The ecology and conservation status of Madagascar's endemic freshwater crayfish (Parastacidae; Astacoides)

1. Freshwater crayfish of the genus Astacoides are endemic to the highlands of eastern Madagascar. Very little is known about their ecology and how this affects their vulnerability to threats. Working in the Fianarantsoa forest corridor, we used a combination of ecological research (>29 000 crayfish caught and released) and interviews (>130 interviews in 38 villages) to investigate the ecology and status of four of the seven described species. 2. Astacoides species studied showed very slow growth, with growth rates of Astacoides granulimanus and Astacoides crosnieri among the slowest known in any species of crayfish. We found individuals of all three species for which we had growth data which we estimate at more than 20‐years old. The size at which females became ovigerous varied among species. Astacoides betsileoensis withheld reproduction until a large size (only 30% of females were gravid at 60‐mm carapace length, compared with 90% for A. granulimanus). This is likely to make A. betsileoensis particularly vulnerable to overexploitation and we found that only 10% of individuals measured in a market (n = 909) would have reproduced before being caught compared with 35% of the more common A. granulimanus (n = 30 561). 3. Habitat loss is a serious threat to the genus; even A. granulimanus, the most widespread species, was only found in rivers or streams flanked by natural vegetation. Astacoides caldwelli, the rarest species in this study, was found only at low altitudes (<800 m) in rivers draining forested catchments. Habitat loss is particularly rapid in low elevation forest. A. crosnieri was restricted to swampy land that is rapidly being converted to rice fields. Introduced Asian snakehead fish (Channa maculata) may pose a hitherto unrecognised threat to some species, particularly A. betsileoensis. 4. Madagascar's freshwater habitats have great significance for global biodiversity yet conservation effort, as in much of the world, has focused on terrestrial ecosystems. Until recently almost nothing was known about the ecology of Astacoides crayfish, a diverse and economically important genus. Here, we show that members of the genus vary markedly in their reproductive biology, growth rates, habitat requirements and the threats they face. We suggest that habitat loss is an urgent threat, especially to A. caldwelli and A. crosnieri, while overharvesting is probably the most immediate threat to the larger A. betsileoensis. We call for more attention to be paid to Madagascar's exceptional, yet understudied, freshwater biodiversity.     

The eastern swamp crayfish Gramastacus lacus sp. n. (Decapoda,Parastacidae) a new species of freshwater crayfish from coastal New South Wales,Australia

Gramastacus lacus sp. n., is described from coastal lowlands of the Central and Mid North Coast regions of New South Wales, Australia. Gramastacus lacus has a restricted distribution in ephemeral habitats, being dependent on regular natural flooding and drying cycles, and burrows for survival during temporary dry cycles. Documented are population distributions in lowland habitats (3–48 m, a.s.l.) from Wamberal Lagoon, north along the coastal strip to Wallis Lake. The species is small, reaching a maximum weight of 7 grams and 21.3 mm OCL, and distinguished by a large male genital papilla, large raised post orbital ridges, laterally compressed carapace and elongated chelae.     

Aggressive interactions between three species of freshwater crayfish of the genus Cherax (Decapoda: Parastacidae)

The freshwater crayfish, Cherax destructor Clark, native to southeastern Australia, was first introduced to farm dams in southwestern Western Australia in 1932. The geographic range of the crayfish in Western Australia has increased substantially since then, and in recent years it has become established in natural waterways where it co-occurs with species of freshwater crayfish endemic to southwestern Australia, Cherax cainii and Cherax quinquecarinatus. The potential for competitive exclusion of these endemic species by C. destructor was investigated through laboratory experiments measuring aggressive behaviour. Body mass and species were found to be important factors governing aggressive dominance between C. cainii and C. destructor, with C. cainii winning significantly more interactions only when they were larger in body mass than their opponent. In trials between C. quinquecarinatus and C. destructor of similar body mass, there was no difference between the number of interactions 'won' by the two species. The implications for natural populations are discussed.     

Phylogeographic structure in the gilgie (Decapoda: Parastacidae: Cherax quinquecarinatus): a south‐western Australian freshwater crayfish

The gilgie (Cherax quinquecarinatus) is among the more widespread of the six endemic south‐western Australian freshwater crayfish species. In the present study, the phylogeographic structure of the gilgie was investigated across its distribution to determine whether patterns reflected those identified earlier in a co‐distributed congeneric, the koonac (Cherax preissii). Gilgies were sampled from 20 localities, a 412‐bp fragment of the cytochrome c oxidase subunit I mitochondrial DNA gene was amplified from 75 individuals, and allozyme variation was assayed at nine loci. As in the koonac, three geographically‐restricted lineages were identified: from the north‐western, southern coastal, and intermediate/south‐western regions. Phylogeographic breaks appeared to be congruent with those in the koonac. The extent of genetic differentiation among lineages was comparable to that in the koonac, suggesting temporal congruence of the historical events responsible for the observed structure. A relaxed Bayesian molecular clock suggested that the major clades and lineages in each species diverged in the Late Miocene–Early Pliocene (4.0–9.6 Myr ago), possibly resulting from increasing pulses of aridity. The retrieval of almost‐identical phylogeographic structure in two co‐distributed species suggests that biogeographic regions can be more accurately defined in south‐western Australia. With the geographic fidelity of these lineages, the present data also provide evidence of the translocation of a single individual from the north‐west to the south coast. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 385–402.     

Habitat use of the European mudminnow Umbra krameri and association with other fish species in a disconnected Danube side arm

Fish assemblages along the longitudinal course of an old, disconnected and modified side arm of the Danube floodplain downstream of Vienna, Austria, as well as habitat structure, hydro‐morphological and hydro‐chemical factors, were investigated in order to analyse the key environmental determinants of the European mudminnow Umbra krameri. Generally, U. krameri was the most abundant species in the system. It occurred in disconnected ditches, ponds and pools with dense reed belts and comparatively low nutrient content, indicating its natural association with marsh habitats. At infrequently disturbed sites it was associated with a small group of stagnophilious and highly specialized species with adaptations to strong oxygen fluctuations. At frequently flooded sites, the species was absent or occurred in low abundances, indicating its adaptation to water bodies in older successional stages and its low competitive power in permanently connected floodplain habitats.     

Development of the female reproductive system in the freshwater crayfish Cherax quadricarinatus (Decapoda, Parastacidae)

Abstract. The differentiation of the female reproductive system from a macroscopic and microscopic point of view was studied in Cherax quadricarinatus. For this characterization, 184 females were dissected and processed for the histological analysis. From the differentiation of the ovary up to its maturity, three ovarian morphotypes could be distinguished macroscopically: parallel strands without any contact between them, an H-shaped ovary, and a Y-shaped ovary. These morphotypes were compared within the Astacida. Four ovarian developmental stages were recognized based on ovary color, and the histological structure and relative proportion of cellular types. The post-spawning ovary was also characterized. The components of the female reproductive system sheath were described and its modifications in the ovary and oviducts were determined and compared. Theoretical aspects of the study of sexual differentiation in C. quadricarinatus were discussed within a phylogenetic framework.     

Habitat predictors of genetic diversity for two sympatric wetland‐breeding amphibian species

Population genetic diversity is widely accepted as important to the conservation and management of wildlife. However, habitat features may differentially affect evolutionary processes that facilitate population genetic diversity among sympatric species. We measured genetic diversity for two pond‐breeding amphibian species (Dwarf salamanders, Eurycea quadridigitata; and Southern Leopard frogs, Lithobates sphenocephalus) to understand how habitat characteristics and spatial scale affect genetic diversity across a landscape. Samples were collected from wetlands on a longleaf pine reserve in Georgia. We genotyped microsatellite loci for both species to assess population structures and determine which habitat features were most closely associated with observed heterozygosity and rarefied allelic richness. Both species exhibited significant population genetic structure; however, structure in Southern Leopard frogs was driven primarily by one outlier site. Dwarf salamander allelic richness was greater at sites with less surrounding road area within 0.5 km and more wetland area within 1.0 and 2.5 km, and heterozygosity was greater at sites with more wetland area within 0.5 km. In contrast, neither measure of Southern Leopard frog genetic diversity was associated with any habitat features at any scale we evaluated. Genetic diversity in the Dwarf salamander was strongly associated with land cover variables up to 2.5 km away from breeding wetlands, and/or results suggest that minimizing roads in wetland buffers may be beneficial to the maintenance of population genetic diversity. This study suggests that patterns of genetic differentiation and genetic diversity have associations with different habitat features across different spatial scales for two syntopic pond‐breeding amphibian species.     

Amplification of multiple copies of mitochondrial Cytochrome b gene fragments in the Australian freshwater crayfish,Cherax destructor Clark (Parastacidae: Decapoda)

Non-coding copies of fragments of the mitochondrial genome translocated to the nucleus or pseudogenes are being found with increasing frequency in a diversity of organisms. As part of a study to evaluate the utility of a range of mitochondrial gene regions for population genetic and systematic studies of the Australian freshwater crayfish, Cherax destructor (the yabby), we report the first detection of Cytochrome b (Cyt b) pseudogenes in crustaceans. We amplified and sequenced fragments of the mitochondrial Cyt b gene from 14 individuals of C. destructor using polymerase chain reaction (PCR) with primers designed from conserved regions of Penaeus monodon and Drosophila melanogaster mitochondrial genomes. The phylogenetic tree produced from the amplified fragments using these primers showed a very different topology to the trees obtained from sequences from three other mitochondrial genes, suggesting one or more nuclear pseudogenes have been amplified. Supporting this conclusion, two highly divergent sequences were isolated from each of two single individuals, and a 2 base pair (bp) deletion in one sequence was observed. There was no evidence to support inadvertent amplification of parasite DNA or contamination of samples from other sources. These results add to other recent observations of pseudogenes suggesting the frequent transfer of mitochondrial DNA (mtDNA) genes to the nucleus and reinforces the necessity of great care in interpreting PCR-generated Cyt b sequences used in population or evolutionary studies in freshwater crayfish and crustaceans more generally.     

Evolution underground: A molecular phylogenetic investigation of Australian burrowing freshwater crayfish (Decapoda: Parastacidae) with particular focus on Engaeus Erichson

Phylogenetic relationships and species boundaries of Australian burrowing freshwater crayfish belonging to the genera Engaeus, Engaewa, Geocharax, Gramastacus and Tenuibranchiurus are investigated using combined mitochondrial and nuclear DNA sequence data and Bayesian and Maximum Parsimony methods. Phylogenies are statistically compared to previously published hypotheses. Engaeus, Engaewa, Geocharax, Gramastacus and Tenuibranchiurus form a strongly supported monophyletic clade. This grouping is independently supported by morphology but unites geographically highly disjunct lineages. Our data show two cryptic species in Geocharax, one cryptic species in Gramastacus and two cryptic species within the highly divergent Engaeus lyelli lineage. Using a Bayesian relaxed molecular clock method, the 16S rDNA data show generic-level diversification coinciding with the transition from a wet to arid palaeoclimate near the mid Miocene.     

Habitat use by sympatric populations of Pseudomys novaehollandiae and Mus domesticus in coastal heathland

Abstract Patterns of habitat use by Pseudomys novaehollandiae and Mus domesticus were investigated on a 6. 8 ha area of coastal heathland, almost five years after it last burned in Myall Lakes National Park. One hundred and fifty-one trap stations were positioned on 17 traplines across a mosaic of interlocking macrohabitats. This pattern was a consequence of the site's topography and to a lesser extent fire history. Multivariate statistical procedures were employed to identify those microhabitat parameters that contribute to both individual species’ habitat use and the partitioning of habitat to reduce competition for space. Trapping results show P. novaehollandiae to be almost three times as abundant as M. domesticus at this time, although M. domesticus does show strong regional, and hence local, fluctuations in abundance over time. Both species exhibit habitat selection (P< 0. 05) with individuals preferring dry, tall closed heath and open heathland habitats. Immature P. novaehollandiae (< 15 g) were found mostly in short dense wet heath and may have been dispersing. Species exhibit high between-habitat overlap using Pianka's formula (0. 88). This overlap is substantially reduced within macrohabitats (0. 28), when trap stations are considered as resource states, suggesting a considerable degree of microhabitat separation at the trap station level. Discriminant function and multiple regression analyses identify microhabitat selection that is not strongly focused, suggesting both species to be spatial generalists. Elevation was selected as the most significant variable in these analyses and represents a soil moisture gradient that determines changes in the floristic and structural components of the biotic environment. No single habitat variable was found to explain within-habitat separation convincingly, however, when a factor analysis of habitat variables was used to produce a reduced three-dimensional factor space the outcome was marked. The 95% confidence envelopes for the distribution of each species show an extremely low overlap in factor space, with a zero overlap in the X-Y plane and less than 1% overlap in theY-Z plane. Differential micro-habitat selection does produce effective habitat partitioning, thus facilitating coexistence of these two species. We suspect this occurs primarily by M. domesticus altering habitat selection. There is evidence to suggest that M. domesticus acts as a fugitive species occupying resource niches left vacant by P. novaehollandiae and other small mammal species coexisting on the study area.     

Molecular phylogeny and zoogeography of the freshwater crayfish genus Cherax Erichson (Decapoda: Parastacidae) in Australia

The evolutionary history and biogeography of freshwater-dependent taxa in Australia is of intrinsic interest given the present-day aridity of this continent. Cherax is the most widespread and one of the most species-rich of Australia's nine freshwater crayfish genera. The phylogenetic relationships amongst 19 of the 23 Australian Cherax were established from mitochondrial DNA sequences representing the 12S rRNA and 16S rRNA gene regions. The relationships among species support an initial east–west separation, followed by a north–south divergence in eastern Australia. Molecular clock estimations suggest that these divergences date back to the Miocene. The phylogenetic relationships support endemic speciation within geographical regions and indicate that long-distance dispersal has not led to recent speciation as previously hypothesized. This new evolutionary scenario is consistent with the climatic history of Australia and the evolutionary history of other similarly distributed freshwater-dependent organisms in Australia.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 553–563.     

Habitat use and behavioural ecology of the juveniles of two sympatric damselfishes (Actinopterygii: Pomacentridae) in the south-western Atlantic Ocean

Spatial distribution, microhabitat use and territorial and feeding behaviours were compared between the juveniles of two sympatric territorial damselfishes Stegastes variabilis and Stegastes fuscus on a small tropical reef in the south-western Atlantic Ocean. Juvenile S. variabilis were most abundant at sites subject to stronger hydrodynamics and with mixed benthic cover, whereas juvenile S. fuscus were most abundant at sheltered sites with dense turf algae cover. No differences regarding feeding habits were detected, with both species preferentially feeding on turf algae. Also, despite similarities in territory area and agonistic encounter rates, the identity and proportion of intruders involved in agonistic interactions differed significantly between species. These interdependent traits suggest strong asymmetric competition, with juvenile S. fuscus dominating high-quality sites and evicting juvenile S. variabilis to low quality, marginal areas of the reef.     

Dietary partitioning by five sympatric species of stingray (Dasyatidae) on coral reefs

Dietary characteristics and the degree of dietary partitioning by five species of sympatric stingray were assessed using stomach content and sediment analyses within a coral reef lagoon at Ningaloo Reef, Western Australia (the cowtail Pastinachus atrus, blue‐spotted fantail Taeniura lymma, blue‐spotted mask Neotrygon kuhlii, porcupine Urogymnus asperrimus rays and the reticulate whipray Himantura uarnak). A total of 2804 items were recovered from the stomachs of 170 rays and 3215 individual taxa from the environment, which were used in selectivity analyses. Twenty‐four prey taxa were identified from stomach contents and pooled into 10 taxonomic categories for analysis, of which annelids, prawns, brachyurans and bivalves were the most abundant, together accounting for 96% of the diet. Himantura uarnak had the greatest interspecific dissimilarity in diet, consuming a larger proportion of crustaceans, notably penaeids (41% of total diet) than the other four species of rays, all of which had diets dominated by annelids (71–82% of total diet). Crustacean specialization by H. uarnak may exist to maximize resources and reduce competition among sympatric species. The remaining species may partition resources on the basis of space, rather than diet.     

Structural changes in the spermatophore of the freshwater 'red claw' crayfish Cherax quadricarinatus (Von Martens, 1898) (Decapoda, Parastacidae)

López Greco, L.S. and Lo Nostro, F.L. 2007. Structural changes in the spermatophore of the freshwater ‘red claw’ crayfish Cherax quadricarinatus (Von Martens, 1898) (Decapoda, Parastacidae). —Acta Zoologica (Stockholm) 88 : 000–000 The structure of the spermatophore was studied in Cherax quadricarinatus. Pieces of the distal vas deferens and transferred spermatophore from the females were fixed, cut and stained. Within the distal vas deferens, the primary layer and the secondary layer of the spermatophore were distinguishable. In the latter, two components were detected: cytoplasmic droplets and a homogeneous matrix. During the first 10 minutes post‐extrusion the cytoplasmic droplets drastically changed from looking like ‘empty droplets’; at this time the spermatophore changed from a liquid stage to a sticky one. One hour after extrusion the spermatophore began to harden and within the first 24–48 h post‐mating it was a solid and intense white structure tightly attached to the female; after 72 h it acquired a softer aspect, completely dehiscing between 96 and 120 h post‐mating. Histologically, the primary layer maintained its integrity surrounding the spermatozoa while the secondary layer lost the cytoplasmic droplets. The spermatophore began to hydrate between 24 and 48 h and by 72–96 h many sections of the sperm cord began to coalesce. From 48 h post‐mating some fissures appeared within the matrix that enlarged between 72 and 120 h. We propose that both manipulation by the female and hydration are the mechanisms involved in the release of the spermatozoa from the spermatophore.     

Structural alterations in the male reproductive system of the freshwater crayfish, Cherax quadricarinatus (Decapoda, Parastacidae)

No diseases affecting reproductive performance have been previously reported in freshwater crayfishes. This study aims to characterise one reproductive system abnormality found in males of Cherax quadricarinatus reared in captivity. Fifteen adult males of C. quadricarinatus (70-110 g) were purchased from San Mateo S.A. farm (Entre Ríos, Argentina) each season during 2007. Macroscopic analysis showed that 26.6% of the animals sacrificed in winter presented brownish distal vasa deferentia. Histological analysis showed different levels of structural abnormality in the epithelium of the vasa deferentia and spermatophore. Granular and hyaline haemocytes were identified within the vasa deferentia but no significant differences were found in the sperm count between normal and brownish vas deferens. Histological analysis of the crayfishes sacrificed in autumn also showed these modifications in 22% of the animals, however, they did not show the brownish colour under macroscopic analysis. The similarities between the male reproductive system syndrome in shrimps and the abnormalities found in C. quadricarinatus are notable. An unspecific response to thermic stress is a possible explanation of these structural alterations.     

Phylogeography of the freshwater crayfish Cherax destructor Clark (Parastacidae) in inland Australia: historical fragmentation and recent range expansion

The yabby, Cherax destructor Clark, is the most widespread species in the most widespread genus of Australian freshwater crayfish. It has a distribution that spans several distinct drainage basins and biogeographical regions within semiarid and arid inland Australia. Here we report a study designed to investigate patterns of genetic variation within the species and hypotheses put forward to account for its extensive distribution using DNA sequences from the mitochondrial 16S rRNA gene region. Results of phylogenetic analyses contradicted previous allozyme data and revealed relatively deep phylogenetic structure in the form of three geographically correlated clades. The degree of genetic divergences between clades (8–15 bp) contrasted with the relatively limited haplotype diversity within clades (1–3 bp). Network-based analyses confirmed these results and revealed genetic structure on both larger and more restricted geographical scales. Nevertheless some haplotypes and 1-step clades had large distributions, some of which crossed boundaries between river basins and aquatic biogeographical regions. Thus both older and more recent historical processes, including fragmentation on a larger geographical scale and more recent range expansion on a local scale, appear to be responsible for the observed pattern of genetic variation within C. destructor . These results support elements of alternative hypotheses previously put forward to account for the evolutionary history of C. destructor and the origin of its large distribution.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 539–550.