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.     

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.     

Molecular barcoding of skates (Chondrichthyes: Rajidae) from the southern Northeast Atlantic

Serra‐Pereira, B., Moura, T., Griffiths, A. M., Gordo, L. S. & Figueiredo, I. (2010). Molecular barcoding of skates (Chondrichthyes: Rajidae) from the southern Northeast Atlantic. —Zoologica Scripta, 40, 76–84. Due to their vulnerability to fishing pressure, many species of skate (Rajidae) in the Northeast Atlantic are undergoing declines in abundance. The assessment of stock status and subsequent proposal of management measures are often complicated by high levels of species diversity and endemism, coupled with morphological and ecological conservatism, which makes distinguishing between species difficult. To improve the identification of skates and investigate the phylogenetic position of endemic species the cytochrome c oxidase subunit I (COI) was sequenced in 12 species (Dipturus oxyrinchus, Leucoraja naevus, Leucoraja circularis, Neoraja iberica, Raja brachyura, Raja clavata, Raja maderensis, Raja microocellata, Raja miraletus, Raja montagui, Raja undulata, Rostroraja alba) inhabiting the Portuguese waters. Based on sequence divergence R. maderensis and R. clavata only differ by 1% of the 652 bp COI sequence, questioning the recognition of R. maderensis (considered to be endemic to Madeira and the Azores), as a reproductively isolated species. Otherwise, there was clear phylogenetic support for the different genera and all the remaining species, although the genetic divergence was low compared to other chordates. In particular, COI analysis allowed clear identification of the morphologically similar species R. brachyura and R. montagui.     

Chlorophyll-nutrient relationships in North Island lakes (New Zealand)

A model for the prediction of chlorophyll a in the near surface waters (1 m) of North Island lakes was developed using data from the literature and our own study of 12 North Island lakes. Annual geometric mean concentrations of chlorophyll a, total nitrogen and total phosphorus were used since no distinct growing season was discernable. Annual mean ratios of total nitrogen to total phosphorus in the near surface waters ranged between 10 and 59 (by weight). Strong correlations were obtained between log-transformed values of chlorophyll a and total nitrogen (r² = 0.53, n = 16), chlorophyll a and total phosphorus (r² = 0.71, n = 21), and between total nitrogen and total phosphorus (r² = 0.69, n = 16). However, after correcting for the high interdependency between total nitrogen and total phosphorus, only total phosphorus was found to be important in predicting chlorophyll concentrations. Much of the variance in the chlorophyll-phosphorus relation was attributable to differences in mean lake depth. Lakes with mean depths less than 11 m had significantly more chlorophyll a per unit of total phosphorus ( = 0.54 µg · µg^–1, SE = 0.05, n = 6) than lakes of greater mean depth ( = 0.17 µg · µg^–1, SE = 0.02, n = 14). When the effect of mean depth was taken into account, 89% of the variance in chlorophyll a was explained compared with 71% for the simple linear regression on total phosphorus alone.     

Nothofagus (Fagaceae) and its invertebrate fauna—an overview and preliminary synthesis

The invertebrates directly associated with the southern hemisphere tree genus Nothofagus are described and discussed in terms of their organization into guilds and attributes of their hostplants. Using literature records, comparisons are made of the guild composition between host species and provenances; a full invertebrate-hostplant list is appended. Sap-sucking Homoptera are particularly diverse and usually host-species specific whereas defoliating insects seem less prevalent and less specific. Few seed-eating insects are reported. Some provenances and hostplants exhibit local radiations in certain taxa, such as Eriococcidae in New Zealand, Notophorina psyllids in South America and scolytid beetles on N. dombeyi. Environmental stresses such as drought, disturbance and landslips, can induce outbreaks of certain scale insects and borers, affecting the dynamics of the forest. Generally, however, the influence of invertebrates appears benign, but occasional large-scale defoliation is observed on N. solandri var. cliffortioides, N. moorei, N. antarctica and N. pumilio. Collectively this fauna has two elements: a geographically recurrent Gondwanic element, usually monophagous, and a provenance-specific element (often polyphagous) locally recruited following vicariance of the host genus. The fauna of Nothofagus can be in stark contrast to that of adjacent forest types. In Australia, the insect profile of Nothofagus has more in common with fagaceous trees on other continents than the more recent Eucalyptus forests which surround it. There is almost no evidence of fauna transfer from Nothofagus to Eucalyptus. Relative to New Zealand, the Australian Nothofagus fauna is depauperate and may reflect truncation in Pleistocene glaciations. Typically low insect populations on these trees may partly explain the limited range of insectivorous birds and spiders observed in Australia. Much more data on the phytophagous insects of this biome is needed for comparative purposes. The fauna of the montane tropical subgenus Brassospora pollen group is almost unknown but could hold the key to the origin of fagaceous insect communities more generally. The conservation value of remaining Nothofagus forests is enhanced by recognition of their co-adapted fauna, some of which are arranged in stable functional groups likely to give valuable insight into invertebrate-hostplant interactions originating in the Cretaceous. Elucidation of its role in the processes of these unique forests remains a fertile field for endeavour in the future.     

Walking in skates (Chondrichthyes, Rajidae): anatomy, behaviour and analogies to tetrapod locomotion

Pelvic fin walking in skates is common. However, the structure and function of pelvic fins have not been analysed. Pelvic fins of skates of the genus Psammobatis and Rioraja agassizi are externally divided into an anterior leg-like lobe and a posterior fin-like lobe. Internally, the anterior lobes are supported by a compound radial, a proximal radial and distal radials that resemble a thigh, a calf and a foot, respectively, and three associated radials arising from the pelvic girdle. A highly developed radial condyle on the pelvic girdle enables broad 'limb' movements. The muscular arrangement of the anterior lobes is formed by protractor, retractor, flexor and extensor muscles, clearly departing from the generalized fin muscle arrangement of elasmobranchs. Walking is composed of propulsion and recovery phases. A backward movement of the compound radial in the horizontal plane characterizes the propulsive phase. The proximal radial connects vertically the compoundradial with the foot-like distal radials, which are anchored on the bottom. During the recovery phase, the foot-like structure is lifted off the bottom and the entire limb-like anterior lobe is moved forwards for starting a new cycle. Walking in skates resembles the ancestral tetrapod sprawling locomotion seen in many salamanders and lizards. Pelvic fin anatomy and walking behaviour in skates and hemiscylliid sharksare compared. Ecological and evolutionary implications of walking locomotion in skates are also discussed.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 35–41.     

Distribution and movements of Buller's albatross (Diomedea bulleri) in Australasian seas

Abstract

The distribution and movements of Buller's albatross in Australasian seas are analysed using results of shipborne surveys (13 238 10‐min counts), counts from trawlers, banding data, recoveries on beaches and fishing vessels, and records from the literature. Patterns of marine distribution are documented by monthly accounts and maps. During the breeding season, highest abundances are recorded over shelves and slopes off southern New Zealand (The Snares shelf to 41–43°S off the South Island, D. b. bulleri), around the Chatham Islands and over oceanic subtropical waters east of New Zealand (probably D. b. platei), with marked seasonal variations observed off southern New Zealand. Both subspecies disperse mostly outside Australasian waters during the non‐breeding season. Birds banded on The Snares were recovered off south‐eastern New Zealand (Stewart Island to Cook Strait) and in the eastern tropical Pacific. Immatures accounted for only 0.25% of birds censused during the ship‐borne surveys; they are recorded around the New Zealand mainland in August‐October and February‐May, off south‐eastern Australia and in the Tasman Sea in November‐December, February, and June‐July. Around New Zealand, males predominate among birds recovered along the eastern seaboard, whereas the sex ratio in south‐western waters tends to vary according to water depth and season. Distribution patterns and movements in New Zealand and Australian seas are discussed in relation to breeding events and breeding status.     

Possible occurrence of character displacement in the sympatric skates Raja erinacea and R. ocellata (Pisces: Rajidae)

Synopsis Raja erinacea and R. ocellata are sibling species which are positively correlated with each other by occurrence and numerical abundance. In sympatry the species undergo interactive segregation; R. erinacea feeds on a higher percentage of epifauna and R. ocellata feeds on a higher percentage of infauna.An isolated allopatric population of R. ocellata occurs in the Gulf of St. Lawrence which is phenotypically intermediate between the sympatric populations of R. erinacea and R. ocellata in characters related to feeding e.g. size, number of tooth rows in the upper jaw, and shape of the upper jaw. It appears probable that the allopatric population represents the morphological state of R. ocellata before it became sympatric with R. erinacea; divergence in size, number of tooth rows, and shape of the upper jaw between the two species developed after establishment of sympatry. These divergences in character traits, related to feeding, reduced competition between the two sympatric species and permitted the present wide overlap in their ranges.Character displacement is evidently rare in demersal fishes inhabiting the flat and soft bottoms of the northwestern Atlantic because the three other pairs of sibling species that occur there are parapatrically distributed and thus would not compete for resources. Raja erinacea and R. ocellata may have been restrained from establishing parapatry by another species pair of skates (R. senta and R. radiata) which have a complementary distribution and similar feeding habits of R. erinacea and R. ocellata but which occur in deeper water.     

Evolutionary and biogeographical history of an ancient and global group of arachnids (Arachnida: Opiliones: Cyphophthalmi) with a new taxonomic arrangement

We investigate the phylogeny, biogeography, time of origin and diversification, ancestral area reconstruction and large‐scale distributional patterns of an ancient group of arachnids, the harvestman suborder Cyphophthalmi. Analysis of molecular and morphological data allow us to propose a new classification system for the group; Pettalidae constitutes the infraorder Scopulophthalmi new clade , sister group to all other families, which are divided into the infraorders Sternophthalmi new clade and Boreophthalmi new clade . Sternophthalmi includes the families Troglosironidae, Ogoveidae, and Neogoveidae; Boreophthalmi includes Stylocellidae and Sironidae, the latter family of questionable monophyly. The internal resolution of each family is discussed and traced back to its geological time origin, as well as to its original landmass, using methods for estimating divergence times and ancestral area reconstruction. The origin of Cyphophthalmi can be traced back to the Carboniferous, whereas the diversification time of most families ranges between the Carboniferous and the Jurassic, with the exception of Troglosironidae, whose current diversity originates in the Cretaceous/Tertiary. Ancestral area reconstruction is ambiguous in most cases. Sternophthalmi is traced back to an ancestral land mass that contained New Caledonia and West Africa in the Permian, whereas the ancestral landmass for Neogoveidae included the south‐eastern USA and West Africa, dating back to the Triassic. For Pettalidae, most results include South Africa, or a combination of South Africa with the Australian plate of New Zealand or Sri Lanka, as the most likely ancestral landmass, back in the Jurassic. Stylocellidae is reconstructed to the Thai‐Malay Penisula during the Jurassic. Combination of the molecular and morphological data results in a hypothesis for all the cyphophthalmid genera, although the limited data available for some taxa represented only in the morphological partition negatively affects the phylogenetic reconstruction by decreasing nodal support in most clades. However, it resolves the position of many monotypic genera not available for molecular analysis, such as Iberosiro, Odontosiro, Speleosiro, Managotria or Marwe, although it does not place Shearogovea or Ankaratra within any existing family. The biogeographical data show a strong correlation between relatedness and formerly adjacent landmasses, and oceanic dispersal does not need to be postulated to explain disjunct distributions, especially when considering the time of divergence. The data also allow testing of the hypotheses of the supposed total submersion of New Zealand and New Caledonia, clearly falsifying submersion of the former, although the data cannot reject the latter. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 92–130.     

Phylogenetic investigation and divergence dating of Poa (Poaceae,tribe Poeae) in the Australasian region

The Australasian region contains a significant proportion of worldwide Poa diversity, but the evolutionary relationships of taxa from this region are incompletely understood. Most Australasian species have been placed in a monophyletic Poa subgenus, Poa supersection Homalopoa section Brizoides clade, but with limited resolution of relationships. In this study, phylogenetic relationships were reconstructed for Australasian Poa, using three plastid (rbcL and matK genes and the rpl32‐trnL intergenic spacer) and two nuclear [internal/external transcribed spacer (ITS/ETS)] markers. Seventy‐five Poa spp. were represented (including 42 Australian, nine New Guinean, nine New Zealand and three Australian/New Zealand species). Maximum parsimony, maximum likelihood and Bayesian inference criteria were applied for phylogenetic reconstruction. Divergence dates were estimated using Bayesian inference, with a relaxed clock applied and rates sampled from an uncorrelated log‐normal distribution. Australasian Poa spp. are placed in three lineages (section Brizoides, section Parodiochloa and the ‘X clade’), each of which is closely related to non‐Australasian taxa or clades. Section Brizoides subsection Australopoa is polyphyletic as currently circumscribed. In Australasia, Poa has diversified within the last 4.3 Mya, with divergence dating results broadly congruent with fossil data that record the appearance of vegetation with a prominent grassland understorey or shrubland/grassland mosaic vegetation dating from the mid‐Pliocene. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 523–552.     

Leaf domatia and mites on Australasian plants: ecological and evolutionary implications

Leaf domatia, specialized chambers in the vein axils on the underside of leaves of many plant species, have remained an enigma for over a century. In this study we show a strong association between foliar domatia and mites in 37 plant species in Australasia. Overall, mites accounted for 91% of the arthropods observed in domatia. Across all species, a median of 51% of domatia were occupied and 71% of leaves showed mite evidence in domatia. The level of mite association did not depend on domatia type (pit, pouch, pocket, or tuft) or provenance (Papua New Guinea, Queensland, Victoria, or New Zealand). Mite association with domatia commonly varied between plant species, between individuals within species, and between shoots within individuals. The leaf developmental stage probably explains much of the variation in association for many of these species. The presence of a variety of life history stages of mites within domatia indicates that these structures act as shelters for development and reproduction. Furthermore, in 12 of 13 plant species examined, domatia concentrate mites in particular locations on the leaf. Mite taxa that we classify as largely predaceous (e.g. phytoseiids, stigmaeids and tydeids) or fungivorous (e.g. acarids and oribatids) were most common in domatia and dominated the association in 21 of 24 plant species in which the relative abundance of herbivorous, fungivorous and predaceous groups was quantified. We evaluate hypotheses that explain the role of leaf domatia, including non-functional hypotheses (e.g. architectural constraints), physiological function (e.g. gas exchange and water uptake), bacterial symbiosis and antagonistic and mutualistic associations with mites. Our quantitative results confirm anecdotal accounts of mite association with leaf domatia and are most consistent with Lundströem's century-old hypothesis of plant-mite mutualism in which leaf domatia billet predaceous and fungivorous mites that prey on plant enemies. Leaf domatia are widespread among woody angiosperms and abundant in many temperate and tropical regions of Australasia. Mites, an ancient group of arthropods whose diversity and abundance parallels that of insects, are likely to be important selective agents on terrestrial plants. Our results (1) indicate that mite-domatia association represents a relationship of comparable scope to plant-ant associations mediated by specialized plant structures such as extrafloral nectaries, food bodies and specialized domatia; (2) suggest that sociality is not a necessary prerequisite for widespread and diverse mutualisms between arthropods and plants; and, (3) extend the diversity of organisms that produce specialized mite ‘houses’ from lizards, and wasps and bees to woody angiosperms.     

Development of molecular and morphological markers to improve species-specific monitoring and systematics of Northeast Atlantic and Mediterranean skates (Rajiformes)

The Northeast Atlantic and Mediterranean skates (Rajidae) showed remarkable species diversity but with high morphological and ecological conservatism. Since skates are particularly vulnerable to the bottom trawl fishery, species-specific demographic surveys as well as studies defining life history and evolutionary traits are important in prioritising conservation programs. However, the identification of juveniles and adults of some species may be difficult using referenced guidelines and identification keys. Therefore, we attempt to develop markers for species identification through the parallel analysis of a 16S rDNA gene sequence and of several morphological characters on 135 individuals collected by trawl surveys in the Adriatic Sea and putatively assigned to six taxa. Species-specific haplotypes were defined for Raja miraletus, Raja montagui, Dipturus oxyrinchus, since a solid accordance between species boundaries and well-differentiated haplotypes was observed. Comparative analysis of 16S rDNA sequences allowed the identification of three juvenile specimens of Leucoraja circularis, a species that rarely occurs in the Adriatic Sea. On the contrary, morphological traits and haplotype distribution were largely discordant in Raja asterias and R. clavata. While all putative R. clavata individuals showed a unique haplotype (H-CLA), only 8 of 30 putative R. asterias individuals possessed a second weakly divergent haplotype (H-AST). The remaining 22 R. asterias carried the H-CLA. The multivariate analyses of morphometric and meristic characters in putative R. clavata and R. asterias revealed the clustering of individuals regardless of haplotypes. However, a bimodal distribution of R. asterias and R. clavata samples would suggest that two separated taxa might exist, both sharing the two 16S rDNA haplotypes. The haplotype distribution appeared to be significantly correlated only to the standardised disc length/total length (DL/TL) variation. Three alternative explanations may support this scenario: (i) an incomplete lineage sorting process in two morphologically yet distinct taxa; (ii) a recent hybridisation between the two taxa; (iii) the two taxa are morphologically plastic species and all considered morphological characters may be misleading in discriminating between them at all maturity stages, except for the DL/TL. However, further analyses on larger data sets and using molecular key markers (i.e. nuclear genes) will be needed to definitely resolve the status of these taxa. Molecular relationships among rajid taxa are largely consistent with systematics based on internal and external anatomical features. This multidisciplinary study contributed to defining the pattern of species diversity and abundance of rajids in the Adriatic Sea.     

Spatio-temporal distribution of sperm whales (Physeter macrocephalus) off Kaikoura,New Zealand,in relation to bathymetric features

Using a shore-based station we monitored the position of sperm whales (Physeter macrocephalus) within the Kaikoura submarine canyon from 2010 to 2012. We tracked sperm whales using a theodolite station for a total of 290 days. We extracted the distance from the nearest coast, the depth and the bathymetric slope using ArcGIS 10.1. We estimated the seasonal spatial distribution of sperm whales using general additive models. The distribution varied significantly between seasons; individuals were found in deeper water and further offshore in the spring than in winter. This study improved our understanding of the variability of sperm whale distribution patterns off Kaikoura. We determined that the distribution was linked to the bathymetric features and we hypothesized that whales adapted their use of the submarine canyon in relation to food aggregation. We would encourage further studies to evaluate the sperm whale relationship with oceanographic variables off Kaikoura.     

Process and pattern in the biogeography of New Zealand – a global microcosm?

Aim  To describe New Zealand's historical terrestrial biogeography and place this history in a wider Southern Hemisphere context.
Location  New Zealand.
Methods  The analysis is based primarily on literature on the distributions and relationships of New Zealand's terrestrial flora and fauna.
Results  New Zealand is shown to have a biota that has broad relationships, primarily around the cool Southern Hemisphere, as well as with New Caledonia to the north. There are hints of ancient Gondwanan taxa, although the long-argued predominance of taxa derived by vicariant processes, driven by plate tectonics and the fragmentation of Gondwana, is no longer accepted as a principal explanation of the biota's origins and relationships.
Main conclusions  Most of the terrestrial New Zealand flora and fauna has clearly arrived in New Zealand much more recently than the postulated separation of New Zealand from Gondwana, dated at c. 80 Ma. There is a view that New Zealand may have disappeared completely beneath the sea in the early Cenozoic, and acceptance of this would mean derivation of the entire biota by transoceanic dispersal. However, there are elements in the biota that seem to have broad distributions that date back to Gondwanan times, and also some that are thought unlikely to have been able to disperse to New Zealand across ocean gaps, especially freshwater organisms. Very strong connections to the biota of Australia, rather than to South America, are inconsistent with the timing of New Zealand's ancient and early separation from Gondwana and seem likely to have resulted from dispersal.     

Molecular Organization of 5S rDNAs in Rajidae (Chondrichthyes): Structural Features and Evolution of Piscine 5S rRNA Genes and Nontranscribed Intergenic Spacers

The genomic and gene organisation of 5S rDNA clusters have been extensively characterized in bony fish and eukaryotes, providing general issues for understanding the molecular evolution of this multigene DNA family. By contrast, the 5S rDNA features have been rarely investigated in cartilaginous fish (only three species). Here, we provide evidence for a dual 5S rDNA gene system in the Rajidae by sequence analysis of the coding region (5S) and adjacent nontranscribed spacer (NTS) in five Mediterranean species of rays (Rajidae), and in a large number of piscine taxa including lampreys and bony fish. As documented in several bony fish, two functional 5S rDNA types were found here also in the rajid genome: a short one (I) and a long one (II), distinguished by distinct 5S and NTS sequences. That the ancestral piscine genome had these two 5S rDNA loci might be argued from the occurrence of homologous dual gene systems that exist in several fish taxa and from 5S phylogenetic relationships. An extensive analysis of NTS-II sequences of Rajidae and Dasyatidae revealed the occurrence of large simple sequence repeat (SSR) regions that are formed by microsatellite arrays. The localization and organization of SSR within the NTS-II are conserved in Rajiformes since the Upper Cretaceous. The direct correlation between the SSRs extension and the NTS length indicated that they might play a role in the maintenance of the larger 5S rDNA clusters in rays. The phylogenetic analysis indicated that NTS-II is a valuable systematic tool limited to distantly related taxa of Rajiformes. Electronic Supplementary Material Electronic Supplementary material is available for this article at and accessible for authorised users. [Reviewing Editor: Dr. Rafael Zardoya]     

<Emphasis Type="Italic">Hongeo</Emphasis>, a new skate genus (Chondrichthyes: Rajidae), with redescription of the type species

Hongeo gen. nov. is proposed following examinations of several individuals, including adult males, of the type species Raja koreana, previously described from a single adult female specimen from the southern coast of the Korean Peninsula. A number of external (including claspers, squamation, and ampullary and lateral line canal systems) and skeletal characters are newly described. The phylogenetic position of R. koreana among 26 nominal rajid supraspecific taxa plus 2 unnamed supraspecific taxa of Rajidae and 3 outgroups was estimated. According to a strict consensus tree of 20 equally parsimonious trees of 139 steps (CI = 0.626; RI = 0.850; RC = 0.532; HI = 0.374), Hongeo gen. nov. is defined by the two derived character states: rostral cartilage continuous with neurocranium and stout proximally, but very slender and uncalcified distally; and anterior fontanelle broad, extending forward onto the basal part of rostral cartilage slightly beyond the leading edge of nasal capsules. The new genus also differs from all other supraspecific taxa of Rajidae in the following combination of characters: dorsal surface of tail with a row of distinct thorns along midline in both sexes; most thorns directed anteriorly; a pair of longitudinally elongated black blotches having undulated contours on middle of dorsal surface of disc; three groups of outer buccal ampullae and tubules; three groups of nasal ampullae and tubules; hyoidean ampullae and tubules on ventral surface posteriorly extending past cloaca; rostral shaft narrow with filamentous cartilage at base; scapulocoracoid comparatively short and high with dorsoventrally elliptical anterior fenestra and expanded postventral fenestra; external margin of mesopterygium of pectoral fin slightly undulated, not sinuous, and not fused with pectoral radials; and clasper component eperon and pent present.     

Redescription of Closterocerus cruy (Girault) (Hymenoptera: Eulophidae: Entedoninae), with new geographical and host records

Abstract  Closterocerus cruy (Girault) is newly recorded, both from New Zealand and as a parasitoid of lepidopteran leafminers. The species is redescribed from the Girault type and from New Zealand material. Diagnostic character states are provided for the separation of C. cruy from the closely related Australian agromyzid parasitoid C. mirabilis Edwards and LaSalle.     

New species and global revision of Hippasteria (Hippasterinae: Goniasteridae; Asteroidea; Echinodermata)

A molecular phylogenetic analysis of the genus Hippasteria, rooted against Evoplosoma, has provided the basis for taxonomic revisions and provided insight into the biogeography of a widely occurring, cold‐water group of corallivorous asteroids. Herein, we describe three new species, H ippasteria mcknighti sp. nov. , H ippasteria muscipula sp. nov. , and H ippasteria tiburoni sp. nov. , from deep‐water settings. Additionally, in light of taxonomic changes, we further elaborate on distribution data for multiple species. Range extensions for Hippasteria phrygiana and Hippasteria californica are included. Discussions about biogeography, cladogenic events, and morphology are also presented. © 2014 The Linnean Society of London     

Biogeographical affinities of the New Caledonian biota: a puzzle with 24 pieces

Aim The distributions of many New Caledonian taxa were reviewed in order to ascertain the main biogeographical connections with other areas. Location Global. Methods Panbiogeographical analysis. Results Twenty‐four areas of endemism (tracks) involving New Caledonia and different areas of Gondwana, Tethys and the central Pacific were retrieved. Most are supported by taxa of lower and higher plants, and lower and higher animals. Main conclusions Although parts of New Caledonia were attached to Gondwana for some time in the mid‐Cretaceous, most of the New Caledonian terranes formed as oceanic island arcs and sections of sea floor bearing seamounts. The flora and fauna have evolved and survived for tens of millions of years as metapopulations on ephemeral islands. Later, the biotas were juxtaposed and fused during terrane accretion. This process, together with the rifting of Gondwana, explains the biogeographical affinities of New Caledonia with parts of Gondwana, Tethys and the Pacific.     

Characterization and isolation of DNA microsatellite primers in Raja clavata L. (thornback ray,Rajidae)

The thornback ray, Raja clavata, is an elasmobranch (cartilaginous fish). Since the 1950s, its stock has severely declined. In order to investigate the genetic population structure of the species, we developed microsatellite loci. The five loci reported here have eight to 48 alleles per locus and display an observed heterozygosity ranging from 0.32 to 0.98 with no deviation from the Hardy–Weinberg equilibrium. In the test panel of 122 individuals from three populations, no null alleles, band‐stuttering, large allele dropouts or linkage disequilibrium was detected.