Conchometrics, systematics and distribution of Melanopsis (Mollusca: Gastropoda) in the Levant

In this conchometric study, the systematics and distribution of the freshwater gastropod Melanopsis in the Levant are described. Of the ten species found, three are widespread, two have narrow distributions and five are known only from their type locality. Five smooth‐shelled species are recognized (buccinoidea, ammonis, dircaena, khabourensis and meiostoma). Within M. buccinoidea, Jordan Valley populations may belong to a separate subspecies. M. ammonis is clearly differentiated from buccinoidea of the nearby Jordan Valley, but less so from buccinoidea of more distant sites; differences between ammonis and meiostoma are significant but not diagnostic. Five species have ribbed shells. Within costata, four subspecies are recognized (in the northern Orontes, upper Jordan, Sea of Galilee, and in southernmost parts of the Levant). Throughout the Levant, M. buccinoidea frequently hybridizes with M. costata and fossil evidence suggests that these species have been hybridizing for the last 1.5 Myr. M. saulcyi differs from M. costata in its narrower shell and shorter, bumpier ribs and frequently hybridizes with M. buccinoidea throughout the Levant. Specimens from Homs differ from those of the Jordan Valley in their higher figurativity index, fewer ribs and lower rib density. M. germaini differs from M. costata in its more numerous ribs, M. pachya in its shorter ribs, and M. infracincta in its bumpy shell in which each rib has huge tubercles, with a pronounced ridge flanking the columella. Our conclusion that there are ten species in the Levant differs from previous studies that suggested only two subspecies of one species (or superspecies). This difference could stem from (1) our use of nonstandard as well as standard conchometrics, (2) a reappraisal of the importance of the shell vs. the radula in intrageneric systematics, and (3) differences of opinion on the subspecies concept. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144 , 229?260.     

Tropidofusus ypotethys: a new genus and new species of Columbariidae (Mollusca: Gastropoda: Turbinelloidea)

ABSTRACT

The genus Tripidofusus n. gen. is proposed to include Tropidofusus ypotethys n. sp. from bathyal depths off New Zealand, as well as Fulgurofusus aequilonius and Columbarium benthocallis from the abyssal zones of the Bering Sea and Scotia Sea, respectively. Tropidofusus can be distinguished from both Fulgurofusus and Coluzea on the basis of its lower spire, prominent peripheral keel, long, thin siphonal canal, as well as in having a much thinner shell that lacks axial sculpture. The type species, Tropidofusus ypotethys n. sp., differs from its congeners in having a broader peripheral keel, smaller aperture and longer siphonal canal, as well as by inhabiting bathyal rather than abyssal depths. The genus is reviewed and the relationships of Tropidofusus to other supraspecific taxa within Columbariidae are discussed, as are relationships among the families within Turbinelloidea.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:3CA568AD-1B6F-4D4E-9DF7-8377CABB8553     

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.     

Snails of the genus Pachnodus (Mollusca; Gastropoda; Enidae): their origins and evolution

Aim The phylogeny of the enid land-snail genus Pachnodus was determined in order to provide information on biogeographical patterns within the granitic Seychelles islands. Location The genus Pachnodus is endemic to the granitic islands of Seychelles (Mahé, Silhouette, Praslin, La Digue and Fregate). Methods Phylogeny was determined using a cladistic analysis of nineteen shell and soft-body anatomy characters. The outgroup used was the central-east African genus Cerastus. Results The proposed phylogeny indicates that the genus divided into two distinct subgenera early in its history. Subsequent speciation occurred in parallel in the two subgenera, resulting in several islands supporting at least two distinct species representing the two subgenera. Main conclusions The pattern of speciation is largely explicable by vicariance as a result of sea-level rises, followed by habitat specialisation and further speciation. The pattern is in contrast to previously published scenarios for other taxa in the islands and indicates significant differences between evolutionary and habitat factors in the biogeography of the Seychelles fauna     

Diversification and biogeographic history of the Western Palearctic freshwater flatworm genus Schmidtea (Tricladida: Dugesiidae), with a redescription of Schmidtea nova

The freshwater flatworm genus Schmidtea is endemic in the Western Palearctic region, where it is represented by only four species, thus contrasting with the high species diversity of the closely related genus Dugesia within Europe. Although containing an important model species in developmental and regeneration research, viz. Schmidtea mediterranea, no evolutionary studies on the genus Schmidtea have been undertaken. For the first time, we present a well‐resolved molecular phylogenetic tree of the four species of the genus, inferred on the basis of two molecular markers, and provide also the first detailed morphological account of Schmidtea nova. The phylogenetic tree generated corroborates an earlier speciation hypothesis based on karyological data and points to chromosomal rearrangements as the main drivers of speciation in this genus. The high genetic divergence between the four species, in combination with previous dating studies and their current geographic distribution, suggests that Schmidtea could have originated in Laurasia but lost most of its diversity during the Oligocene. Thus, its present distribution pattern may be the result of the expansion of three of its four relictual species over Europe, probably after the Pleistocene glaciations. Our detailed morphological study of S. nova revealed that it shows a number of remarkable features: interconnected testis follicles, parovaria, an ejaculatory duct exiting into the primary as well as the secondary seminal vesicle by means of a nipple, and the wall of the distal section of the ejaculatory duct being sclerotic or chitinized.     

Two Radix spp. (Gastropoda: Lymnaeidae) endemic to thermal springs around Lake Baikal represent ecotypes of the widespread Radix auricularia

In this study, we re‐examine two species of freshwater gastropods of the genus Radix Montfort, 1810 (family Lymnaeidae), endemic to the geothermal springs in the Lake Baikal region in the southern part of eastern Siberia — Lymnaea (Radix) hakusyensis Kruglov et Starobogatov, 1989, and Lymnaea (Radix) thermobaicalica Kruglov et Starobogatov, 1989. The alleged species status of these endemics has been re‐assessed by means of an integrative approach combining molecular genetic taxonomy techniques with the traditional methods based on shell and soft body morphology. Phylogenetic reconstructions were made using both mitochondrial (COI) and nuclear (ITS2) DNA markers. We used topotypic samples of both species and specimens sampled from other sites around Lake Baikal. The results demonstrate that the two endemic species are only synonyms of a widespread Holarctic species, Radix auricularia (Linnaeus, 1758), and represent its intraspecific morph (ecotype) adapted to living in thermal springs. A new synonymy is proposed: Thermoradix Kruglov et Starobogatov, 1989 = Radix Montfort, 1810 (syn. n.).     

Characterization of the bacterial microbiota of Biomphalaria glabrata (Say, 1818) (Mollusca: Gastropoda) from Brazil

Roughly 200 000 000 people in 74 countries infected with schistosomes all share the fact that they came in contact freshwater harbouring infected snails. The aim of the study is to characterize the microbiota of wild and laboratory‐reared snails of Biomphalaria glabrata from Pernambuco, Brazil. The microbiota of these molluscs was identified biochemically by the VITEK 2 automated microbiological system. Antimicrobial susceptibility testing was carried out by the disc diffusion method with ß‐lactam antibiotics, aminoglycosides, quinolones, folate pathway inhibitors, fenicols and tetracyclines. The results showed that all bacteria identified were gram‐negative, including 11 bacterial genera: Aeromonas, Citrobacter, Enterobacter, Cupriavidus, Rhizobium, Stenotrophomonas, Pseudomonas, Klebsiella, Acinetobacter, Vibrio and Sphingomonas. Regarding the antimicrobial susceptibility, all the isolates exhibited resistance to amoxicillin and sensitivity to meropenem (beta‐lactam antimicrobials). The microbiota of the wild snails consisted predominantly of Enterobacter cloacae, while the laboratory‐reared snails predominantly showed Citrobacter freundii and Aeromonas sobria.

Significance and Impact of the Study

Biomphalaria glabrata is a Brazilian freshwater Planorbidae of great medical relevance as an intermediate host of Schistosoma mansoni. About a month after being infected by one or more miracidia larvae of a compatible schistosome, B. glabrata sheds thousands of cercariae into the water where they seek human skin and, if successful, penetrate to establish infection, eventually taking up residence and maturing in blood vessels of the small intestine. Results obtained from this study aim at targeting novel biological control strategies for schistosomiasis such as paratransgenesis. This is the first study on the microbiota of B. glabrata from Brazil.     

A pre‐Miocene Irano‐Turanian cradle: Origin and diversification of the species‐rich monocot genus Gagea (Liliaceae)

The Irano‐Turanian (IT) floristic region is considered an important center of origin for many taxa. However, there is a lack of studies dealing with typical IT genera that also occur in neighboring areas. The species‐rich monocot genus Gagea Salisb. shows a center of diversity in IT region and a distribution in adjacent regions, therefore representing a good study object to investigate spatial and temporal relationships among IT region and its neighboring areas (East Asia, Euro‐Siberia, Himalaya, and Mediterranean). We aimed at (a) testing the origin of the genus and of its major lineages in the IT region, (b) reconstructing divergence times, and (c) reconstructing colonization events. To address these problems, sequences of the ribosomal DNA internal transcribed spacer (ITS) region of 418 individuals and chloroplast intergenic spacers sequences (psbA‐trnH, trnL‐trnF) of 497 individuals, representing 116 species from all sections of the genus and nearly its entire distribution area were analyzed. Divergence times were estimated under a random molecular clock based on nrITS phylogeny, which was the most complete data set regarding the representation of species and distribution areas. Ancestral distribution ranges were estimated for the nrITS data set as well as for a combined data set, revealing that Gagea most likely originated in southwestern Asia. This genus first diversified there starting in the Early Miocene. In the Middle Miocene, Gagea migrated to the Mediterranean and to East Asia, while migration into Euro‐Siberia took place in the Late Miocene. During the Pleistocene, the Arctic was colonized and Gagea serotina, the most widespread species, reached North America. The Mediterranean basin was colonized multiple times from southwestern Asia or Euro‐Siberia. Most of the currently existing species originated during the last 3 Ma.     

Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca,Gastropoda), with descriptions of three new species

The taxonomy of common northern nudibranch molluscs of the genus Dendronotus in the vast cold regions of Eurasia remains largely unknown. Abundant material collected in many localities from the Barents Sea, via the Arctic region, to the north‐west Pacific was analysed for the first time. An integrated approach combining morphological and ontogenetic data with molecular four‐gene (COI, 16S, H3, and 28S) analysis reveals seven species, including three previously undescribed. Dendronotus frondosus (Ascanius, 1774) and Dendronotus dalli Bergh, 1879 were commonly considered as amphiboreal species; however, according to this study they are restricted to the North Atlantic and the North Pacific, respectively. In the north‐west Pacific two new species were discovered, D endronotus kamchaticus sp. nov. and D endronotus kalikal sp. nov. , that are externally similar to D. frondosus, but that show significant distance according to molecular analysis and are considerably different in radular morphology. In the North Atlantic a new species D endronotus niveus sp. nov. , sibling to North Pacific D. dalli, is revealed. The separate status of North Atlantic Dendronotus lacteus (Thompson, 1840) is confirmed, including considerable range extension. The essential similarity of early ontogenetic stages of radular development common for species with disparate adult radular morphology (such as D. frondosus and D. dalli) is shown, and its importance for taxonomy is discussed. © 2015 The Linnean Society of London     

Molecular and morphological criteria for revision of the genus Microcoleus (Oscillatoriales,Cyanobacteria)

Ninety‐two strains of Microcoleus vaginatus (=nomenclatural‐type species of the genus Microcoleus Desmazières ex Gomont) and Phormidium autumnale Trevisan ex Gomont from a wide diversity of regions and biotopes were examined using a combination of morphological and molecular methods. Phylogenies based on the 16S rDNA and 16S‐23S ITS (partial) demonstrated that the 92 strains, together with a number of strains in GenBank, were members of a highly supported monophyletic clade of strains (Bayesian posterior probability = 1.0) distant from the species‐cluster containing the generitype of Phormidium. Similarity of the 16S rRNA gene exceeded 95.5% among all members of the Microcoleus clade, but was less than 95% between any Microcoleus strains and species outside of the clade (e.g., Phormidium sensu stricto). These findings, which are in agreement with earlier studies on these taxa, necessitate the revision of Microcoleus to include P. autumnale. Furthermore, the cluster of Phormidium species in the P. autumnale group (known as Group VII) must be moved into Microcoleus as well, and these nomenclatural transfers are included in this study. The main diacritical characters defining Microcoleus are related to the cytomorphology of trichomes, including: narrowed trichome ends, calyptra, cells shorter than wide up to more or less isodiametric, and facultative presence of sheaths. The majority of species are 4–10 μm in diameter. The possession of multiple trichomes in a common sheath is present facultatively in many but not all species.     

Morphology of sensory papillae on the feeding proboscis of cone snails (Mollusca,Gastropoda)

Cone snails, predatory marine gastropods, have developed a specialized prey capture method in which a long, distensible proboscis is used to identify prey and inject venom via a hydraulically propelled hollow radular tooth. Using brightfield, epifluorescence, confocal, and transmission and scanning electron microscopy, we describe the morphology of ciliated sensory structures concentrated on the tip of the proboscis. The number and morphology of these sensory papillae are linked to the type of preferred prey: cone snails feeding on worms and mollusks have short, cone or finger‐shaped papillae, whereas fish‐hunting cone snails have long tubular papillae in addition to short conical papillae. Sensory papillae are well positioned to provide information necessary to locate, identify, and dispatch prey. Proboscis tips and their sensory papillae regenerated within 10 d following experimental ablation, and snails with regenerated proboscis tips were able to locate and envenomate prey. The remarkable intrageneric variation found in the morphology of these sensory structures is probably linked to the specialized prey that cone snail species have evolved to hunt.     

A new genus of cave‐dwelling microcrustaceans from the Dinaric Region (south‐east Europe): adaptations of true stygobitic Cladocera (Crustacea: Branchiopoda)

We revise ‘true’ stygobitic cladocerans and lift three species from Alona Baird, 1843 (Cladocera: Chydoridae). Species of Brancelia gen. nov. are inhabitants of saturated karst, collected in pools of residual water in the amphibious zones of a few caves in the Dinaric Region, Europe. All species are blind (regression of eye and ocellus), have elongated sensorial equipment (aesthetascs) and a short rostrum, reduced antennal spines, and a globular body. In contrast to earlier hypotheses, there is no epigean chydorid taxon from the Palaearctic that can be linked to Brancelia gen. nov. The new genus may be an offshoot of six‐limbed Aloninae, but a littoral‐benthic ancestor is not apparent and most likely extinct. Evolution of Brancelia gen. nov. parallels that of other subterranean Cladocera like Phreatalona Van Damme, Brancelj & Dumont, 2009. We discuss the functional morphology of Brancelia gen. nov. and compare its adaptations to a subterranean life mode with those of Phreatalona. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 31–52.     

A nomenclator of extant and fossil taxa of the Valvatidae (Gastropoda,Ectobranchia)

A compilation of all supra- and (infra-) specific taxa of extant and fossil Valvatidae, a group of freshwater operculate snails, is provided, including taxa initially described in this family and subsequently classified in other families, as well as names containing errors or misspellings. The extensive reference list is directly linked to the available electronic source (digital view or pdf-download) of the respective papers.     

Miocene diversification of an open‐habitat predatorial passerine radiation,the shrikes (Aves: Passeriformes: Laniidae)

Diversification of avifaunas associated with savannah and steppes appears to correlate with open habitats becoming available, starting in the Miocene. Few comparative analyses exist for families for which all species are predominantly adapted to these habitats. One such group is Laniidae (Passeriformes), which are small‐ to medium‐sized predatory passerines known for their distinctive behaviour of impaling prey. We used multispecies coalescent‐based and concatenation methods to provide the first complete species‐level phylogeny for this group, as well as an estimate of the timing of diversification. Our analyses indicate that Laniidae as currently delimited is not monophyletic, as the genus Eurocephalus is not closely related to the remaining species. The two species currently assigned to the monotypic genera Urolestes and Corvinella are part of the same clade as the Lanius species, and we propose that they are included in the genus Lanius, making Laniidae monogeneric. The initial diversification of the clade is inferred to have occurred very rapidly, starting about 7.2–9.1 million years ago, timing depending on calibration method, but in either case coinciding with the expansion of C4 grasses. An African origin is inferred in the biogeographic analysis. In the redefined Laniidae, cooperative breeding is inferred to be restricted to a single clade, characterized by gregarious behaviour and rallying. Migratory behaviour evolved multiple times within the family.