Molecular analysis of Southern Ocean skates (Bathyraja) reveals a new species of Antarctic skate

Two regions of mtDNA, cytochrome b and cytochrome c oxidase subunit 1, were sequenced in nine species of Bathyraja from the Southern Ocean and New Zealand. Based on sequence divergence, the species that has been referred to as Bathyraja eatonii from the Antarctic continental shelf and slope is a species distinct from B. eatonii from the Kerguelen Plateau (the type locality) and is a new and undescribed species Bathyraja sp. (cf. eatonii). There was no sequence divergence among samples of Bathyraja sp. (dwarf) from the Ross Sea and the South Atlantic. However, for both Bathyraja sp. (cf. eatonii) and Bathyraja maccaini in the Ross Sea and the South Atlantic Ocean, the DNA sequence divergences indicate differentiation among ocean basins and within Bathyraja sp. (cf. eatonii) divergences are similar to those among recognized species of Bathyraja in the North Pacific Ocean.     

Trophic niche partitioning of five skate species of genus Bathyraja in northern and central Patagonia,Argentina

Overexploitation of marine communities can lead to modifications in the structure of the food web and can force organisms like elasmobranchs to change their feeding habits. To evaluate the impact that fisheries have on food webs and on the interactions between species, it is necessary to describe and quantify the diet of the species involved and follow it through time. This study compares the diet of five skate species using the data obtained from the by-catch of the Argentine hake (Merluccius hubbsi) fishery in north and central Patagonia, Argentina. Diet composition was assessed by analysing the digestive tract contents and trophic overlapping between species of the genus Bathyraja: Bathyraja albomaculata, Bathyraja brachyurops, Bathyraja macloviana, Bathyraja magellanica and Bathyraja multispinis. A total of 184 stomachs were analysed. The diets of B. albomaculata and B. macloviana mainly comprised annelids, whereas that of B. brachyurops primarily comprised fish, including hake heads discarded by the fishery. The diets of B. magellanica and B. multispinis were largely based on crustaceans. Despite the morphological similarities and their shared preference for benthic habitats, no complete diet overlaps were found between the different species. These results suggest that these skate species have undergone a process of diet specialisation. This is a common feeding strategy that occurs to successfully eliminate competition when resources are limited, which corresponds to the conditions found in an environment being affected by the pressures of overfishing.     

Two new North Pacific skates (Rajidae) and a revised key toBathyraja in the area

Two new species of the skates,Bathyraja hubbsi andB. pseudoisotrachys are described from the North Pacific. Among the North PacificBathyraja, B. hubbsi most closely resemblesB. violacea Suvorov in possessing a disc without thorns, a median tail thorns irregularly spaced, and minute fine prickles covering the entire dorsal surface. The two species are distinguished from each other by their dorsal color pattern, proportional measurement of tail in relation to total length, clasper components, and number of precaudal vertebrae.Bathyraja pseudoisotrachys has long been confused withB. isotrachys (Günther) by many Japanese authors, but is distinguished from other North PacificBathyraja by dorsal squamation and unique clasper components such as two clefts, terminal bridge and ridge.Bathyraja kincaidii Garman is synonymized with the valid speciesB. interrupta Gill et Townsend.Bathyraja abyssicola Gilbert andB. trachura Gilbert are redescribed in detail. Provisionally, 21 species of the genusBathyraja are recognized in the present study from this region, and a revised key is given to the North Pacific species of the genus.     

First records of advanced embryos and egg capsules of Bathyraja skates from the deep north-eastern Atlantic

Five variously developed embryos [142–279 mm total length (L_T)] from egg capsules from the Porcupine Seabight, north-eastern Atlantic (1541 m depth) are used to establish the ontogeny and early life history of Bathyraja richardsoni. The capsules of this species and two half-formed ones from the shell glands of a large (1620 mm L_T) female Bathyraja pallida taken off Ireland (c. 1900 m depth) are described and illustrated. The varying degree of yolk sac absorption found in the B. richardsoni embryos is discussed in relation to hatching size and its seeming size independence over c. 20–50 mm within the embryos' total length range. The conservative variation in external morphology with development from advanced embryos to adults among such deep stenobathic Bathyraja skates is commented upon, as is the bathymetric segregation of adults from levels in which egg capsules are deposited and young develop.     

Reproductive biology of skates, Bathyraja(Ishiyama), along the eastern Bering Sea continental slope

Eight Bathyraja skate species, Bathyraja aleutica, Bathyraja interrupta, Bathyraja lindbergi, Bathyraja maculata, Bathyraja minispinosa, Bathyraja parmifera, Bathyraja taranetzi and Bathyraja trachura, collected along the eastern Bering Sea continental slope were examined for reproductive features. The sex ratio for six of eight Bathyraja skate species studied was c. 1 : 1 with males dominating for B. interrupta and B. parmifera. Five of eight species examined, B. aleutica, B. lindbergi, B. parmifera, B. taranetzi and B. trachura approximately equalled or exceeded the maximum reported total length, L_T. Maximum L_T for three species (B. interrupta, B. minispinosa and B. parmifera) revealed that the largest individuals were males and that in three other species (B. lindbergi, B. maculata and B. trachura) both females and males grew to approximately the same size. Both females and males matured at approximately the same L_T for seven of eight species. Only female B. aleutica matured at a much larger size than males. All species matured within a narrow size range following a protracted juvenile stage and once mature these skates appeared to grow very little. First maturity for all species occurred at ≥80% of their maximum L_T. The number of oocytes did not significantly differ between the left and right ovary for any species. The total number of mature oocytes increased with L_T for six of eight species examined. The number of oocytes for one species, B. aleutica, increased in number until 145 cm L_T, then declined in those individuals greater in length. The egg cases of all species are described, of which four (B. lindbergi, B. parmifera, B. taranetzi and B. trachura) have never been previously reported. Evidence is presented that suggest skates may be limited to as little as one and as many as 13 spawning seasons, depending on the species, and that senescence may occur in older individuals.     

Skate Bathyraja spp. egg predation in the eastern Bering Sea

Predation on skate eggs by snails was examined for three skate species at seven nursery sites in three regions (north, middle and south) of the eastern Bering Sea. Mean predation levels were 6·46% for the Alaska skate Bathyraja parmifera , 2·65% for the Aleutian skate Bathyraja aleutica and 22·25% for the Bering skate Bathyraja interrupta . Predation levels were significantly higher at the middle and north sites than the south sites for all species combined. Predation levels decreased with increasing egg-case densities at all nursery sites, and the highest predation levels occurred where egg-case densities were very low. Predated egg-case density increased with increasing snail densities across all nursery sites examined. The Oregon triton Fusitriton oregonensis was the most abundant snail species at all nursery sites and displayed ability to drill holes in the egg case of B. parmifera. Holes left by predatory snails in egg cases of B. parmifera were significantly larger, and of different shape at the middle site compared to the south site. Holes in B. parmifera were also significantly larger than those in egg cases of B. interrupta across all sites examined. Egg cases of B. aleutica possess surface spines that cover the egg case and may inhibit predation by snails at a greater rate than that of the B. parmifera and B. interrupta , which have a smoother egg-case surface.     

The First Record of a Spawning Ground of the Mud Skate Bathyraja taranetzi (Dolganov, 1983) and the Okhotsk Skate B. violacea (Suvorov, 1935) in Pacific Waters off the Northern Kuril Islands

Russian Journal of Marine Biology - A spawning ground of the Mud Skate Bathyraja taranetzi and the Okhotsk Skate B. violacea was recorded for the first time from Pacific waters off the northern...     

Distribution and Size and Sex Composition of the Mud Skate Bathyraja taranetzi in the Pacific Waters off the Northern Kuril Islands and Coasts of Southeastern Kamchatka in the Summer Period

Journal of Ichthyology - In the summer period, the mud skate Bathyraja taranetzi lives at depths from ~100 to 757 m in the Pacific waters off the northern Kuril Islands and southeastern Kamchatka;...     

Molecular systematics of the skate subgenus Arctoraja (Bathyraja: Rajidae) and support for an undescribed species, the leopard skate, with comments on the phylogenetics of Bathyraja

Sequence variability in the cytochrome c oxidase I (COI) gene from 226 samples of the species previously considered Bathyraja parmifera (Rajidae) revealed three distinct haplotypes, one of which represents an undescribed species, the leopard skate. Further genetic examination of four closely related North Pacific and Bering Sea skate species, Bathyraja parmifera, B. simoterus, B. smirnovi, and the leopard skate in comparison with 19 related species indicates that together these four species comprise the subgenus Arctoraja. Phylogenetic analysis suggests that Arctoraja is monophyletic, but that the genus Bathyraja may be paraphyletic due to the phylogenetic position of Rhinoraja.     

Identification of multiple nursery habitats of skates in the eastern Bering Sea

The use of more than a single nursery habitat type is examined for oviparous elasmobranchs using data summarized from studies conducted on the Alaska skate Bathyraja parmifera and the Aleutian skate Bathyraja aleutica in the eastern Bering Sea. The eastern Bering Sea skate species use two discrete areas as nurseries, one for egg deposition and a second for newly emergent juveniles. Egg deposition sites were located along the outer shelf and upper slope near canyons in the eastern Bering Sea. Newly emergent juveniles were found along the outer and middle shelf for B. parmifera and deep‐slope for B. aleutica, suggesting that habitat used by newly emergent juvenile skates is distinct from habitat used for egg deposition and embryo development. In reviewing many studies on oviparous elasmobranchs, similar patterns emerge of habitat use during their early life history. To distinguish these distinct habitats, appropriate terminology is proposed. Egg case nursery is suggested for areas of egg deposition and juvenile nursery is suggested for areas where juveniles aggregate after emergence. Criteria to describe each habitat type are outlined.     

Effects of intrinsic and extrinsic factors on the diet of Bathyraja macloviana,a benthophagous skate

The effects of intrinsic (sex, maturity stage and body size) and extrinsic (depth and region) factors on the diet of Bathyraja macloviana, in the south‐west Atlantic Ocean, were evaluated using a multiple‐hypothesis modelling approach. Bathyraja macloviana fed mainly on polychaetes followed by amphipods, isopods and decapods. Effects of intrinsic and extrinsic factors on diet composition of this species were found. The consumption of polychaetes had a humped relationship with total length (L_T), and isopods and decapods increased with increasing L_T. Immature individuals preyed on amphipods more heavily than mature individuals. Furthermore, region and depth had an important effect on the consumption of isopods, decapods and amphipods. Such ontogenetic changes and spatial patterns may provide insights into understanding the regulatory mechanisms of marine communities.     

HERMAPHRODITIC FLOWERING IN GAMBEL OAK

Gambel oak (Quercus gambelii Nutt.) is normally monoecious, flowering once a year in spring. In Utah, August 1978, several individuals were observed producing hermaphroditic flowers in riparian habitats. This unusual event was preceded by a general freeze of male catkins in May which resulted in low acorn production. In 1979 the same individuals flowered normally, with no evidence of freeze damage or hermaphroditic flowers being observed. Acorns were produced on these individuals in both years. It is suggested that hermaphroditic flowering in the Fagaceae is an ancestral condition, still present in the genome, but normally suppressed, which may be evoked by unusual environmental conditions.     

Marginal increment analysis: a new statistical approach of testing for temporal periodicity in fish age verification

This paper proposes a new and flexible statistical method for marginal increment analysis that directly accounts for periodicity in circular data using a circular–linear regression model with random effects. The method is applied to vertebral marginal increment data for Alaska skate Bathyraja parmifera. The best fit model selected using the AIC indicates that growth bands are formed annually. Simulation, where the underlying characteristics of the data are known, shows that the method performs satisfactorily when uncertainty is not extremely high.     

Reduced recombination in gynodioecious populations of a facultative apomictic orchid

Many plants combine sexual reproduction with some form of asexual reproduction to different degrees, and lower genetic diversity is expected with asexuality. Moreover, the ratios of sexual morphs in species with gender dimorphism are expected to vary in proportion to the reproductive success of the sexual process. Hence, sex ratios can directly influence the genetic structure and diversity of a population. We investigated genotypic diversity in 23 populations of a facultative, apomictic gynodioecious orchid, Satyrium ciliatum, to examine the effect on genotypic diversity of variation in the frequency of females and in the amount of sexual reproduction. The study involved one pure female, seven gynodioecious (both females and hermaphrodites present) and 15 hermaphroditic populations. Pollinia receipt was higher in hermaphroditic than in gynodioecious populations. Analyses of variation in ISSRs demonstrated that genotypic diversity was high in all populations and was not significantly different between hermaphroditic and gynodioecious populations. We used character compatibility analysis to determine the extent to which recombination by sexual reproduction contributed to genotypic diversity. The results indicate that the contribution of recombination to genotypic diversity is higher in hermaphroditic than in gynodioecious populations, consistent with the finding that hermaphroditic populations received higher amounts of pollinia. Our finding of reduced recombination in gynodioecious populations suggests that maintenance of sex in hermaphrodites plays an important role in generating genotypic diversity in this apomictic orchid.     

The role of sex separation in neutral speciation

Neutral speciation mechanisms based on isolation by distance and assortative mating, termed topopatric, has recently been shown to describe the observed patterns of abundance distributions and species–area relationships. Previous works have considered this type of process only in the context of hermaphroditic populations. In this work, we extend a hermaphroditic model of topopatric speciation to populations where individuals are explicitly separated into males and females. We show that for a particular carrying capacity, speciation occurs under similar conditions, but the number of species generated is lower than in the hermaphroditic case. As a consequence, the species–area curve has lower exponents, especially at intermediate scales. Evolution results in fewer species having more abundant populations.     

Intra-locus sexual conflict and sexually antagonistic genetic variation in hermaphroditic animals

Intra-locus sexual conflict results when sex-specific selection pressures for a given trait act against the intra-sexual genetic correlation for that trait. It has been found in a wide variety of taxa in both laboratory and natural populations, but the importance of intra-locus sexual conflict and sexually antagonistic genetic variation in hermaphroditic organisms has rarely been considered. This is not so surprising given the conceptual and theoretical association of intra-locus sexual conflict with sexual dimorphism, but there is no a priori reason why intra-locus sexual conflict cannot occur in hermaphroditic organisms as well. Here, I discuss the potential for intra-locus sexual conflict in hermaphroditic animals and review the available evidence for such conflict, and for the existence of sexually antagonistic genetic variation in hermaphrodites. I argue that mutations with asymmetric effects are particularly likely to be important in mediating sexual antagonism in hermaphroditic organisms. Moreover, sexually antagonistic genetic variation is likely to play an important role in inter-individual variation in sex allocation and in transitions to and from gonochorism (separate sexes) in simultaneous hermaphrodites. I also describe how sequential hermaphrodites may experience a unique form of intra-locus sexual conflict via antagonistic pleiotropy. Finally, I conclude with some suggestions for further research.     

Differences in sex expression and mating systems in three pollination morphs of Cimicifuga simplex

Cimicifuga simplex (Ranunculaceae) has three genetically distinct pollination morphs. Here, we report that each of the three pollination morphs of C. simplex differs from the others with regard to sex expression and mating system: morph I consists mostly of ramets with hermaphroditic flowers and ramets with only female flowers, morph II consists of ramets with hermaphroditic flowers and ramets with hermaphroditic and male flowers, and morph III consists mostly of ramets with hermaphroditic flowers. A microsatellite analysis of seed DNA showed that morph III has a high self-fertilization rate. Flowering season and flower visitor assemblages, which also differ among the three morphs, may influence the evolution and maintenance of the differences in sex expression and mating systems in the morphs.     

Organisation of the Hb 1 genes of the Antarctic skate Bathyraja eatonii: new insights into the evolution of globin genes

An extensive investigation of the organisation of globin genes has greatly contributed to the understanding of universal mechanisms of gene evolution and expression. Cartilaginous fish are the first organisms that have evolved the tetrameric form of hemoglobin (Hb). So far, there has been absolute lack of data about globin genes in chondrichthyans. Bathyraja is the dominant rajid south of 60 degrees S. In the framework of the investigations on globin genes of Antarctic red-blooded and Hb-less fish we obtained the cloning of the alpha- and beta-globin cDNAs of the main Hb (Hb 1) of the skate Bathyraja eatonii. Then, a genomic fragment of 6.2 kb was isolated where the Hb 1 alpha and beta genes are linked in a tail-to-head (3' to 5') orientation. The beta-globin gene promoter region and the chromosomal organisation of the Hb 1 genes of B. eatonii have been compared to their homologues in other vertebrates. The finding of a tail-to-head linkage of the Hb 1 alpha- and beta-globin genes in B. eatonii is the first characterisation of the organisation of globin genes in chondrichthyes; such finding offers a novel contribution to the understanding of the evolution of this class of genes. Moreover, the characterisation of chondrichthyan genes is very important for gaining insight into the ancestral state of vertebrate genomes.     

The Effect of Breeding System on Polymorphism in Mitochondrial Genes of Silene

Gynodioecy is a breeding system characterized by the co-occurrence of hermaphrodite and female individuals, generally as the result of nuclear–cytoplasmic interactions. The question remains whether the genetic factors controlling gynodioecy are maintained in species over long evolutionary timescales by balancing selection or are continually arising and being replaced in epidemic sweeps. If balancing selection maintains these factors, then neutral cytoplasmic diversity should be greater in gynodioecious than hermaphroditic species. In contrast, epidemic sweeps of factors controlling gynodioecy should decrease cytoplasmic diversity in gynodioecious relative to hermaphroditic species. We took a comparative approach in which we sequenced two mitochondrial genes, cytochrome b (cob) and cytochrome oxidase (cox1), for multiple populations of several hermaphroditic, gynodioecious, and dioecious species in the genus Silene. Breeding system was predictive of polymorphism. Gynodioecious species harbor many old haplotypes while hermaphroditic and dioecious species have little to no nucleotide diversity. The genealogical structure of neither gene departed from neutral expectations. Taken together, our results suggest that balancing selection acts on cytoplasmic male-sterility factors in several gynodioecious species in the genus.