A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia)

The phylogenetic relationships of the cryptobranch dorids are studied based on morphological characters of species belonging to all previously described genera. The phylogenetic hypothesis supports the cryptobranch dorids as a monophyletic group. There are two major clades within the Cryptobranchia: the radula‐less dorids (Porostomata), and the radula‐bearing dorids ( Labiostomata new taxon ). Labiostomata consists of those taxa sharing a more recent common ancestor with Actinocyclus than with Mandelia, and includes several monophyletic groups: Actinocyclidae, Chromodorididae, Dorididae and Discodorididae. The traditional group Phanerobranchia is probably paraphyletic. The new classification proposed for the Cryptobranchia addresses concepts of phylogenetic nomenclature, but is in accordance with the rules of the International Code of Zoological Nomenclature. The following genera of cryptobranch dorids are regarded as valid: Doris Linnaeus, 1758, Asteronotus Ehrenberg, 1831, Atagema J. E. Gray, 1850, Jorunna Bergh, 1876, Discodoris Bergh, 1877, Platydoris Bergh, 1877, Thordisa Bergh, 1877, Diaulula Bergh, 1878, Aldisa Bergh, 1878, Rostanga Bergh, 1879, Aphelodoris Bergh, 1879, Halgerda Bergh, 1880, Peltodoris Bergh, 1880, Hoplodoris Bergh, 1880, Paradoris Bergh, 1884, Baptodoris Bergh, 1884, Geitodoris Bergh, 1891, Gargamella Bergh, 1894, Alloiodoris Bergh, 1904, Sclerodoris Eliot, 1904, Otinodoris White, 1948, Taringa Er. Marcus, 1955 , Sebadoris Er. Marcus & Ev. Marcus, 1960, Conualevia Collier & Farmer, 1964, Thorybopus Bouchet, 1977, Goslineria Valdés, 2001, Pharodoris Valdés, 2001, Nophodoris Valdés & Gosliner, 2001. Several genera previously considered as valid are here regarded as synonyms of other names: Doridigitata d’Orbigny, 1839, Doriopsis Pease, 1860, Staurodoris Bergh, 1878, Fracassa Bergh, 1878, Archidoris Bergh, 1878, Anoplodoris Fischer, 1883, Etidoris Ihering, 1886, Phialodoris Bergh, 1889, Montereina MacFarland, 1905, Ctenodoris Eliot, 1907, Carryodoris Vayssière, 1919, Austrodoris Odhner, 1926, Guyonia Risbec, 1928, Erythrodoris Pruvot‐Fol, 1933, Neodoris Baba, 1938, Siraius Er. Marcus, 1955, Tayuva Ev. Marcus & Er. Marcus, 1967, Nuvuca Ev. Marcus & Er. Marcus, 1967, Doriorbis Kay & Young, 1969, Pupsikus Er. Marcus & Ev. Marcus, 1970, Percunas Ev. Marcus, 1970, Verrillia Ortea & Ballesteros, 1981 . The genera Artachaea Bergh, 1882, Carminodoris Bergh, 1889 and Homoiodoris Bergh, 1882 have been poorly described and no type material is known to exist. They are regarded as incertae sedis until more material becomes available. The genus names Xenodoris Odhner in Franc, 1968 and Cryptodoris Ostergaard, 1950 are unavailable within the meaning of the Code. Hexabranchus Ehrenberg, 1831 is not a cryptobranch dorid, as suggested by other authors, because of the lack of a retractile gill. Other nomenclatural and taxonomic problems are discussed, and several type species, neotypes and lectotypes are selected. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society, 2002, 136 , 535?636.     

Sperm ultrastructure in the nudibranch genus Halgerda with reference to other Discodorididae and to Chromodorididae (Mollusca: Opisthobranchia)

Comparative sperm ultrastructure within the molluscan nudibranch genus Halgerda (Discodorididae) was examined for the first time using transmission electron microscopy (TEM), based on 17 of the 35 known species. In addition, observations on two other discodorids are made to facilitate outgroup comparison with Halgerda, including one species of Discodoris (D. boholiensis) and Asteronotus cespitosus (currently accepted as the closest sister taxon to Halgerda). Comparison was also made with some genera of the Chromodorididae in view of sperm similarities. Spermatozoa of all species examined were of the complex, helical, elongate ( approximately 300-400 micro m) type characteristic of most heterobranch gastropods. These cells exhibit the following discrete regions (in anteroposterior sequence) : an acrosomal complex (composed of a rounded, membrane-bound vesicle and a column-like pedestal); a solid, helical nucleus; an elongate, helical midpiece (composed of an axoneme and associated nine coarse fibers, an enveloping mitochondrial derivative of matrix, and paracrystalline materials and glycogen helix); an annular complex; and a short glycogen piece. Of these regions, the midpiece is by far the longest, occupying over 90% of the total sperm length. Comparison with other members of the radula-bearing cryptobranch dorids reveals several sperm similarities to other genera in the clade, particularly those of other Discodorididae and also with the Chromodorididae. Comparison with previously studied genera reveals noteworthy sperm differences within the Discodorididae. The most notable differences are the internal structure of the acrosomal pedestal (long and homogeneous in Halgerda, Discodoris; short and homogeneous in Asteronotus; long and finely striated in Rostanga; oblong with angular electron-lucent striations in Jorunna) and the internal structure of the glycogen piece. The pronounced helical keels of most Halgerda and Discodoris nuclei contrast with the weakly helical nucleus of Asteronotus. Sperm features alone do not provide a means of defining the genus Halgerda or the family Discodorididae nor do they support the monophyletic status of the caryophyllidia-bearing dorids. Important sperm characters such as the acrosome, nucleus, and midpiece can often still be determined from specimens that have been initially fixed in formalin, then stored in ethanol for extended periods of time (i.e., museum material). Of all sperm features, the mitochondrial derivative of the midpiece is the most resistant to long-term fixation : the survival of acrosomal, nuclear, and axonemal components is variable, presumably a factor of prefixation autolysis, varied primary fixation times and temperatures, formalin quality, and duration of alcohol storage.     

Phylogeny of the radula-less dorids (Mollusca, Nudibranchia), with the description of a new genus and a new family

This paper studies the phylogenetic relationships of the nudibranch dorids that lack a radula. Numerous specimens belonging to 29 different species, representing all genera of this group and other related taxa, were examined anatomically. Details of the foregut were examined with SEM. From this anatomical study a total of 62 characters were considered. These characters have been polarized using the genus Berthella as the outgroup. Eight species of dorids with a radula and the genus Armina have been also included in the analysis for comparative purposes. The phylogeny obtained supports the hypothesis that the radula has been lost once in the Doridina and that the radula-less dorids are a monophyletic group. Cryptobranchia is monophyletic when the radula-less dorids are included. The new genus Mandelia , introduced on the basis of a new species from South Africa, is the sister group of the rest of the radula-less dorids, and is described as a new family, Mandeliidae. The radula-less dorids (currently placed in a separate superfamily) actually constitute an internal branch of other cryptobranch dorids. Phyllidia has no synapomorphies that distinguish if from Fryeria , which has one autoapomorphy, the ventral position of the anus. Therefore, both genera are regarded as synonyms. The evolution of several characters among the radula-less dorids is discussed, and the present phylogeny is compared with prior studies.     

The taxonomy and biology of further aeolidacean and arminacean nudibranch molluscs with symbiotic zooxanthellae

The occurrence of symbiotic zooxanthellae in further aeolid and arminacean nudibranch molluscs is described for the first time. The aeolid Aeolidiopsis ransoni Pruvot-Fol is redescribed, and a new species of Aeolidiopsis , also feeding on the colonial zoantharian Palythoa , is described. Both have symbiotic zooxanthellae. The taxonomy of the family Aeolidiidae is discussed and the genus Berghia Trinchese, 1877 is considered a synonym of Spurilla Bergh, 1864. Spurilla major (Eliot, 1903) and a new species of Spurilla are reported to have zooxanthellae while another new species of Spurilla is without zooxanthellae. The glaucid aeolid Pteraeolidia ianthina (Angas) is shown to have symbiotic zooxanthellae, as is the arminacean Doridomorpha gardineri Eliot, which is reported to feed on the alcyonarian blue coral, Heliopora. In all cases, the morphological adaptations developed for this symbiosis are described. Further notes on the Porites-fetding arminacean Pinufius rebus Marcus & Marcus and the aeolid Phestilla lugubris (Bergh) are included and a facultative symbiosis with zooxanthellae is suggested for the latter. The tion of symbiosis with zooxanthellae within nudibranchs is discussed and it is suggested that the relationship has evolved independently on several occasions.     

Aeolid opisthobranch molluscs (Glaucidae) from the Indian Ocean and the south-west Pacific

Eight aeolid opisthobranch molluscs of the subfamilies Facelininae, Favorininae, and Herviellinae are reported from Tanzanian waters, and two species from Northwestern India. New records from Queensland, Australia greatly extend the range of two species reported from Tanzania. Phidiana militaris (Alder & Hancock) and P. indica (Bergh) are shown to be distinct and a species from New Zealand, originally identified as P. militaris , is shown to be new. P. bourailli (Risbec), previously reported only from New Caledonia, is described from Tanzania, as is a new species of Phidiana. Favorinus japonicus Baba is reported from Tanzania, the first published record outside Japan, a new species of Godiva is described from Tanzania and Queensland, and three new species of Sakuraeolis are described, one from India and two from Tanzania. A new species of Herviella is described from Tanzania.     

A new species of Monoplacophora, redescription of the genera Veleropilina and Rokopella, and new information on three species of the class

Vema levinae Warén, sp. n. is described from a submarine volcano off western Mexico. The type species of Rokopella and Veleropilina are redescribed, the usage of the names is discussed and Rokopella and Veleropilina are recognized as valid genera in Neopilinidae. Rokopella euglypta (Dautzenberg & Fischer, 1897) is redescribed from shells and a single live specimen taken from seamounts south of the Azores, from a depth of 1200–1600 m. Tectura reticulata Seguenza, 1876 (Gastropoda, 'Acmaeidae', southern Italy, Upper Pliocene/Lower Pleistocene) is considered to have been based on the monoplacophoran species previously known as Neopilina zografi from the Mediterranean, and is classified in the genus Veleropilina. Veleropilina zografi (Dautzenberg & Fischer, 1896) is redescribed based on shells from several seamounts south of the Azores. It is considered distinct from Rokopella euglypta and classified in the genus Veleropilina. An undescribed species of Veleropilina from a seamount off southern Baja California is reported, figured and discussed but not formally described.     

Systematic revision of Jorunna Bergh, 1876 (Nudibranchia: Discodorididae) with a morphological phylogenetic analysis

The genus Jorunna is characterized by a dorsum covered withcaryophyllidia, a prostate with two sections, a penis usuallyunarmed but occasionally armed with hooks, a copulatory spine,the presence of an accessory gland and a labial cuticle smoothor armed with jaw elements. The examination of 216 non-typespecimens, 30 types, and a review of the literature show thatthere are 16 valid species of the genus Jorunna: J. tomentosa(Cuvier, 1804); J. funebris (Kelaart, 1859); J. pantherinaAngas,1864; J. rubescens (Bergh, 1876); J. labialis (Eliot, 1908);J. parva (Baba, 1938); J. spazzola (Marcus, 1955); J. hartleyi(Burn, 1958); J. alisonaeMarcus, 1976; J. lemchei (Marcus, 1976);J. evansi (Eliot, 1906); J. pardusBehrens & Henderson, 1981;J.ramicolaMiller, 1996 and J. onubensis Cervera, García-Gómez& García, 1986. In addition, two new species fromthe Eastern Pacific are described: J. osae n. sp. and J. tempisquensisn. sp. We propose two new combinations: Jorunna parva and J.evansi. New records for the genus Jorunna are provided fromItaly, Algeria, Seychelles, Madagascar, Thailand, Marshall Islands,New Caledonia, Île de la Réunion, Sudan, PapuaNew Guinea, Indonesia, Panama, Costa Rica, Bahamas, and SouthernMexico. We present the first preliminary phylogenetic analysisof this cryptobranch dorid genus, based on morphological anatomicaldata, and discuss the biogeography and evolution of severalcharacters in this group. The phylogeny supports the hypothesisthat the genus Jorunna is a monophyletic group and shows thatKentrodoris is nested within it. (Received 31 December 2004; accepted 10 January 2008)     

Molecular phylogeny of the scincid lizards of New Caledonia and adjacent areas: evidence for a single origin of the endemic skinks of Tasmantis

We use approximately 1900bp of mitochondrial (ND2) and nuclear (c-mos and Rag-1) DNA sequence data to recover phylogenetic relationships among 58 species and 26 genera of Eugongylus group scincid lizards from New Caledonia, Lord Howe Island, New Zealand, Australia and New Guinea. Taxon sampling for New Caledonian forms was nearly complete. We find that the endemic skink genera occurring on New Caledonia, New Zealand and Lord Howe Island, which make up the Gondwanan continental block Tasmantis, form a monophyletic group. Within this group New Zealand and New Zealand+Lord Howe Island form monophyletic clades. These clades are nested within the radiation of skinks in New Caledonia. All of the New Caledonian genera are monophyletic, except Lioscincus. The Australian and New Guinean species form a largely unresolved polytomy with the Tasmantis clade. New Caledonian representatives of the more widespread genera Emoia and Cryptoblepharus are more closely related to the non-Tasmantis taxa than to the endemic New Caledonian genera. Using ND2 sequences and the calibration estimated for the agamid Laudakia, we estimate that the diversification of the Tasmantis lineage began at least 12.7 million years ago. However, using combined ND2 and c-mos data and the calibration estimated for pygopod lizards suggests the lineage is 35.4-40.74 million years old. Our results support the hypothesis that skinks colonized Tasmantis by over-water dispersal initially to New Caledonia, then to Lord Howe Island, and finally to New Zealand.     

Frugivore gape size and the evolution of fruit size and shape in southern hemisphere floras

Abstract The present study uses differences among frugivore faunas of the southern hemisphere landmasses to test whether frugivore characteristics have influenced the evolution of fruit traits. Strong floristic similarities exist among southern landmasses; for example, 75% of New Zealand vascular plant genera also have species in Australia. However, plants in Australia and South America have evolved in the presence of a range of mammalian frugivores, whereas those in New Zealand, New Caledonia and the Pacific Islands have not. In addition, the avian frugivores in New Zealand and New Caledonia are generally smaller than those of Australia. If frugivore characteristics have influenced the evolution of fruit traits, predictable differences should exist between southern hemisphere fruits, particularly fruit size and shape. Fruit dimensions were measured for 77 New Zealand species and 31 Australian species in trans‐Tasman genera. New Zealand fruits became significantly more ellipsoid in shape with increasing size. This is consistent with frugivore gape size imposing a selective pressure on fruit ingestability. This result is not a product of phylogenetic correlates, as fruit length and width scaled isometrically for Australian species in genera shared with New Zealand. Within‐genus contrasts between New Zealand and Australian species in 20 trans‐Tasman genera showed that New Zealand species have significantly smaller fruits than their Australian counterparts. Within‐genus contrasts between New Zealand and South American species in nine genera gave the same result; New Zealand species had significantly smaller fruits than their South American counterparts. No difference was found in fruit size or shape between New Zealand and New Caledonia congeneric species from 12 genera. These results are consistent with the broad characteristics of the frugivore assemblage influencing the evolution of fruit size and shape in related species. The smaller‐sized New Zealand frugivore assemblage has apparently influenced the evolution of fruit size of colonizing taxa sometimes within a relatively short evolutionary timeframe.     

ON THE STATUS OF THE GENERIC NAMES OF THE NUDIBRANCH GENERA CATRIONA, CRATENA, HERVIA, RIZZOLIA AND TRINCHESIA

The genus Catriona Winckworth (1941) is synonymous with TrinchesiaPruvot-Fol (1951) but not with Trinchesia von Jhering (1879). It has priority over Trinchesia Pruvot-Fol. The genus Cratena Bergh (1864) is valid and synonymous withRizzolia Trinchese (1877) but not with Hervia Bergh (1871). The genus Hervia Bergh (1871) is synonymous with Facelina Alderand Hancock (1855), but not all species of Hervia can be includedin the latter genus, and these should, therefore, be distributedamong other genera, some of which are listed. (Received 8 October 1954;     

Monophyletic origin of Caryophyllia (Scleractinia,Caryophylliidae), with descriptions of six new species

The genus Caryophyllia Lamarck, 1816 is the most diverse genus within the azooxanthellate Scleractinia comprising 66 Recent species and a purported 195 nominal fossil species. Examination of part of the deep-sea scleractinian collection made by the Paris Museum off New Caledonia and part of the material collected by CSIRO off Australian waters revealed the occurrence of 23 species of Caryophyllia, of which six are new to science. All new records, including the new species, are described, and synonyms, distribution, type locality, type material and illustration are provided for each species. An identification key to all Recent species of Caryophyllia is presented. In addition, the validity of the genus Caryophyllia was investigated by phylogenetic analyses of a dataset consisting of partial mitochondrial 16S rRNA sequences from 12 species assigned to this genus together with seven species representing some of the most morphologically similar caryophylliid genera, and 14 non-caryophyllid species representing 14 scleractinian families. Irrespective of the method of analysis employed, all of the Caryophyllia species formed a well-supported clade together with Dasmosmilia lymani and Crispatotrochus rugosus. Although based on a subset of the Recent Caryophyllia species, these results are consistent with Caryophyllia being a valid genus, but call for a reexamination of Dasmosmilia and Crispatotrochus.     

Two new species of nudibranchiate molluscs from the west coast of North America, with a revision of the family Cuthonidae

Two species of nudibranch molluscs are described from the northeastern Pacific Ocean on the west coast of North America. These are: a new aeolid species of Cuthona Alder & Hancock, 1855 (family Cuthonidae) from the Oregonian marine zoogeographical province and a new dorid species of Acanthodoris Gray, 1850 (family Onchidorididae) from the Panamic province in the Gulf of California. The internal anatomy and external features of each species are compared with related species. Differences in structure of the radula, reproductive system and other morphological aspects are described. A revision of the family Cuthonidae is included. Evidence is presented to include the genera Trinchesia Ihering, 1879, and Precuthona Odhner, 1929, within the genus Cuthona. Cuthona alpha Baba & Hamatani, 1963, is synonymized with Catriona columbiana (O'Donoghue, 1922). We consider the New World Cuthonidae to be composed of the following five genera: Catriona Winckworth, 1941; Cuthona Alder & Hancock, 1855; Embletonia Alder & Hancock, 1851; Tenellia Costa, 1877; and Tergipes Cuvier, 1805.     

Integrative taxonomy of New Caledonian beetles: species delimitation and definition of the Uloma isoceroides species group (Coleoptera,Tenebrionidae, Ulomini), with the description of four new species

New Caledonia is an important biodiversity hotspot with much undocumented biodiversity, especially in many insect groups. Here we used an integrative approach to explore species diversity in the tenebrionid genus Uloma (Coleoptera, Tenebrionidae, Ulomini), which encompasses about 150 species, of which 22 are known from New Caledonia. To do so, we focused on a morphologically homogeneous group by comparing museum specimens with material collected during several recent field trips. We also conducted molecular phylogenetic analyses based on a concatenated matrix of four mitochondrial and three nuclear genes for 46 specimens. The morphological study allowed us to discover and describe four new species that belong to the group of interest, the Uloma isoceroides group. Molecular analyses confirmed the species boundaries of several of the previously described species and established the validity of the four new species. The phylogenetic analyses also provided additional information on the evolutionary history of the group, highlighting that a species that was thought to be unrelated to the group was in fact a member of the same evolutionary lineage. Molecular species delimitation confirmed the status of the sampled species of the group and also suggested some hidden (cryptic) biodiversity for at least two species of the group. Altogether this integrative taxonomic approach has allowed us to better define the boundaries of the Uloma isoceroides species group, which comprises at least 10 species: Uloma isoceroides (Fauvel, 1904), Uloma opacipennis (Fauvel, 1904), Uloma caledonica Kaszab, 1982, Uloma paniei Kaszab, 1982, Uloma monteithi Kaszab, 1986, Uloma robusta Kaszab, 1986, Uloma clamensae sp. n., Uloma condaminei sp. n., Uloma jourdani sp. n., and Uloma kergoati sp. n. We advocate more studies on other New Caledonian groups, as we expect that much undocumented biodiversity can be unveiled through the use of similar approaches.     

On the Paleozoic bryozoans of the genus <Emphasis Type="Italic">Ascopora</Emphasis> Trautschold

Based on the study of the neotype and the astogeny of the type specimens of the species Ascopora nodosa (Fischer von Waldheim, 1837), the diagnosis and species composition of the genus Ascopora Trautschold, 1876 are emended. The genus Ascopora and three species out of the five included in this genus—A. nodosa (Fischer von Waldheim, 1837); A. sokolovae Schulga-Nesterenko, 1955; and A. latiaxis Schulga-Nesterenko, 1955—are described. A new genus, Tetrasella gen. nov., with the type species T. blanda sp. nov. is described. This genus comprises five species, transferred from the composition of the genus Ascopora. An emended diagnosis of the genus Ascoporella Krutchinina, 1986, is provided. This genus comprises nine species, of which two, A. lecta sp. nov. and A. mera sp. nov., are described as new species.     

Stigmatomyces from New Zealand and New Caledonia: new records, new species and two new host families

Seven new records and three new species of Stigmatomyces (S. australis, S. baeopteri and S. clasiopellae), a genus of Laboulbeniales associated only with Diptera, are described from New Zealand and New Caledonia. Two new host families for Laboulbeniales are recorded, and a dichotomous key to Stigmatomyces species in New Zealand and New Caledonia is presented.     

A taxonomic revision of Paradoris sea slugs (Mollusca, Gastropoda, Nudibranchia, Doridina)

Paradoris , traditionally a generic taxon of discodorid sea slugs, is revised for the first time. One hundred and thirty specimens were examined, including all types and most of the nontype specimens available. New records for Paradoris are provided: South Africa, Tanzania, Seychelles, Western Australia, Thailand, Philippines, shallow waters of New Caledonia, southern Japan, and Hawaii. The individual variation of all taxonomic characters is thoroughly evaluated. Prior to the present study, Paradoris included 15 nominal species: 12 valid names and three synonyms. Three additional names, Discodoris erythraeensis , D. lora , and D. cavernae , are re-allocated to Paradoris , based on phylogenetic analysis. A phylogenetic diagnosis is provided for Paradoris : in particular, two new synapomorphies are described. Eight species names are regarded as valid: P. araneosa , P. dubia , P. erythraeensis , P. indecora , P. liturata , P. lopezi , P. mulciber , and P. tsurugensis . However, most of these species are poorly known, i.e. from very few specimens, and their taxonomic status might change when more individuals are available. P. lora is regarded as a nomen dubium . Six new synonymies are proposed, and explained by the fact that: (1) species names were created for one or a few specimens, without considering individual variation; (2) authors have not worked within a phylogenetic framework and have created new species names without considering all the existing species names already available within Paradoris . Three new morphospecies are described, but not formally named because their taxonomic status is still uncertain for several reasons (e.g. lack of knowledge of individual variation for some critical features).  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 147 , 125–238.