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.     

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.     

Genetic Connectivity among Populations of an Endangered Snake Species from Southeastern Australia (Hoplocephalus bungaroides, Elapidae)

For endangered species that persist as apparently isolated populations within a previously more extensive range, the degree of genetic exchange between those populations is critical to conservation and management. A lack of gene flow can exacerbate impacts of threatening processes and delay or prevent colonization of sites after local extirpation. The broad-headed snake, Hoplocephalus bungaroides, is a small venomous species restricted to a handful of disjunct reserves near Sydney, Australia. Mark-recapture studies have indicated low vagility for this ambush predator, suggesting that gene flow also may be low. However, our analyses of 11 microsatellite loci from 163 snakes collected in Morton National Park, from six sites within a 10-km diameter, suggest relatively high rates of gene flow among sites. Most populations exchange genes with each other, with one large population serving as a source area and smaller populations apparently acting as sinks. About half of the juvenile snakes, for which we could reliably infer parentage, were collected from populations other than those in which we collected their putative parents. As expected from the snakes' reliance on rocky outcrops during cooler months of the year, most gene flow appears to be along sandstone plateaux rather than across the densely forested valleys that separate plateaux. The unexpectedly high rates of gene flow on a landscape scale are encouraging for future conservation of this endangered taxon. For example, wildlife managers could conserve broad-headed snakes by restoring habitats near extant source populations in areas predicted to be least affected by future climate change.     

The impact of fire regimes on populations of an endangered lizard in montane south‐eastern Australia

The Blue Mountains water skink (Eulamprus leuraensis; Scincidae) is restricted to less than 40 fragmented swamp sites, all within the Blue Mountains and Newnes Plateau areas of New South Wales, Australia. Climate change is expected to increase fire frequency in the area, potentially degrading habitat quality for this endangered reptile. We quantified lizard abundances in 12 swamps using standardized surveys, and constructed a Global Information System (GIS) database to determine fire‐histories for each swamp since 1967. The abundance of Blue Mountains water skinks was negatively correlated with fire frequency, but not with time since fire. Indirect impacts of fire (mediated via shifts in vegetation density, moisture levels, prey availability and post‐fire predation) may be more important than direct effects in these cool, moist habitats. Although lizards were less common in swamps close to urban areas, and less common in frequently burnt areas, viable populations of this endangered reptile still persist even in anthropogenically disturbed swamps and in swamps that have experienced up to four fires in 20 years. Future research could usefully extend these analyses to other swamps in the locality, and explore the broader impacts of fire regimes on the distinctive flora and fauna of this threatened ecological community.     

Relationships between seed germination requirements and ecophysiological characteristics aid the recovery of threatened native plant species in Western Australia

Summary One of the foremost technical issues addressed in reintroduction and restoration projects is the feasibility of establishing living plants. To advance the recovery process, the germination requirements of 201 threatened Western Australian seed‐bearing taxa representing a range of life forms, families and ecophysiological characteristics were studied. Procedures used to stimulate germination in otherwise dormant seed involved pretreatment using thermal shock, scarification, seed coat removal, soaking in an aqueous smoke solution and/or additions of the growth hormone gibberellic acid (GA₃). Sixty‐one taxa germinated under the basic trial conditions of light (12‐ h photoperiod), temperature (constant 15°C) and moisture, without additional pretreatments. These taxa were generally those with canopy‐stored seeds in the families Proteaceae and Casuarinaceae, and small‐seeded taxa in Myrtaceae. Taxa with soil‐stored seeds required single or multiple cues to stimulate germination. Seeds in the families Fabaceae and Mimosaceae were dependent on cracking of the seed coat, mechanically through nicking of the testa or through thermal shock, as were several non‐leguminous species of the Sterculiaceae and Liliaceae. Complete or partial removal of seed coats, in conjunction with GA₃ enhanced germination percentage in some taxa of the Myoporaceae, Lamiaceae and Myrtaceae. Application of GA₃ also enhanced germination percentage in members of the Epacridaceae. Several taxa previously stimulated by aqueous smoke solutions were equally responsive to additions of GA₃ after complete seed coat removal. In general, species with seed weights greater than 10 mg germinated better under a range of conditions than those with lighter seeds. There was no difference in germinability between resprouter and seeder species, and no obvious relationship between seed weight and germination rate. In the light of previous studies these results indicate that the relationship between germination requirements and ecophysiological characteristics is similar for both threatened and common species. These data will enable better prediction of likely dormancy breaking cues for other related species and will greatly assist the process of recovery and restoration work for mining operations and community bushland regeneration as well as single species reintroductions.     

Genetic analysis of populations of threatened snake species using RAPD markers

Snakes are a particularly threatened vertebrate taxon, with distributions of many species and populations becoming increasingly fragmented. At present, little is known about the degree of genetic differentiation that exists between isolated populations even though such information may be critical to their survival and conservation. As an example of how recently developed RAPD genetic markers can be used in conservation genetics, we present preliminary results from a study which used these DNA-based markers to assess population divergence in two threatened Canadian snakes, the black rat snake ( Elaphe o. obsoleta ) and the eastern massasauga rattlesnake ( Sistrurus c. catenatus ). We present information on the levels of variation and reliability of amplification for fragments generated from five primers. We then use a recently developed analytical technique to estimate levels of nucleotide diversity within populations and sequence divergence between populations. Our results show that intrapopulation levels of divergence as estimated by the methods of Clark & Lanigan ( Molecular Biology and Evolution 1993, 10 , 1096–1111) approximate those found for mtDNA in vertebrates and that diversity between snake populations is small and non-significant when tested using randomization procedures. Thus, our study provides an example of how RAPDs can be applied to conservation genetic studies of vertebrates and suggest that the snake populations we examined have only recently become isolated and maybe considered genetically equivalent from a conservation perspective, although this conclusion needs to be confirmed with other DNA-based markers.     

Habitat preference in the critically endangered yellow‐tailed woolly monkey (Lagothrix flavicauda) at La Esperanza,Peru

Habitat loss is one of the main threats to wildlife. Therefore, knowledge of habitat use and preference is essential for the design of conservation strategies and identification of priority sites for the protection of endangered species. The yellow‐tailed woolly monkey (Lagothrix flavicauda Humboldt, 1812), categorized as Critically Endangered on the IUCN Red List, is endemic to montane forests in northern Peru where its habitat is greatly threatened. We assessed how habitat use and preference in L. flavicauda are linked to forest structure and composition. The study took place near La Esperanza, in the Amazonas region, Peru. Our objective was to identify characteristics of habitat most utilized by L. flavicauda to provide information that will be useful for the selection of priority sites for conservation measures. Using presence records collected from May 2013 to February 2014 for one group of L. flavicauda, we classified the study site into three different use zones: low‐use, medium‐use, and high‐use. We assessed forest structure and composition for all use zones using 0.1 ha Gentry vegetation transects. Results show high levels of variation in plant species composition across the three use zones. Plants used as food resources had considerably greater density, dominance, and ecological importance in high‐use zones. High‐use zones presented similar structure to medium‐ and low‐use zones; thus it remains difficult to assess the influence of forest structure on habitat preference. We recommend focusing conservation efforts on areas with a similar floristic composition to the high‐use zones recorded in this study and suggest utilizing key alimentation species for reforestation efforts.     

Reproductive patterns of a biologically invasive species: the brown tree snake (Boiga irregularis) in eastern Australia

Boiga irregularis , a snake native to Australasia, is a biologically invasive species that has become established on Guam and threatens to spread to other Pacific Islands. In eastern Queensland, we found that the snake breeds seasonally and gonadal development of males and females is temporally dissociated in time. Although the period of mating in this snake is not known, these data suggest that obligatory sperm storage in the reproductive tract occurs during the reproductive cycle. Such sperm storage may partially explain the success of this species as a biological invader.     

Novel microsatellite loci identified from the Australian eastern small‐eyed snake (Elapidae: Rhinocephalus nigrescens) and cross species amplification in the related genus Suta

A total of 15 microsatellite primers pairs were developed for the Australian small‐eyed snake Rhinoplocephalus nigrescens. Five primers were used to screen 93 individuals of R. nigrescens and were also tested against eight species of the closely related genus Suta. Allelic diversity in R. nigrescens was high in three loci (12–27) and there was high heterozygosity (0.58–0.82). Observed heterozygosity did not deviate from Hardy–Weinberg expectations for the five loci tested. These primers will be useful in studies of population genetics and mating systems of small‐eyed snakes and related species.     

Microhabitat relationships among five lizard species associated with granite outcrops in fragmented agricultural landscapes of south‐eastern Australia

A fundamental part of developing effective biodiversity conservation is to understand what factors affect the distribution and abundance of particular species. However, there is a paucity of data on ecological requirements and habitat relationships for many species, especially for groups such as reptiles. Furthermore, it is not clear whether habitat relationships for particular species in a given environment are transferable to other environments within their geographical range. This has implications for the type of ‘landscape model’ used to guide management decisions in different environments worldwide. To test the hypothesis that species‐specific habitat relationships are transferable to other environments, we present microhabitat models for five common lizard species from a poorly studied habitat – insular granite outcrops, and then compared these relationships with studies from other environments in south‐eastern Australia. We recorded twelve species from five families, representing 699 individuals, from 44 outcrops in the south‐west slopes of New South Wales. Five lizard species were abundant and accounted for 95% of all observations: Egernia striolata, Ctenotus robustus, Cryptoblepharus carnabyi, Morethia boulengeri and Carlia tetradactyla (Scincidae). Linear regression modelling revealed suites of different variables related to the abundance patterns of individual species, some of which were broadly congruent with those measured for each species in other environments. However, additional variables, particular to rocky environments, were found to relate to reptile abundance in this environment. This finding means that species' habitat relationships in one habitat may not be readily transferable to other environments, even those relatively close by. Based on these data, management decisions targeting reptile conservation in agricultural landscapes, which contain rocky outcrops, will be best guided by landscape models that not only recognize gradients in habitat suitability, but are also flexible enough to incorporate intraspecies habitat variability.     

Edge effects across boundaries between natural and restored jarrah (Eucalyptus marginata) forests in south‐western Australia

Edge effects are a widespread and ubiquitous ecological phenomenon, yet they remain poorly studied across edges between restored and natural forests. To address this lack of knowledge, we studied vertebrate communities across edges between 3‐year old restored mine‐pits and adjacent unmined forest in the jarrah (Eucalyptus marginata) forest of south‐western Australia. We found that mammal communities showed no edge response but reptile communities did. Overall reptile abundance and Morethia obscura abundance were higher in unmined forest along edges, Egernia napoleonis abundance was lower in unmined forest along edges, while Pogona minor abundance was lower in restored mine‐pits along edges. Predictive models were unable to predict species edge responses, due to the lack of knowledge of the ecology of jarrah forest reptiles, but proved useful in identifying potential ecological mechanisms behind observed edge responses and suggested that potential mechanisms were likely different for each species. Our study is the first to show edge responses in both habitats forming the edge between restored and natural forests, emphasizing the importance of studying both habitats forming the edge. Our results also suggest that, despite being poorly studied, edge responses are common across edges between restored and natural forest and result from a variety of ecological mechanisms. An increased understanding of the ecological mechanisms driving edge responses across edges between restored and natural forests will improve our ability to integrate restored areas into cross‐landscape management and, ultimately, improve our ability to manage landscapes for biodiversity conservation.     

Habitat requirements and host selectivity of Thesium species (Santalaceae)

The aim of this study was to identify the habitat requirements and host specificity of three root parasites in the genus Thesium (T. linophyllon, T. bavarum and T. ebracteatum). We conducted a vegetation survey and tested whether some plant species occur significantly more often in plots with Thesium spp. than expected, thereby representing potential hosts of the parasites. To test if the potential hosts are the species to which Thesium is physically attached, we also conducted an excavating and sowing experiment. We found a positive association with surrounding species for T. linophyllon, but not for the other two species. The number of haustoria of T. linophyllon attached to roots of host species was significantly affected by the number of roots of the surrounding species and by their identity (species, family and plant group). Thesium was, however, attached to 94% of all species occurring in the plots. We suggest that Thesium spp. do not specifically select hosts, but rather occur in microhabitats with specific conditions. This conclusion is also supported by the fact that there were differences in the microhabitats with and without Thesium and that germination of Thesium in the sowing experiment was higher in pots without any host. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 394–408.     

Management of threatened insect species in Australia, with particular reference to butterflies

Abstract   Advances in determination of conservation status of insects through evaluation of threats (rather than simple 'rarity' alone) help to set priorities for allocation of conservation resources, and formulate effective legislative treatment and field management for the most needy species. Summaries of several recent conservation studies on butterflies exemplify the variety of practical needs, amongst which habitat protection and enhancement is paramount. However, the paucity of biological and distributional information on most Australian insects necessitates considerable flexibility in approach, so that perceptions of conservation need may be highly subjective. Listing species for protection and conservation priority must be viewed as a dynamic process, with opportunity for rapid action, either to list or de-list taxa as research and recovery actions lead to change in perceived or real status and conservation need.     

Ultrastructure of the placentae of the natricine snake,Virginia striatula (Reptilia: Squamata)

Virginia striatula is a viviparous snake with a complex pattern of embryonic nutrition. Nutrients for embryonic development are provided by large, yolked eggs, supplemented by placental transfer. Placentation in this species is surprisingly elaborate for a predominantly lecithotrophic squamate reptile. The embryonic-maternal interface consists of three structurally distinct areas, an omphalallantoic placenta and a regionally diversified chorioallantoic placenta. The chorioallantoic placenta over the embryonic hemisphere (paramesometrial region) of the egg, features close apposition of embryonic and uterine blood vessels because of the attenuate form of the interceding epithelial cells. The periphery of the chorioallantoic placenta, which is adjacent to the omphalallantoic placenta, is characterized by a simple cuboidal uterine epithelium apposed to a stratified cuboidal chorionic epithelium. There are no sites with attenuate epithelial cells and close vascular apposition. The morphology of the omphalallantoic placenta is similar to that of the peripheral chorioallantoic placenta, except that the height of uterine epithelial cells is greater and allantoic blood vessels are not associated with the embryonic epithelium. The functional capabilities of the three placental regions are not known, but structural characteristics suggest that the omphalallantoic placenta and peripheral zone of the chorioallantoic placenta are sites of nutritional provision via histotrophy. The paramesometrial region of the chorioallantoic placenta is also nutritive, in addition to functioning as the primary embryonic respiratory system. The structure of the chorioallantoic placenta of V. striatula is a new placental morphotype for squamate reptiles that is not represented by a classic model for the evolution of reptilian placentation.     

Preliminary Insights into the Habitat Preferences of the Centralian Bandy Bandy （Vermicella vermiformis）（Squamata： Elapidae） in Central Australia

Bandy Bandy’s（Vermicella spp.）are a striking,black-and-white ringed genus of small elapid snakes endemic to Australia.All taxa are burrowers and little is known of their biology and ecology.We investigated the habitat preferences of the only arid-dwelling species,the centralian bandy bandy（Vermicella vermiformis）,in the MacDonnell Ranges west of Alice Springs in the Northern Territory.Using systematic road-cruising,we encountered 16V.vermiformis over a 12 months period between 2009 and 2010.We used logistic regression to model the occurrence of the species against a range of different habitat variables collected at multiple scales.Despite the small sample size,V.vermiformis exhibited a clear preference for acacia shrubland habitats,with acacia variables present in all AICc ranked models in the 95%confidence set.The factors driving this association,together with the preference for habitat not burnt in the most recent wildfires,may be related to the abundance of their only known prey,blind snakes（Ramphotyphlops spp.）.     

Benchmarks and predictors of coarse woody debris in native forests of eastern Australia

Fallen coarse woody debris (CWD) is critical to forest biodiversity and function. Few studies model factors that influence CWD availability, although such investigations are critically needed to inform sustainable forest management. We assess benchmark levels of CWD in unharvested native forests and those harvested for timber, across a range of forests in north‐east New South Wales, Australia. We found timber‐harvesting was the dominant driver of CWD, with almost double the count (pieces ha^?1) and volume (m³ ha^?1) of total CWD in selectively harvested than unharvested sites. This pattern was consistent across wet and dry forest types. Harvested sites had greater counts of hollow‐bearing logs, and greater volumes of small and medium‐sized CWD (15–50 cm diameter) than unharvested sites. There was no effect of harvesting on the volume of large CWD (>51 cm diameter). Total volumes of CWD (>15 cm diameter) varied from 114 to 166 m³ ha^?1. We found few differences in CWD counts and volumes between forest types, with grassy woodlands and forests containing less CWD than other dry and shrubby forest types, reflecting lower potential input rates. The CWD levels recorded here are similar to those recorded in dry and wet sclerophyll forests elsewhere in Australia and are typical of global estimates for ‘old growth’ forests. Using general linear models we captured up to 57% of the variation in CWD across sites, and found that timber harvesting, topography and the numbers of standing hollow‐bearing and dead trees were significant predictors of CWD. Values for unharvested forest provide a benchmark that could be used to inform retention guidelines for CWD in managed forests in this region. Further assessment of the effect of repeat timber harvesting is needed to fully understand its impact on CWD dynamics, especially if forest residues resulting from timber harvesting are removed from native forests for bioenergy production.