Zoogeography and relationships of Australasian skates (Chondrichthyes: Rajidae)

Aim To investigate distributional patterns and derivation of skates in the Australasian realm. Location Australasia. Methods Genus‐group skate taxa were defined for this region for the first time and new systematic information, as well as bathymetric and geographical data, used to identify distribution patterns. Results The extant skate fauna of Australasia (Australia, New Zealand, New Caledonia and adjacent subAntarctic dependencies) is highly diverse and endemic with sixty‐two species from twelve currently recognized, nominal genus‐group taxa. These include the hardnose skate (rajin) groups Anacanthobatis, Amblyraja, Dipturus, Okamejei, Rajella and Leucoraja, and softnose skate (arhynchobatin) genera Arhynchobatis, Bathyraja, Insentiraja, Irolita, Pavoraja and Notoraja. Additional new and currently unrecognized nominal taxa of both specific and supraspecific ranks also occur in the region. The subfamily Arhynchobatinae is particularly speciose in Australasia, and the New Zealand/New Caledonian fauna is dominated by undescribed supraspecific taxa and species. The Australian fauna, although well represented by arhynchobatins, is dominated by Dipturus‐like skates and shows little overlap in species composition with the fauna of New Zealand and New Caledonia. Similarly, these faunas exhibit no overlap with the polar faunas of the Australian subAntarctic dependencies (Heard and Macdonald Islands) to the south. Skates appear to be absent from the Macquarie Ridge at the southern margin of the New Zealand Plateau. Their absence off New Guinea probably reflects inadequate sampling and the subsequent poor knowledge of that region's deepwater fish fauna. Main conclusions Skates appear to have existed in the eastern, Australasian sector of Gondwana before fragmentation in the late Cretaceous. The extant fauna appears to be derived from elements of Gondwanan origin, dispersal from the eastern and western Tethys Sea, and intraregional vicariance speciation.     

Standardized diet compositions and trophic levels of skates (Chondrichthyes: Rajiformes: Rajoidei)

Skates by virtue of their abundance and widespread occurrence appear to play an influential role in the food webs of demersal marine communities. However, few quantitative dietary studies have been conducted on this elasmobranch group. Therefore, to better understand the ecological role of skates, standardized diet compositions and trophic level (TL) values were calculated from quantitative studies, and compared within and among skate and shark taxa. Prey items were grouped into 11 general categories to facilitate standardized diet composition and TL calculations. Trophic level values were calculated for 60 skate species with TL estimates ranging from 3.48 to 4.22 (mean TL = 3.80 ± 0.02 SE). Standardized diet composition results revealed that decapods and fishes were the main prey taxa of most skate species followed by amphipods and polychaetes. Correspondingly, cluster analysis of diet composition data revealed four major trophic guilds, each dominated by one of these prey groups. Fish and decapod guilds were dominant comprising 39 of 48 species analyzed. Analysis of skate families revealed that the Arhynchobatidae and Rajidae had similar TL values of 3.86 and 3.79 (t-test, P = 0.27), respectively. The Anacanthobatidae were represented by a single species, Cruriraja parcomaculata, with a TL of 3.53. Statistical comparison of TL values calculated for five genera (Bathyraja, Leucoraja, Raja, Rajella, Rhinoraja) revealed a significant difference between Bathyraja and Rajella (t-test, P = 0.03). A positive correlation was observed between TL and total length (L _T) with larger skates (e.g. >100 cm L _T) tending to have a higher calculated TL value (>3.9). Skates were found to occupy TLs similar to those of several co-occurring demersal shark families including the Scyliorhinidae, Squatinidae, and Triakidae. Results from this study support recent assertions that skates utilize similar resources to those of other upper trophic-level marine predators, e.g. seabirds, marine mammals, and sharks. These preliminary findings will hopefully encourage future research into the trophic relationships and ecological impact of these interesting and important demersal predators.     

Dietary comparisons of six skate species (Rajidae) in south-eastern Australian waters

The diet of six skate species caught as bycatch in south-eastern Australian waters was examined over a 2-year period. The skates were segregated into two regions (continental shelf and continental slope) based on prey species and depth of capture. The shelf group consisted of four species, Dipturus sp. A, D. cerva, D. lemprieri and D. whitleyi, while the slope group comprised two species, Dipturus sp. B and D. gudgeri. The two groups varied in feeding strategies with the shelf species generally occupying a broader feeding niche and preying on a larger diversity of prey including a variety of crustaceans (brachyurans, anomurans, achelates, carideans and dendobranchiates), cephalopods, elasmobranchs and teleosts. Within the slope group, Dipturus sp. B and D. gudgeri were more specialised. Dipturus sp. B preyed primarily on anomurans (galatheids) and bachyurans (homolids), whereas D. gudgeri preyed primarily on teleosts. A size related change in diet was evident for all species with the exception of D. gudgeri in which all sizes preyed predominantly on teleosts. Smaller representatives of the four shelf species all preyed on numerous amounts of caridean shrimps, in particular Leptochela sydniensis. In contrast, the continental slope species, Dipturus sp. B consumed anomurans when small, shifting to brachyurans with increasing size. Of the six skate species examined in this study, three were secondary consumers (trophic level <3) and the remaining three tertiary consumers (trophic level >4). Although ANOSIM found significant differences in dietary composition between species within groups, there was some overlap in prey species amongst co-existing skates, which suggests that there is some degree of resource partitioning amongst them.     

Observations on the electric organ discharge of two skate species (Chondrichthyes: Rajidae) and its relationship to behaviour

Synopsis The electric organ discharge (EOD) of the little skate,Raja erinacea and winter skate,R. ocellata was recorded both from isolated individuals and from small groups using methods that allowed for the identification of individuals producing EODs. Pulse duration, train lengh, frequency, and pulse patterns are characterized and correlated with behaviour. The two species,R. erinacea andR. ocellata, were found to have characteristically different EOD pulse durations of 70 ms and 217 ms respectively. Isolated skates rarely discharged whereas groups of skates were found to discharge regularly. The EOD was evoked by tactile prodding, physical contact with other skates and electrical stimulation. Skates also discharged reflexively in response to an artificially induced head-positive DC stimulus, sine wave and monopolar square pulses. During approach and contact, skates responded to each other with interacting EOD displays. EOD interaction and pulse duration differences between other species suggest a possible intra-specific communication function of the EOD inRaja.     

Skates and rays (Elasmobranchii,Batomorphii) from the Eocene La Meseta and Submeseta formations,Seymour Island,Antarctica

Abstract

Eocene deposits of the famous La Meseta Formation of Seymour Island, Antarctic Peninsula, yielded the most diverse Paleogene fossil elasmobranch association of the Southern Hemisphere. In this assemblage, sharks clearly dominate the fauna, whereas batoids are very rare components. Herein, we describe two new taxa of cold water tolerant skates, Marambioraja leiostemma gen. et sp. nov., and Mesetaraja maleficapelli gen. et sp. nov., two new species of the genus Raja, Raja amphitrita sp. nov. and Raja manitaria sp. nov., as well as remains of warm water adapted myliobatiforms. It is, however, not possible to unambiguously assign these remains either to Myliobatidae or Rhinopteridae, or to any specific genus. Previously reported remains of Raja/Bathyraja sp. are assigned to the new described species Raja manitaria sp. nov. The biogeographic distribution of extant and extinct rays and skates clearly shows that both groups are more widely distributed today than in the past, and additionally seem to have been more diverse in the Northern than the Southern Hemisphere. The occurrence, albeit rare of isolated teeth of skates (Rajidae) and rays (Myliobatidae) in the La Meseta Formation representes a minimum age constraint for their first appearance in the Southern Ocean.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:E507D86C-FFEC-4047-A337-AE77606BB1A6     

Captive breeding of the barndoor skate (Dipturus laevis) at the Montreal Biodome,with comparison notes on two other captive‐bred skate species

In 1997, the Montreal Biodome obtained five barndoor skates (Dipturus laevis) from the waters off Boston, Massachusetts. Six years later, those specimens began reproducing, and the first egg case was collected in November 2003. Since then, 73 hatchlings have been born and raised. Egg cases were observed year round, and annual fecundity was measured for the first time: one female laid 69 eggs in 2005, 85 in 2006 and 115 in 2007. Egg incubation was longer than believed previously, ranging from 342 to 494 days. Hatching occurred throughout the year. Hatchlings averaged 193 mm total length and 128 mm disk width and weighed 32 g. They were fed krill and diced fish. All but one survived the first month. A photo identification system was useful in recognizing two groups of 10 specimens during their first year, and transponders could be inserted in the wing muscles of 1‐year‐old skates. Total lengths at birth and at age 2 were similar to the data reported from the wild, suggesting a similar growth pattern in captivity. The reproduction characteristics of the barndoor skate were compared with those of two other skate species currently bred at the Montreal Biodome, the winter skate (Leucoraja ocellata) and the thorny skate (Amblyraja radiata). Zoo Biol 27:145–153, 2008. © 2008 Wiley‐Liss, Inc.     

A conservation assessment of the freshwater crabs of southern Africa (Brachyura: Potamonautidae)

Recent taxonomic revisions of the freshwater crabs of southern Africa (Angola, Botswana, Lesotho, Mozambique, Namibia, South Africa, Swaziland, Zambia and Zimbabwe) allow accurate depictions of their diversity, distribution patterns and conservation status. The southern African region is home to nineteen species of freshwater crabs all belonging to the genus Potamonautes (family Potamonautidae). These crabs show high levels of species endemism (84%) to the southern African region and to the country of South Africa (74%). The conservation status of each species is assessed using the IUCN (2003) Red List criteria, based on detailed compilations of the majority of known specimens. The results indicate that one species should be considered vulnerable, fifteen species least concern and three species data deficient. The results have been utilized by the IUCN for Red Lists, and may prove useful when developing a conservation strategy for southern Africa’s endemic freshwater crab fauna.     

Reanalysis of long‐term surveys on the ecology and biology of mud skate (Rhinoraja taranetzi Dolganov, 1985) in the northwestern Pacific (1993–2002)

Based on data sampled in 1992–2002, the occurrence, spatial distribution, bathymetry, bottom temperatures preferences, size composition, feeding, and some features of reproductive biology of mud skate Rhinoraja taranetzi in the Pacific waters off the northern Kuril Islands and southeastern Kamchatka are considered. Throughout the year, the mud skate was most abundant in the central part of the study area, from the coast of Onekotan Island to the southern tip of Kamchatka peninsula. The proportion of this species in bottom trawl catches in different seasons has changed slightly. However, maximum catch occurred in September–December. In April–May mud skates occupied shallower depths (mean 230–270 m), moving deeper in the summer period (mean 340–390 m). In December–March the skate occurred at lower bottom temperatures (mean 0.8–1.6°C) whereas the rest of the year it inhabited warmer waters with mean bottom temperatures of 2.5–3.1°C. Throughout the year decreasing body weight with depth was observed, indicating that adult and juvenile mud skates inhabit different depths. Total length of mud skates in catches ranged from 17 to 70 cm with a mean of 51.71 cm. Relation between total length (TL, cm) and body weight (W, g) was: W = 0.0029TL^3.1614 (r² = 0.978). Males were more abundant among small skates (<30 cm) only, whereas females predominated among larger skates (>30 cm) comprising about 70% in 60–70 cm length class. Female mud skates were longer and heavier than males (mean length 56.9 vs 51.2 cm and mean weight 1206 vs 807 g, respectively). This species is considered to be benthophagous, consuming mostly amphipods (34.9% by weight), polychaete worms (27.6%), decapod crustaceans (12.7%), and fishery discards (13.9%). Small skates (20–40 cm) fed mostly on amphipods (85.4%); medium‐sized (40–60 cm) ate amphipods (40.2%), polychaetes (29.1%) and decapods (19.3%); the largest individuals (>60 cm) consumed fishery offal (27.9%) and less amphipods (26.6%) and polychaetes (27.7%). Preliminary data on the maturation of the species in the western Bering Sea showed that males and females become mature at lengths above 61 cm TL.     

Seamount egg‐laying grounds of the deep‐water skate Bathyraja richardsoni

Highly localized concentrations of elasmobranch egg capsules of the deep‐water skate Bathyraja richardsoni were discovered during the first remotely operated vehicle (ROV) survey of the Hebrides Terrace Seamount in the Rockall Trough, north‐east Atlantic Ocean. Conductivity–temperature–depth profiling indicated that the eggs were bathed in a specific environmental niche of well‐oxygenated waters between 4·20 and 4·55° C, and salinity 34·95–35·06, on a coarse to fine‐grained sandy seabed on the seamount's eastern flank, whereas a second type of egg capsule (possibly belonging to the skate Dipturus sp.) was recorded exclusively amongst the reef‐building stony coral Solenosmilia variabilis. The depths of both egg‐laying habitats (1489–1580 m) provide a de facto refuge from fisheries mortality for younger life stages of these skates.     

Reply to Hussey et al.: The requirement for accurate diet-tissue discrimination factors for interpreting stable isotopes in sharks

Carbon and nitrogen stable isotope analyses have improved our understanding of food webs and movement patterns of aquatic organisms. These techniques have recently been applied to diet studies of elasmobranch fishes, but isotope turnover rates and isotope diet–tissue discrimination are still poorly understood for this group. We performed a diet switch experiment on captive sandbar sharks (Carcharhinus plumbeus) as a model shark species to determine tissue turnover rates for liver, whole blood, and white muscle. In a second experiment, we subjected captive coastal skates (Leucoraja spp.) to serial salinity reductions to measure possible impacts of tissue urea content on nitrogen stable isotope values. We extracted urea from spiny dogfish (Squalus acanthias) white muscle to test for effects on nitrogen stable isotopes. Isotope turnover was slow for shark tissues and similar to previously published estimates for stingrays and teleost fishes with low growth rates. Muscle isotope data would likely fail to capture seasonal migrations or diet switches in sharks, while liver and whole blood would more closely reflect shorter term movement or shifts in diet. Nitrogen stable isotope values of skate blood and skate and dogfish white muscle were not affected by tissue urea content, suggesting that available diet–tissue discrimination estimates for teleost fishes with similar physiologies would provide accurate estimates for elasmobranchs.     

Gill Memorial Medal Address Holding the mirror up to nature

Since the early 1980s, there have been large decreases in numbers breeding in South Africa of three seabirds that compete with fisheries for food: African Penguin Spheniscus demersus, Cape Cormorant Phalacrocorax capensis and Bank Cormorant P. neglectus. By contrast, two other species that compete with fisheries but which have a larger foraging range when breeding or are nomadic between breeding localities, Cape Gannet Morus capensis and the nominate race of Swift Tern Thalasseus bergii bergii, have increased. Five seabirds that do not compete with fisheries for prey have maintained stable populations or increased. The decreases in the three species have led to deterioration in the overall conservation status of South Africa's seabirds, of which 10 of 15 species (67%) are presently classified as Threatened or Near-threatened, including six of the seven species that are endemic to the Benguela ecosystem off western southern Africa.     

Distribution patterns of the Diaptomidae (Calanoida: Copepoda) in southern Africa

All freshwater calanoids in southern Africa belong to the family Diaptomidae of which there are two subfamilies, the Paradiaptominae (Lovenula, Paradiaptomus and Metadiaptomus) and Diaptominae (Tropodiaptomus and Thermodiaptomus). The Paradiaptominae are endemic to Africa, while the Diaptominae have one endemic African genus Thermodiaptomus, and the other genus Tropodiaptomus, occurs in other parts of the world as well as Africa. Knowledge of distribution of the Diaptomidae has been hampered by incomplete taxonomic knowledge and lack of co-ordination of available information. Distribution patterns of 19 described species and 11 new species confirm that the majority of species have a limited distribution, and that the Paradiaptominae are arid-adapted and belong to temporary pool communities. Six species are endemic to coastal pans and vleis of the western and southern Cape Province. Only those species with a wider distribution have been recorded as colonising man-made lakes. The four Lovenula species are predators and often co-exist with a Metadiaptomus species as prey. Synapomorphies of widely separated species of the Paradiaptominae, provide interesting insights into how their evolution may have been governed by vicariance, dispersal and palaeoenvironments of the African continent.     

Trophic structure of a coastal fish community determined with diet and stable isotope analyses

A combination of dietary guild analysis and nitrogen (δ¹⁵N) and carbon (δ¹³C) stable‐isotope analysis was used to assess the trophic structure of the fish community in Rhode Island and Block Island Sounds, an area off southern New England identified for offshore wind energy development. In the autumn of 2009, 2010 and 2011, stomach and tissue samples were taken from 20 fish and invertebrate species for analysis of diet composition and δ¹⁵N and δ¹³C signatures. The food chain in Rhode Island and Block Island Sounds comprises approximately four trophic levels within which the fish community is divided into distinct dietary guilds, including planktivores, benthivores, crustacivores and piscivores. Within these guilds, inter‐species isotopic and dietary overlap is high, suggesting that resource partitioning or competitive interactions play a major role in structuring the fish community. Carbon isotopes indicate that most fishes are supported by pelagic phytoplankton, although there is evidence that benthic production also plays a role, particularly for obligate benthivores such as skates Leucoraja spp. This type of analysis is useful for developing an ecosystem‐based approach to management, as it identifies species that act as direct links to basal resources as well as species groups that share trophic roles.     

Characterisation of polymorphic microsatellite markers for skates (Elasmobranchii: Rajidae) from expressed sequence tags

Simple sequence repeat markers (SSRs, microsatellites) were characterised for skates (Elasmobranchii: Rajidae) from published expressed sequence tags (ESTs) of Leucoraja erinacea. These were tested in four European species (Raja clavata, Raja montagui, Dipturus batis, and Leucoraja naevus). Thirteen of the fourteen amplified loci were polymorphic in at least one species. Polymorphic loci possessed on average 4.5–5.9 alleles per species, and expected heterozygosity ranged from 0.05 to 0.88. Possible null alleles were detected at three loci, while one locus showed significant deviation from Hardy–Weinberg Equilibrium proportions. Three locus-pairs exhibited significant linkage disequilibrium in one or more species. This marker set will be valuable for population genetic analyses of the focal taxa, and may prove useful for studies of other skate species.     

Food habits of the longnose skate, <Emphasis Type="Italic">Raja rhina</Emphasis> (Jordan and Gilbert, 1880), in central California waters

Feeding studies can provide researchers with important insights towards understanding potential fishery impacts on marine systems. Raja rhina is one of the most common elasmobranch species landed in central and northern California demersal fisheries, yet life history information is extremely limited for this species and aspects of its diet are unknown. Specimens of R. rhina were collected between September, 2002 and August, 2003 from fisheries-independent trawl surveys. Percent Index of Relative Importance values indicated that the five most important prey items in 618 stomachs of R. rhina were unidentified teleosts (31.6% IRI), unidentified shrimps (19.6% IRI), unidentified euphausiids (10.9% IRI), Crangonidae (7.4% IRI), and Neocrangon resima (6.0% IRI). There were significant dietary shifts with increasing skate total length and with increasing depths. Smaller skates ate small crustaceans and larger skates ate larger fishes and cephalopods. With increasing depths, diet included bentho-pelagic teleosts and more cephalopods and euphausiids. The findings of this study are consistent with previous researchers that report similar diet shifts in skate species with size and depth.     

Biodiversity and systematics of skates (Chondrichthyes: Rajiformes: Rajoidei)

Skates (Rajiformes: Rajoidei) are a highly diverse fish group, comprising more valid species than any other group of cartilaginous fishes. The high degree of endemism exhibited by the skates is somewhat enigmatic given their relatively conserved body morphology and apparent restrictive habitat, e.g. soft bottom substrates. Skates are primarily marine benthic dwellers found from the intertidal down to depths in excess of 3,000 m. They are most diverse at higher latitudes and in deepwater, but are replaced in shallower, warm temperate to tropical waters by stingrays (Myliobatodei). The number of valid skate species has increased exponentially, with more species having been described since 1950 (n = 126) than had been described in the previous 200 years (n = 119). Much of the renaissance in skate systematics has largely been through the efforts of a few individuals who through author–coauthor collaboration have accounted for 78 of the 131 species described since 1948 and for nine of 13 genera named since 1950. Furthermore, detailed regional surveys and accounts of skate biodiversity have also contributed to a better understanding of the diversity of the skates. A checklist of the living valid skate species is presented.     

Phylogenetic placement and population structure of Indo‐Pacific bottlenose dolphins (Tursiops aduncus) off Zanzibar,Tanzania, based on mtDNA sequences

Phylogenetic placement of bottlenose dolphins from Zanzibar, East Africa and putative population differentiation between animals found off southern and northern Zanzibar were examined using variation in mtDNA control region sequences. Samples (n= 45) from animals bycaught in fishing gear and skin biopsies collected during boat surveys were compared to published sequences (n= 173) of Indo‐Pacific bottlenose dolphin, Tursiops aduncus, from southeast Australian waters, Chinese/Indonesian waters, and South African waters (which recently was proposed as a new species) and to published sequences of common bottlenose dolphin, Tursiops truncatus. Bayesian and maximum parsimony analyses indicated a close relationship between Zanzibar and South African haplotypes, which are differentiated from both Chinese/Indonesian and Australian T. aduncus haplotypes. Our results suggest that the dolphins found off Zanzibar should be classified as T. aduncus alongside the South African animals. Further, analyses of genetic differentiation showed significant separation between the T. aduncus found off northern and southern Zanzibar despite the relatively short distance (approximately 80 km) between these areas. Much less differentiation was found between southern Zanzibar and South Africa, suggesting a more recent common evolutionary history for these populations than for the northern and southern Zanzibar populations.     

Spatial Segregation in Eastern North Pacific Skate Assemblages

Skates (Rajiformes: Rajoidei) are common mesopredators in marine benthic communities. The spatial associations of individual species and the structure of assemblages are of considerable importance for effective monitoring and management of exploited skate populations. This study investigated the spatial associations of eastern North Pacific (ENP) skates in continental shelf and upper continental slope waters of two regions: central California and the western Gulf of Alaska. Long-term survey data were analyzed using GIS/spatial analysis techniques and regression models to determine distribution (by depth, temperature, and latitude/longitude) and relative abundance of the dominant species in each region. Submersible video data were incorporated for California to facilitate habitat association analysis. We addressed three main questions: 1) Are there regions of differential importance to skates?, 2) Are ENP skate assemblages spatially segregated?, and 3) When skates co-occur, do they differ in size? Skate populations were highly clustered in both regions, on scales of 10s of kilometers; however, high-density regions (i.e., hot spots) were segregated among species. Skate densities and frequencies of occurrence were substantially lower in Alaska as compared to California. Although skates are generally found on soft sediment habitats, Raja rhina exhibited the strongest association with mixed substrates, and R. stellulata catches were greatest on rocky reefs. Size segregation was evident in regions where species overlapped substantially in geographic and depth distribution (e.g., R. rhina and Bathyraja kincaidii off California; B. aleutica and B. interrupta in the Gulf of Alaska). Spatial niche differentiation in skates appears to be more pronounced than previously reported.     

The rust mycobiota of southern Africa: Species richness, composition, and affinities

The rust mycobiota (Uredinales, Basidiomycota) of southern Africa (Botswana, Namibia, and South Africa) is analysed with regard to species richness, generic composition, and similarities to the rust mycobiotas of the remaining African continent and other regions of the world. Southern Africa is home to about 546 rust species: ca 522 species have been reported from South Africa, 73 from Namibia, and less than ten from Botswana. Thirty-two species were considered to be exotics. Two hundred and twenty-five of the species are restricted to southern Africa, suggesting an endemism rate of ca 44 %. At present, the rust fungus:host ratio is 1:38.5, which is much lower than expected from other regions of the world. This low ratio may partly be due to under-exploration of the area, but the results presented here indicate that a natural paucity of rust fungi on certain, especially species-rich plant taxa centred in southern Africa and possibly environmental factors are more important reasons. The predominant genera are Puccinia and Uromyces accounting for ca 59 % of the rust species. The genera Hemileia, Phakopsora and especially Ravenelia, centred in tropical regions, are well represented and sum up to 8 % of the species. Members of Melampsoraceae and Phragmidiaceae, common in temperate regions of the Northern Hemisphere, are scarce. Most of the other 28 recorded teleomorph genera are only represented by three or less species. In an African context, most species are shared with central and east Africa (almost 16 %). Only a few species are disjunct between southern and West Africa or Madagascar. Ca 10 % of the species are shared only with other parts of the paleotropics, especially the Indian subcontinent. Disjunctions of native species with the New World, Australia/New Zealand, or Europe are rare.