were (1) horizontal gradient between body and neck, (2) posture and (3) vertical gradients from body core to body surface.
The role of behavioral, physiological and physical factors in thermoregulation are discussed.     

Habitat Preference, Reproduction and Diet of the Earthworm Eel, Chendol keelini (Teleostei: Chaudhuriidae)

The ecology of the earthworm eel, Chendol keelini, was studied in the field over a period of nine months. In addition this information was supplemented by aquarium observations. The species was most abundant in pools where it was associated with leaf litter and mats of fine tree roots along the banks. It fed on benthic invertebrates, especially chironomid and ephemeropteran larvae. C. keelini is sexually dimorphic; adult males develop a headhump and grow to a larger size than females. Reproduction was seasonal; the reproductive phase coincided with the wet season and lasted for several months. Fecundity was around 40 eggs per clutch. The eggs were spherical, between 1.2 and 1.5mm in diameter, and possessed a pair of long filaments for adhesion to the substrate. Females probably spawned more than once during the breeding season. The length frequency distributions and juvenile growth suggest that C. keelini is a short-lived species that matures during the first year with few individuals surviving to the second breeding season.     

Uncoupling ecological innovation and speciation in sea snakes (Elapidae, Hydrophiinae, Hydrophiini)

The viviparous sea snakes (Hydrophiini) are by far the most successful living marine reptiles, with ～ 60 species that comprise a prominent component of shallow-water marine ecosystems throughout the Indo-West Pacific. Phylogenetically nested within the ～ 100 species of terrestrial Australo-Melanesian elapids (Hydrophiinae), molecular timescales suggest that the Hydrophiini are also very young, perhaps only ～ 8-13 Myr old. Here, we use likelihood-based analyses of combined phylogenetic and taxonomic data for Hydrophiinae to show that the initial invasion of marine habitats was not accompanied by elevated diversification rates. Rather, a dramatic three to six-fold increase in diversification rates occurred at least 3-5 Myr after this transition, in a single nested clade: the Hydrophis group accounts for ～ 80% of species richness in Hydrophiini and ～ 35% of species richness in (terrestrial and marine) Hydrophiinae. Furthermore, other co-distributed lineages of viviparous sea snakes (and marine Laticauda, Acrochordus and homalopsid snakes) are not especially species rich. Invasion of the oceans has not (by itself) accelerated diversification in Hydrophiini; novelties characterizing the Hydrophis group alone must have contributed to its evolutionary and ecological success.     

Molecular phylogeny of the Australian venomous snake genus Hoplocephalus (Serpentes, Elapidae) and conservation genetics of the threatened H. stephensii

The Australian elapid snakeHoplocephalus stephensii (Stephens' BandedSnake) is patchily distributed in disjunctforest remnants in eastern Australia and islisted as threatened in both states in which itoccurs (Qld and NSW). Here we focus on thephylogeography of H. stephensii toaddress (1) the genetic distinctiveness of thistaxon within its genus and (2) the level ofgenetic diversity present within and betweendisjunct populations from throughout thespecies' range. We sequenced an approximately900 base pair DNA fragment of the mitochondrialgenome that includes half of the ND4 gene andthree tRNA genes. We obtained sequence datafrom 15 H. stephensii individuals drawnfrom four populations, plus representatives ofthe other Hoplocephalus species.Phylogenetic analyses of the data produced asingle fully resolved tree. The two coastaltaxa (H. bungaroides and H. stephensii)are very closely related (2.6–3.1% sequencedivergence) whereas the inland taxon H.bitorquatus is more distantly related to theother two (7.6% vs H. bungaroides;7.8–8.3% vs H. stephensii). Geneticdiversity is low within H. stephensii(nine mitochondrial haplotypes with 1–3haplotypes with only single base pairdifferences within populations). The largestsplit (1.7% sequence divergence) occursbetween the northern population and the threesouthern populations and corresponds to thespecies distribution north and south of theMcPherson Range on the Queensland-New SouthWales border. The three southern populationsdisplay much less molecular divergence (maximumof 0.6% sequence divergence), consistent withthe presence of generally continuous forestthroughout the species' range until Europeaninvasion of Australia 200 years ago, and withradiotelemetric studies that have found highvagility in these arboreal snakes. Thus, onthe basis of genetic distinctiveness we arguethat (1) Hoplocephalus bitorquatus shouldreceive high conservation priority; and (2)managers should treat the Queensland and NSWpopulations of H. stephensi as separateconservation units.     

Diversity in Reproduction and General Morphology between Two Glossiphonia Species (Hirudinea) in Lake Esrom,Denmark*

According to Bennike (1943) the leech fauna in Lake Esrom, Denmark, includes two types of Glossiphonia complanata (L.), namely G. complanata typica Bennike, 1943 and G. complanata concolor (Apáthy, 1888). The results of the present investigation, which were based on material collected in 1975, 1976, and 1980, strongly indicated that we are dealing with two species in Lake Esrom, namely Glossiphonia complanata (L.) and G. (Clepsine) concolor (Apáthy). The two species have a rather similar life history, but G. complanata is on average larger than G. concolor. Differences in reproduction included size of eggs and timing of breeding, but the number of offspring seemed to be similar. Tank experiments revealed that all offspring of G. complanata had a conspecific parent, and likewise for G. concolor. The general appearance of the two species is significantly different, and the average ratio of body width to body height (W/H ratio) was ca. 30% higher for preserved individuals of G. complanata than for G. concolor. The ecological distribution of the two types, and the reliability of pigmentation as a diagnostical tool in leech systematics are discussed. Characters used for the identification of the species are included.     

Notes on the Australian sea-snake Ephalophis greyi M. Smith (Serpentes:Elapidae, Hydrophiinae) and the origin and classification of sea-snakes

The type and two other specimens of Ephalophis greyi are described. The venom gland musculature is of the Demansia type, with an isolated muscle running from the quadrate to the rear of the venom gland. The only other hydrophiine with the Demansia pattern of venom gland musculature, Hydrophis mertoni Roux, is referred, for this and other reasons, to the genus Ephalophis as E. mertoni (Roux). Some notes on a skull of E. mertoni are presented, along with a revised diagnosis of Ephalophis. The Demansia pattern of venom gland muscles seen in Ephalophis is primitive for the Hydrophiinae and other members of that subfamily have secondarily taken on a Glyphodon type of musculature of the venom gland by loss of the muscle slip from quadrate to gland. The Demansia group of terrestrial elapids, particularly the genus Drepanodontis Worrell, stand close to the origin of the Hydrophiinae. A classification of the Hydrophiinae into three generic groups is presented, with Ephalophis, Aipysurus , and Emydocephalus referred to an Aipysurus group.     

Reproduction of the Australian freshwater turtle Emydura krefftii (Chelonia: Chelidae)

A study of the reproduction of Krefft's river tortoise, Emydura krefftii , was conducted in the perched dune lakes of Fraser Island, Queensland. Mature male specimens exhibit a postnuptial pattern of spermatogenesis typical of temperate-zone turtles elsewhere, with a peak in spermatogenic activity in autumn and a cessation of activity during the breeding season in spring and early summer. The spermatogenic cycle is paralleled by seasonal variation in testicular weight (standardized for body size) and in the diameter of the seminiferous tubules. Sperm are abundant in the epididymal canals throughout the year. Mating was observed in autumn, late winter and spring.
Females have a cyclic reproductive pattern, with distinct phases of follicular enlargement, ovulation and oviducal period, and quiescence. Yolk begins to accumulate in the ovaries in late summer, and the accumulation continues unabated through the winter, presumably by the transfer of material from fat stores to the ovaries. Ovulations occur from late winter to mid-summer. Atresia of follicles that fail to ovulate was demonstrated histologically.
Emydura krefftii lay up to three clutches of hard-shelled ellipsoid eggs per season. Each clutch contains between four and 10 eggs; the number is strongly correlated with maternal body size. Reproductive potential ranges from 12 eggs per annum for a female that has recently matured (carapace length c. 150 mm), to 30 eggs per annum for a full-sized female (length c. 250 mm). Selected life-history traits of Emydura krefftii are discussed in the context of findings for other populations of the species and for other species of freshwater turtle.     

Diet and Mandibular Morphology in African Apes

Investigations seeking to understand the relationship between mandibular form, function, and dietary behavior have focused on the mandibular corpus and symphysis. African apes vary along a gradient of folivory/frugivory, yet few studies have evaluated the morphology of the mandibular corpus and symphysis in these taxa, and the investigations have yielded mixed results. Specifically, studies using external metrics have identified differences in mandibular proportions that analysis of cortical bone distribution has not substantiated. I contribute to the ongoing debate on the relationship between jaw form and dietary behavior by comparing mandibular corporal and symphyseal shapes in African apes. Importantly, and in contrast to previous studies of African ape internal geometry, I include the Virunga mountain gorillas (Gorilla beringei beringei), the ape most specialized toward a folivorous diet. I test the hypotheses that 1) Gorilla beringei beringei always has significantly more robust mandibular corpora and symphyses, relative to mandibular length, than all other African apes and 2) all gorillas have significantly more robust mandibular corpora and symphyses, relative to mandibular length, than Pan. Results demonstrate that the folivorous mountain gorillas consistently exhibit a relatively wider mandibular symphysis and corpus than all other African apes. Furthermore, all gorillas consistently exhibit relatively more robust mandibular corporal and symphyseal dimensions than Pan. The results indicate that among African apes, mountain gorillas are better able to counter lateral transverse bending (wishboning) loads at the symphysis and torsional loads at the corpus. All gorillas are likewise better able to resist wishboning and vertical bending at the symphysis, and sagittal bending and torsion at the corpus, than Pan, findings that are consistent with masticating relatively tougher foods, repetitive loading of the jaws, or both. I offer possible explanations for the lack of concordance in results between studies that have analyzed the biomechanical properties of African ape mandibles and others that have relied on external metrics. More comprehensive study of the internal geometry of the mandible is needed to resolve whether African apes differ morphologically in ways predicted by dietary variation.

Death,Funerals and Emotion in an Australian Aboriginal Community

This paper discusses emotions as intersubjective states that develop in the interactions between individuals. Central to the discussion is the funeral of an Aboriginal elder in Cape York Peninsula, Australia. The actions of certain of her kin are described, and demonstrate their concern to bridge the gap between the loss of such elders and the uncertainties of a future without their guidance. Funerals are clearly events marking the social change associated with death and allow for intense expressions of grief, but they are also part of the trajectory of everyday life. In the social context discussed, beliefs about intersubjectivity between the spirits of the dead and their living kin inform all spheres of action, including the economic, and require observance of the cultural value of ‘respect’. Understanding emotions as interactive states engaged in such locally significant events sheds light on the relationship between them and collective political aspirations.     

A multivariate morphometric analysis and systematic review of Pseudonaja (Serpentes,Elapidae, Hydrophiinae)

Pseudonaja is a clade of seven nominal species of elapid snakes distributed throughout Australia and in southern New Guinea. The species‐level systematics of this group is generally considered to be problematic. A recent phylogenetic analysis of mitochondrial DNA sequences for a geographically extensive series of Pseudonaja specimens revealed nine major clades, of which six largely coincide with nominal species (P. affinis, P. guttata, P. inframacula, P. ingrami, P. modesta and P. textilis). The three remaining clades are composed of specimens currently referred to P. nuchalis. This paper presents a multivariate analysis of 30 morphometric variables recorded for 220 specimens, representing the P. affinis, P. inframacula, P. textilis and three P. nuchalis clades (P. guttata, P. ingrami and P. modesta are well‐demarcated species and, accordingly, were not considered). The morphometric data readily separate these putative lineages, affording compelling evidence that they constitute evolutionary species. The names aspidorhyncha and mengdeni are resurrected for two of the three species presently recognized as P. nuchalis. These species, P. affinis, P. inframacula, P. nuchalis and P. textilis are redescribed. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 171–197.     

Reproduction and perennation of Chondrilla juncea L. (skeleton weed) in the Western Australian wheatbelt

Survivorship of first-year plants of Chondrilla juncea was considerably higher on a deep yellow sand than on a yellow duplex soil. Mortality on the latter soil was associated with drying of the Al horizon during early summer. No plants produced seeds in their first year of growth on the duplex soil. However, during both years of the study, first-year individuals of both forms of C. juncea produced viable seeds when growing on the deep sand. Higher population growth rates could be expected on deep sands, owing to a combination of earlier reproduction and greater survivorship through the first summer. It is suggested that whilst the entire wheatbelt might be climatically suitable for C. juncea, the invasive potential of this weed might be restricted by edaphic characteristics over large areas.