Unique digestive system,trophic specialization,and diversification in the deep‐sea gastropod genus Scaphander

Dietary specialization is known to be important for the evolution of Cephalaspidea gastropods, but still little is known about the overall trophic interactions of the group and the putative role of trophic ecology on diversification. The genus Scaphander is a group of predominantly deep‐sea, infaunal cephalaspids with about 40% of its species (eight) occurring on the Atlantic Ocean. They are carnivorous and have a unique digestive system with a large, strongly muscularized gizzard containing three sizable and heavily calcified plates. This work aims to describe the diet of Scaphander, to evaluate if there is a functional relation between the anatomy of the digestive tract and exploitation of novel food resources, and to assess if dietary specialization may have played a role in the diversification of the Atlantic species of Scaphander. Gut contents were studied from 31 specimens representing seven of the Atlantic species using scanning electron microscopy and light microscopy. The chemical composition of the gizzard plates was analysed by X‐ray microanalysis and X‐ray powder diffraction. Foraminiferans, including agglutinating forms, were shown to be the most important food item for Scaphander; bivalves, gastropods, scaphopods, and polychaetes with calcareous tubes were also found to form part of the diet. The gizzard plates were shown to consist of a phosphate‐rich amorphous component and a crystalline component identified as fluorite (hardness 4; Moh's scale). It is suggested that the ability of Scaphander gastropods to prey upon organisms with hard tests, particularly agglutinating foraminiferans, is not only due to the hardness of the gizzard plates but to the cumulative effect of shape and hardness of the gizzard plates and relative size of the gizzard and associated musculature. No interspecific differences were found in the diet and morphology of the digestive tract, indicating that dietary specialization probably has not played a significant role in the diversification of Atlantic species of Scaphander. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 512–525.     

Revision of the bathyal cusk-eels of the genus Bassogigas (Ophidiidae) with description of a new species from off Guam, west Pacific Ocean

The bathyal genus Bassogigas (Teleostei: Ophidiidae) is revised based on 25 specimens, 18 from the west Atlantic Ocean and seven from the Indo-west Pacific Ocean. One specimen, from off Guam, west Pacific Ocean, represents a new species, Walker's cusk eel Bassogigas walkeri. The other 24 specimens all belong to the type species, Gills cusk eel Bassogigas gillii. A comparison between the Atlantic and the Indo-west Pacific Ocean specimens of B. gillii showed no differences in meristic and morphometric characters, but in two of the Indo-west Pacific Ocean specimens the sagittal otolith varied somewhat from the remaining specimens. The two Bassogigas species differ in the length of the lateral line, the number of scales in the midline of the body, the form of the median basibranchial tooth patches and in the thickness of the otolith.     

Systematic revision and molecular phylogeny of the land snail genus Fruticocampylaea (Gastropoda: Hygromiidae) from the Caucasus region

This paper presents a systematic revision and a molecular phylogenetic analysis of the Caucasian land snail genus Fruticocampylaea. The genus is newly delimited based on the reduction of the cavities adjoining the seminal duct in the penial papilla. Shell and genitalia of all five species (F. narzanensis, F. kobensis, F. tushetica sp. nov., F. christophori, F. daghestana) are described and figures provided. All synonyms and all locality records are listed. Maximum likelihood and Bayesian analyses of mitochondrial and nuclear DNA sequences (fragments of cox1, 16S rDNA, ITS2 and 28S rDNA) confirm the monophyly of Fruticocampylaea. The reduction of the dart apparatus and the conical plug, via which the dart apparatus inserts into the vagina, as well as the molecular phylogenetic analyses, suggests a sister group relationship between Fruticocampylaea and Circassina (without Abchasohela). Furthermore, the molecular phylogenetic analyses indicate that the Fruticocampylaea species originated in a rapid radiation. The uplift of the Greater Caucasus in the Late Miocene or Pliocene or climatic changes at the end of the Pliocene or in the early Pleistocene may have caused the radiation of Fruticocampylaea. Low intraspecific variability can be explained by population bottlenecks during Pleistocene glacial periods followed by postglacial population increase.

http://zoobank.org/urn:lsid:zoobank.org:pub:AB15158D-21A3-4945-8D49-F7DE8E406E2B     

New representatives of the genus Ancorabolina George, 2006 (Copepoda,Harpacticoida, Ancorabolidae) including remarks on ancorabolid phylogeny

Four new species of Ancorabolina George, 2006 (Copepoda: Harpacticoida: Ancorabolidae) are described: Ancorabolina belgicae sp. nov. (from the Porcupine Seabight, north‐east Atlantic Ocean), Ancorabolina anaximenesi sp. nov. and Ancorabolina galeata sp. nov. (both from Anaximenes Mountain, eastern Mediterranean Sea), and Ancorabolina divasecunda sp. nov. (from Cape and Guinea Basins, south‐east Atlantic Ocean). Furthermore, the generic diagnosis for Ancorabolina is provided and the position of the taxon within Ancorabolidae is discussed. The characters originally employed to allocate Ancorabolina to the subfamily Ancorabolinae Sars, 1909, are critically evaluated and two true autapomorphies for a monophyletic Ancorabolinae (including Ancorabolina) are recognized. Two autapomorphies supporting Ancorabolina are proposed: (1) cephalothorax posteriorly with pair of ventrolateral cuticular processes turned backwards and (2) loss of the third setal element subapically on the antennary endopod. The urgent need for a phylogenetic re‐evaluation of the paraphyletic subfamily Laophontodinae Lang, 1944 is stressed and evidenced by three characters widespread in this subfamily but also present in Ancorabolina. This may be an indication of a closer relationship of Ancorabolina with certain members of Laophontodinae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 16–55.     

Systematic revision of the living species of Bullidae (Mollusca: Gastropoda: Cephalaspidea), with a molecular phylogenetic analysis

Bullidae are a worldwide family of marine shelled cephalaspidean gastropods with a mainly tropical distribution, but also with some representatives in temperate waters. The taxonomy of the group has in the past been based only on shell characters, and the few anatomical accounts available have not addressed more than one to three species, so there has been no agreement about the number of valid species. Seventy‐two specific names and 16 varietal names have been proposed worldwide. The systematics of the family Bullidae are revised, based not only on shells but also on anatomy of all extant species and on DNA sequence data. Twelve species are recognized worldwide, including one new species here described, and all are assigned to the genus Bulla. Two species occur in the eastern Atlantic, B. striata and B. mabillei; two in the western Atlantic, B. occidentalis and B. solida; two in the eastern Pacific, B. gouldiana and B. punctulata; and six in the Indo‐West Pacific, B. ampulla, B. arabica sp. nov. , B. orientalis, B. peasiana, B. quoyii and B. vernicosa. Full synonymies and taxonomic histories are provided for each species. In order to promote taxonomic stability, neotypes are designated for B. striata, B. solida, B. nebulosa (valid name B. gouldiana) and B. vernicosa, and lectotypes for B. occidentalis, B. mabillei, B. punctulata, B. ampulla and B. quoyii. The type locality of B. ampulla is restricted to Mauritius. Bullidae show a general morphological stasis, with anatomy being very similar between species. However, there are high levels of intraspecific variability in the shell, radula and male genital system. In some cases species could only be separated based on molecular data . After defining the characters and geographical range of each species it became clear that sympatric species (a maximum of three) show distinctive shells and reproductive structures, which makes identification straightforward. This study employs an integrative approach, combining information on shells, anatomy, DNA and geographical distribution, in order to resolve the systematics of a difficult taxonomic group. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 453–543.     

Systematic revision of the genus Everettia Godwin‐Austen, 1891 (Mollusca: Gastropoda: Dyakiidae) in Sabah,northern Borneo

The species of the snail genus Everettia in the Malaysian state of Sabah are superficially similar and difficult to distinguish by their shells. This paper presents new data on the taxonomy and distribution of Everettia in Sabah that have accumulated since the revision by Godwin‐Austen in 1891. By using morphological and molecular phylogenetic approaches, we reveal at least seventeen species of Everettia in Sabah, of which eleven are new to science, namely: Everettia layanglayang sp. nov. , Everettia lapidini sp. nov. , Everettia paulbasintali sp. nov. , Everettia occidentalis sp. nov. , Everettia jasilini sp. nov. , Everettia safriei sp. nov. , Everettia interior sp. nov. , Everettia jucundior sp. nov. , Everettia planispira sp. nov. , Everettia monticola sp. nov. , and Everettia dominiki sp. nov. , and one new subspecies, namely, Everettia corrugata williamsi ssp. nov. Phylogenetic analysis of mitochondrial COI and 16S, and nuclear ITS‐1 sequences demonstrates the monophyly of most of the morphologically well‐defined species. Our results show that certain aspects of classical morphology‐based taxonomy for Everettia species, especially with regard to the unique combination of shell surface sculptures, animal head colour, and mantle pigmentation, are solid. A dichotomous key to the Sabah species and subspecies of Everettia is provided. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 515–550.     

Diversity of isopods (Crustacea): new data from the Arctic and Atlantic Oceans

New data on the diversity pattern of isopods (Crustacea) from the northern most part of the North Atlantic and the Arctic Oceans is presented. The pattern of diversity with depth is similar at depths <1000m, but differs considerably below about 1000m. In the Arctic the diversity of isopods (expressed both as numbers of species per sled and expected number of species) increased with increased depth to a maximum at depths of about 320 to 1100m, but then declined towards deeper waters. There was a significant increase in numbers per sled and in the expected number of species with increased depth in the northernmost part of the North Atlantic Ocean. Additionally, changes occurred in the relative composition of the shallow and deep water fauna, with asellote isopods being relatively larger part of the isopod fauna in the Arctic than in the northern most part of the North Atlantic. This indicates major faunistic changes occurring at the Greenland-Iceland-Faeroe Ridge, possibly caused by rapid changes in the temperature. Furthermore, that the low diversity of the Arctic deep-sea is a regional phenomenon, and not a part of a large scale latitudinal pattern in the North Atlantic.     

A phylogeographic investigation of the kelp genus Laminaria (Laminariales,Phaeophyceae), with emphasis on the South Atlantic Ocean

The genus Laminaria has a wide distribution range compared with other kelp genera because it is found in both the North and the South Atlantic, on both sides of the North Pacific, as well as in the Mediterranean. Hypotheses behind this biogeographical pattern have been discussed by several authors but have not yet been fully evaluated with time‐calibrated phylogenies. Based on the analysis of four molecular markers (ITS2, rbcL, atp8 and trnWI), our goal was to reassess the Laminaria species diversity in South Africa, assess its relationship with the other species distributed in the South Atlantic and reconstruct the historical biogeography of the genus. Our results confirm the occurrence of a single species, L. pallida, in southern Africa, and its sister relationship with the North Atlantic L. ochroleuca. Both species belonged to a clade containing the other South Atlantic species: L. abyssalis from Brazil, and the Mediterranean L. rodriguezii. Our time‐calibrated phylogenies suggest that Laminaria originated in the northern Pacific around 25 mya, followed by at least two migration events through the Bering Strait after its opening (~5.32 mya). Today, the first is represented by L. solidungula in the Arctic, while the second gave rise to the rest of the Atlantic species. The colonization of the North Atlantic was followed by a gradual colonization southward along the west coast of Europe, into the Mediterranean (~2.07 mya) and two recent, but disconnected, migrations (~1.34 and 0.87 mya) across the equator, giving rise to L. abyssalis in Brazil and L. pallida in southern Africa, respectively.     

Evolutionary diversification of spiny rats (genus Trinomys, Rodentia: Echimyidae) in the Atlantic Forest of Brazil

Phylogenetic and phylogeographic relationships within and among species of the Atlantic Forest spiny rat Trinomys (family Echimyidae) were examined using cytochrome b sequence data. Levels of sequence divergence among species of Trinomys are as high as those found among taxa of echimyids that are recognized as different genera. Trinomys contains three distinct monophyletic clades that show a striking concordance with vegetational distribution. Haplotypes of clade 1 are distributed along the coastal margins of southeastern Brazil, following the moist tropical forest. Members of clade 2 are found in the semi-deciduous tropical forest. T. albispinus represents clade 3 and is found in a more xeric vegetation. Estimates of divergence times separating the three clades are very deep and range from 1.6 to 7.4 millions of years, predating the climatic fluctuations of the Pleistocene. Therefore, the proposed Late Pleistocene refugia in the Atlantic Forest cannot account for the divergence of the clades of Trinomys , but most likely shaped the modern distribution of species. The current taxonomy of this group does not reflect the diversity and phylogenetic relationships of the named species. However, morphological characters are congruent with the phylogeny uncovered by the molecular data. An extensive taxonomic rearrangement is suggested, reflecting phylogenetic relationships of monophyletic entities within the genus Trinomys , degree of sequence differences, and morphological diagnosability.     

Species in the genus Turritopsis (Cnidaria, Hydrozoa): a molecular evaluation

Mitochondrial ribosomal gene sequences were used to investigate the status of several populations of hydromedusae belonging to the genus Turritopsis (family Oceaniidae). Several nominal species have been described for this genus, but most of them had been synonymized and attributed to one cosmopolitan species, Turritopsis nutricula . A recent revision based on morphological and reproductive characters, however, has shown that many different populations can be distinguished and that several of the nominal Turritopsis species are likely valid biological species. Our investigation using molecular sequence data of 16S mitochondrial gene confirms these results. The Mediterranean Turritopsis must be attributed to Turritopsis dohrnii and the Turritopsis of New Zealand must be referred to Turritopsis rubra . The situation of the Japanese Turritopsis is more complex, though all sampled populations are clearly distinct from T. nutricula , a species likely confined to the Western Atlantic. The Japanese Turritopsis fall into three widely separated lineages. One of them, corresponding likely to Turritopsis pacifica , is closely related to T. rubra . A second clade, which potentially represents an as yet undescribed species, produces smaller medusae than T. pacifica and is morphologically distinguishable from it. Finally, a third group was distinguished by a single haplotype sequence that is identical with a Mediterranean sample of T. dohrnii . It is postulated that the last group of Japanese Turritopsis is likely a recent introduction, most probably by human activity. A survey of all known and potentially valid Turritopsis species is given in table format to facilitate identifications and future revisory work.     

Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores,including one new genus and three new species

The Hippasterinae is a subfamily within the Goniasteridae, consisting of five genera and 26 species, which occur in cold‐water settings ranging from subtidal to abyssal depths. All known genera were included in a cladistic analysis resulting in two most parsimonious trees, supporting the Hippasterinae as monophyletic. Our review supports Sthenaster emmae gen. et sp. nov. as a new genus and species from the tropical Atlantic and two new Evoplosoma species, Evoplosoma claguei sp. nov. and Evoplosoma voratus sp. nov. from seamounts in the North Pacific. Hippasteria caribaea is reassigned to the genus Gilbertaster, which previously contained a single Pacific species. Our analysis supports Evoplosoma as a derived deep water lineage relative to its continental‐shelf, shallow water sister taxa. The genus Hippasteria contains approximately 15 widely distributed, but similar‐looking species, which occur in the northern and southern hemispheres. Except for Gilbertaster, at least one species in each genus has been observed or is inferred to prey on deep‐sea corals, suggesting that this lineage is important to the conservation of deep‐sea coral habitats. The Hippasterinae shares several morphological similarities with Circeaster and Calliaster, suggesting that they may be related. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 266–301.     

A molecular phylogenetic appraisal of the systematics of the Aglaopheniidae (Cnidaria: Hydrozoa,Leptothecata) from the north‐east Atlantic and west Mediterranean

The hydrozoan family Aglaopheniidae (Cnidaria) is widespread worldwide and contains some of the most easily recognizable hydroids because of their large colony size and characteristic microscopic structure. The systematics of the group has, however, been controversial and dedicated molecular analyses are lacking. We therefore analysed existing and new 16S rRNA sequences of Aglaopheniidae, in a total of 98 16S sequences corresponding to 25 putative species (25 nominal and three undescribed) from seven genera. The monophyly of the subfamilies Gymnangiinae and Aglaopheniinae, and tribes Aglaopheniini and Cladocarpini were not verified with 16S sequence data. The genera Gymnangium and Aglaophenia can only be considered valid if both Gymnangium gracicaule and Aglaophenia latecarinata are removed from their respective genera. The phenotypically similar Cladocarpus and Streptocaulus are probably monophyletic and clearly distinct genetically. The genus Lytocarpia may be polyphyletic. The nominal species Aglaophenia pluma, Aglaophenia tubiformis, and Aglaophenia octodonta are probably conspecific, as are also the species Aglaophenia acacia and Aglaophenia elongata. The 16S data revealed the existence of two potentially unnamed species of Aglaophenia respectively from the Azores and Madeira. The phylogeographical structure of the taxa with the greatest representation of haplotypes from the north‐east Atlantic and Mediterranean, revealed the influence of Mediterranean waters in Madeira and the Azores, and gene flow between deep waters of the Mediterranean and Atlantic. The last glaciations in Europe may have caused genetic bottlenecks but also high intraspecific haplotype diversity. Finally, Macrorhynchia philippina was detected in samples from Madeira and possibly represents an invasive species. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 717–727.     

Reassessment of the trophic position of Bullidae (Gastropoda: Cephalaspidea) and the importance of diet in the evolution of cephalaspidean gastropods

A review of the available data on the diet of species of Bullidae is presented. Their trophic level is reassessed through gut content analysis of worldwide species from various habitats. Gut contents were identified by light and scanning electron microscopy. The previous claims of omnivory and carnivory were not supported. Worldwide species of Bullidae feed preferentially on algae and secondarily on diatoms, and are therefore herbivorous. The influence of dietary specialization on the evolution of cephalaspidean gastropods was assessed by superimposing the diet onto a molecular family-level phylogeny. Herbivory was found to be the plesiomorphic condition, with carnivory having arisen independently two or three times. This evidence suggests that dietary specialization has played a major role in the adaptive radiation of the group.     

Amphi-Atlantic distribution of the Mancocumatinae (Cumacea: Bodotriidae), with description of a new genus dwelling in marine lava caves of Tenerife (Canary Islands)

A new cumacean genus and species, Speleocuma guanche gen et sp. nov. , belonging to the subfamily Mancocumatinae Watling 1977 , is described from marine lava caves of Tenerife, Canary Islands. This genus differs from others in the subfamily in having two pairs of pleopods in males and exopods on first to third pereopods, but not on fourth pereopod, in both sexes. Correlation between phylogenetic relationships of the new genus and the geological history of the Canarian archipelago points to a probable origin of this endemism: the subfamily Mancocumatinae show a disjunct distribution on both sides of the Atlantic Ocean, a pattern which has been associated with fauna of Tethyan origin. It is suggested that present-day genera of the subfamily evolved from a common shallow water ancestor living in the late Mesozoic ( c . 120 Myr BP), when the opening of the Atlantic Ocean had begun. It survived in shallow waters off the west African coast before the emergence of the Canary Islands from the sea floor. Later on, between the late Cretaceous and the Mio-Pliocene, it was able to colonize crevicular habitats of the eastern Canary Islands through continental or land bridges. Finally, their numbers increased to occupy more recent biotops like flooded lava tube caves as well as westward islands such as Tenerife. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 134 , 453–461.     

Species range size distributions for some marine taxa in the Atlantic Ocean. Effect of latitude and depth

The range size distributions of 6643 species in ten different fish and invertebrate taxa dwelling in pelagic (latitudinal range sizes) and benthic (latitudinal and depth range sizes) habitats on both sides of the Atlantic Ocean (80°N−70°S) were studied. The objectives were to analyse: (1) the range size distribution patterns for the various taxa and whether they have right/left skewed or lognormal distributions; (2) the geographical species distributions, to ascertain whether the distribution ranges change with latitude (Rapoport's rule); and (3) the relationship between the depth ranges of benthic species and their maximum depth of occurrence and how depth range size distributions change with latitude. The pelagic taxa exhibited larger range sizes than did the benthic taxa, continental slope/rise species excepted. On the other hand, the boundaries between geographical provinces for both benthic taxa and pelagic taxa tended to occur in association with major oceanographic processes. The shape of the latitudinal range frequency distributions (LRFDs) of the pelagic organisms were distinctly left‐skewed, and the LRFDs for most taxa were significantly different from lognormal. There was no common pattern for the distributions of the benthic organisms, which were lognormal in Cephalopoda, Stomatopoda, and Crustacea Decapoda and tended to be left‐skewed and significantly different from lognormal in Pisces. The applicability of Rapoport's rule was not clearly inferable from the results, and the rule appears to be conditioned by the location of biogeographical boundaries and the endemism rate in the different biogeographical provinces. A clear increase in depth range size with maximum depth range was observable for benthic species, confirming previous studies. Species’ depth range distributions displayed a discernible latitudinal pattern, right‐skewed at high latitudes and left‐skewed at low latitudes. The location of biogeographical boundaries, and endemism rate by biogeographical province were considered to be the factors most useful in explaining species’ distribution patterns and their conformity or nonconformity to Rapoport's rule. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80 , 437–455.     

Out-of-Africa again: A phylogenetic hypothesis of the genus Charaxes (Lepidoptera: Nymphalidae) based on five gene regions

Despite the long popularity of Charaxes among collectors and researchers, their evolutionary history is largely unknown. The current and accepted species groupings and relationships within the genus are based exclusively on adult morphology and life histories. Here, we examine the monophyly and evolutionary affinities of the species-groups within the genus Charaxes and explore how they relate to members of their closest genera (Euxanthe, Polyura and Palla) using 4167 bp of sequence data from five (1 mitochondrial and 4 nuclear) gene regions. Within the proposed phylogenetic framework, we estimate ages of divergence within the genus and also reconstruct their historical biogeography. We included representatives of all known species-groups in Africa and Asia, all known species of Euxanthe and Palla and two exemplar species of Polyura. We found the genus Charaxes to be a paraphyletic group with regard to the genera Polyura and Euxanthe, contrary to the earlier assumption of monophyly. We found that 13 out of 16 morphologically defined species-groups with more than one species were strongly supported monophyletic clades. Charaxes nichetes is the sister group to all the other Charaxes. Polyura grouped with the Zoolina and Pleione species-groups as a well-supported clade, and Euxanthe grouped with the Lycurgus species-group. Our results indicated that the common ancestor of Charaxes diverged from the common ancestor of Palla in the mid Eocene (45 million years ago) in (Central) Africa and began diversifying to its extant members 15 million years later. Most of the major diversifications within the genus occurred between the late Oligocene and Miocene when the global climates were putatively undergoing drastic fluctuations. A considerable number of extant species diverged from sister species during the Pliocene. A dispersal–vicariance analysis suggests that many dispersal rather than vicariance events resulted in the distribution of the extant species. The genus Polyura and the Indo-Australian Charaxes are most likely the results of three independent colonizations of Asia by African Charaxes in the Miocene. We synonymize the genera Polyura (syn. nov.) and Euxanthe (syn. nov.) with Charaxes, with the currently circumscribed Charaxes subdivided into five subgenera to reflect its phylogeny.