Enzyme electrophoresis,genetic identity and description of a new genus and species of heteronemertean (Nemertea,Anopla) from northwestern Spain and North Wales

The anoplan order Heteronemertea, particularly the genera Cerebratulus, Lineus and Micrura, contains a very large number of nominate species, many of which are inadequately described. As a consequence, systematic difficulties are encountered with the identification of many taxa in this group, especially those originally established primarily on the basis of their external features. The present paper concerns heteronemerteans collected from two locations, the Foz Estuary (north-western Spain) and Llandudno (North Wales). The Spanish collection included specimens identified as Lineus longissimus (Gunnerus), whilst samples from Llandudno contained large numbers of Lineus viridis (Müller); samples of a third similar but apparently undescribed species were found at both locations. Starch gel electrophoresis showed that samples of the apparent third species were genetically almost identical from each of the two locations, but were clearly different from the two described Lineus species. Histological studies of the unknown specimens revealed anatomical characters, including the unique feature of a proboscis epithelium ciliated throughout its length, which exclude it from any known heteronemertean taxon; it is accordingly placed in a new genus and species, for which the name Riseriellus occultus is proposed.     

Nemerteans of the Great Barrier Reef 1. Anopla Palaeonemertea

Two new palaeonemerteans, Carinoma patriciae sp. nov. , and Hubrechtella queenslandica sp. nov. , from the Great Barrier Reef province of Australia, are described and illustrated. The systematic relationships of these nemerteans are discussed and emended diagnoses are given for the genera Carinoma (Carinomidae) and Hubrechtella (Hubrechtidae).     

Six new species of palaeonemerteans (Nemertea) from Hong Kong

One new genus and six new species of palaeonemerteans are described and illustrated from Hong Kong and the New Territories; these are Hubrechtella sinimarinus sp. nov. and Parahubrechtia jillae gen. et sp. nov. in the Hubrechtidae, and Callinera bergendali sp. nov., Carinina sinensis sp. nov., Tubulanus hylbomi sp. nov. and Tubulanus longivasculus sp. nov. in the Tubulanidae. These bring the known number of pa-laeonemertean species from Hong Kong to 11; a key to these species, based primarily upon features distinguishable in the living animals, is provided.     

Vestigial prototroch in a basal nemertean, Carinoma tremaphoros (Nemertea; Palaeonemertea)

Nemerteans have been alleged to belong to a protostome clade called the Trochozoa that includes mollusks, annelids, sipunculids, echiurids, and kamptozoans and is characterized by, among other things, the trochophore larva. The trochophore possesses a prototroch, a preoral belt of specialized ciliary cells, derived from the trochoblast cells. Nemertea is the only trochozoan phylum for which presence of the trochophore larva possessing a prototroch had never been shown. However, so little is known about nemertean larval development that comparing it with development of other trochozoans is difficult. Development in the nemertean clade Pilidiophora is via a highly specialized planktonic larva, the pilidium, and most of the larval body is lost during a drastic metamorphosis. Other nemerteans (hoplonemerteans and palaeonemerteans) lack a pilidium, and their development is direct, forming either an encapsulated or planktonic "planuliform" larva, producing a juvenile without a dramatic change in body plan. We show that early in the development of a member of a basal nemertean assemblage, the palaeonemertean Carinoma tremaphoros, large squamous cells cover the entire larval surface except for the apical and posterior regions. Although apical and posterior cells continue to divide, the large surface cells cleavage arrest and form a contorted preoral belt. Based on its position, cell lineage, and fate, we suggest that this belt corresponds to the prototroch of other trochozoans. Lack of differential ciliation obscures the presence of the prototroch in Carinoma, but differentiation of the trochoblasts is clearly manifested in their permanent cleavage arrest and ultimate degenerative fate. Our results allow a meaningful comparison between the development of nemerteans and other trochozoans. We review previous hypotheses of the evolution of nemertean development and suggest that a trochophore-like larva is plesiomorphic for nemerteans while a pilidium type of development with drastic metamorphosis is derived.     

Assessing the diversity and distribution of Cephalothrix species (Nemertea: Palaeonemertea) in European waters by comparing different species delimitation methods

Soft‐bodied marine taxa, like ribbon worms (Nemertea), often lack clear diagnostic morphological characters impeding traditional species delimitation. Therefore, recent studies concentrated on molecular genetic methods to solve taxonomic issues. Different delimitation methods were employed to explore species boundaries and the presence of cryptic species. However, the performance of the different delimitation methods needs to be tested. A particularly promising nemertean genus in this regard is the palaeonemertean genus Cephalothrix that is commonly found in European waters. In order to gain information on the number and distribution of European cephalotrichids and to test different tree‐based and non‐tree‐based delimitation methods, we analyzed a dataset comprising the barcoding region of the mitochondrial cytochrome c oxidase subunit I (COI) of 215 European Cephalothrix specimens, of which 78 were collected for this study. Our results show the presence of 12–13 European lineages of which several can be assigned to known European species. Analyzing a second dataset comprising 74 additional sequences from the Pacific and the Atlantic Oceans helped identify some of the unassigned European specimens. One resulting clade seems to represent a non‐native introduced Cephalothrix species, while another has never been recorded from Europe before. In our analysis, especially the tree‐based methods and the phylogenetic analysis proved to be a useful tool when delimiting species. It remains unclear whether the different identified clades result from cryptic speciation or from a high genetic variability of the COI gene.     

The partial mitochondrial genome of the Cephalothrix rufifrons (Nemertea, Palaeonemertea): characterization and implications for the phylogenetic position of Nemertea

A continuous 10.1kb fragment of the Cephalothrix rufifrons (Nemertea, Palaeonemertea) mitochondrial genome was sequenced and characterized to further assess organization of protostome mitochondrial genomes and evaluate the phylogenetic potential of gene arrangement and amino acid characters. The genome is A-T rich (72%), and this biased base composition is partly reflected in codon usage. Inferred tRNA secondary structures are typical of those reported for other metazoan mitochondrial DNAs. The arrangement of the 26 genes contained in the fragment exhibits marked similarity to those of many protostome taxa, most notably molluscs with highly conserved arrangements and a phoronid. Separate and simultaneous phylogenetic analyses of inferred amino acid sequences and gene adjacencies place the nemertean within the protostomes among coelomate lophotrochozoan taxa, but do not find a well-supported sister taxon link.     

A new genus and species of freshwater monostiliferous hoplonemertean (Nemertea, Enopla) from the People's Republic of China

A new genus and species of freshwater monostiliferous hoplonemertean, Limnemertes poyangensis gen. et sp. nov., from Poyang Lake, People's Republic of China, is described and illustrated. The taxon is compared and contrasted with previously described freshwater hoplonemerteans. This is the fourth species of freshwater nemertean to be described from China and the first recorded from Poyang Lake.     

A new genus and species of monostiliferous hoplonemertean (Nemertea: Enopla: Monostilifera) from Japan

A new genus and species of monostiliferous hoplonemertean, Diopsonemertes acanthocephala gen. et sp. nov., is described from Otsuchi Bay, Japan. Significant anatomical features of the new form include a body wall longitudinal musculature anteriorly divided into inner and outer layers by connective tissue, no pre-cerebral septum, the presence of a thin coat of diagonal muscle fibres between the body wall longitudinal and circular muscle layers in the foregut body region, cephalic retractor muscles derived only from the inner portion of the divided longitudinal muscles and a rhynchocoel more than half the body length.     

Protonephridia in the larvae of the paleonemertean species Carinoma mutabilis (Carinomidae,Nemertea) and Cephalothrix (Procephalothrix) filiformis (Cephalothricidae,Nemertea)

During spiralian development, the first pair of nephridia forms anterior to the mouth. Each organ consists of a few cells, which is characteristic for spiralian larvae. In nemerteans, one of the unambiguously spiralian taxa, so far protonephridia, has been reported only in advanced pilidium larvae, where they likely persist as juvenile and adult nephridia. These organs have not been recorded in larvae of the basally branching nemertean taxa. In search for these organs, we examined the ultrastructure of pelagic planuliform larvae of the palaeonemerteans Carinoma mutabilis and Cephalothrix (Procephalothrix) filiformis. In both species, a pair of protonephridia is located at the level of the stomodaeum. Each protonephridium of C. mutabilis consists of two terminal cells, two duct cells and one nephropore cell, while that of C. filiformis consists of three terminal cells, three duct cells and one nephropore cell. In C. mutabilis and in C. filiformis, all terminal cells contribute to forming a compound filtration structure. In both species, the protonephridia seem to develop subepidermally, since in C. filiformis, the nephropore cells pierce the larval epidermis and in C. mutabilis, the nephropores are initially covered by the binucleated multiciliated trophoblast cells. On the fifth day, these cells degenerate, so that the protonephridium becomes functional. The occurrence of protonephridia in the larvae of both paleonemertean species is in accordance with the hypothesis that a common ancestor of Nemertea and Trochozoa had a larval stage with a pair of protonephridia. This does not contradict previous hypotheses on placing the Nemertea as an ingroup of the Trochozoa or Spiralia (= Lophotrochozoa). Whether these protonephridia are restricted to the larval phase or whether they are transformed into the adult protonephridia, like those of the pilidium larva, remains to be answered.     

Estimation of the density ofBuccinum undatum (Gastropoda) off Douglas,Isle of Man

The density of the common whelk (Buccinum undatum L.) off Douglas, Isle of Man, was determined by four methods: (1) pot sampling, (2) diving, (3) mark-recapture experiment, and (4) underwater television. Although the values obtained by these methods were comparable, the last two methods yielded overestimations ofBuccinum density. The results from diving survey and from pot sampling showed a good agreement, indicating that pot sampling can be used to determine the density of the common whelk, provided a good estimate of the pot attraction area is available. The range of whelk density between February 1989 and August 1990 resulting from pot sampling was between 0.08 and 0.38 individuals m⁻². The temporal fluctuations of the whelk densities are discussed in detail.     

Polymorphism hides cryptic species in Oerstedia dorsalis (Nemertea, Hoplonemertea)

The marine hoplonemertean Oerstedia dorsalis is considered to be a highly polymorphic species with extensive geographic distribution. We show, based on mitochondrial cytochrome c oxidase subunit I, 16S and nuclear internal transcribed spacer sequences, that there are genetic subsets withing this species. Seventy-one specimens of various colours from different geographic localities (in Europe) were sequenced and analysed using statistical parsimony and Bayesian analysis. Both analyses supported nine major clades. We conclude that O. dorsalis hides different species with geographic resolution. These species, however, appear to be polymorphic as well, and we find no diagnostic features in pigmentation or external characters to separate species within this complex.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 556–567.     

Phylogenetic analysis of a group of palaeonemerteans (Nemertea) including two new species from Queensland and the Great Barrier Reef, Australia

Sundberg, P., Gibson, R. & Olsson, U. (2003). Phylogenetic analysis of a group of palaeonemerteans (Nemertea) including two new species from Queensland and the Great Barrier Reef, Australia. — Zoologica Scripta, 32, 279–296.
Based on 18S rDNA nucleotide sequences and morphological characters, we reconstruct the phylogeny for a group of palaeonemerteans estimated to be monophyletic. Two new palaeonemertean species from Queensland and the Great Barrier Reef, Australia are included in the phylogenetic analysis. The results confirm that one of the species, Cephalothrix queenslandica sp. n., is part of the Cephalothrix–Cephalotrichella–Procephalothrix group. These genera are redefined phylogenetically under the name Cephalothrix based on the cladistic analysis. The other species, Balionemertes australiensis gen. et sp. n., is placed in a new genus which forms a sister taxon to Cephalothrix . The morphology of both new species is described in detail.     

A new species ofFumana (Cistaceae) from Rif, Morocco

A new species ofFumana (Dunal) Spach subgenusFumana, F. fontqueri, is described from the region of Rif, Morocco. Notes on its morphology, ecology, distribution and taxonomic relationships are presented. It differs fromF. procumbens (Dunal) Gren. etGodr. andF. baetica Güemes, by the indument of the stem, the disposition of pedicels, and the size of the flowers.     

A new species of Ornithocephalus (Orchidaceae) from Oaxaca,Mexico

Ornithocephalus torresii Salazar & Soto Arenas, a new species from the Pacific slope of Oaxaca. Mexico, is described and illustrated. It is distinguished by the nearly closed, globose flowers and the cymbiform lip with a small, irregularly dentate-lacerate lobule at each side of the base. A key to all the species ofOrnithocephalus hiterto recorded from Mexico is provided.