Cosmopolitan versus cryptic meiofaunal polychaete species: an approach to a molecular taxonomy

Polychaete taxonomy is characterised by a high number of apparently cosmopolitan species. Detection of subtle but diagnostic ultrastructural differences and – in recent years – investigations at the molecular level have revealed that many of these "species" are actually complexes of morphologically identical or almost identical cryptic species. To disregard their existence would lead to an underestimation of global meiofauna diversity and undermine the value of many scientific studies. Therefore, we strongly recommend that they be given formal taxonomic recognition, beyond their published presentation as "operational taxonomic units", "types" or by alphabetic or numerical designators. Since there are neither generally accepted practical procedures nor any established consensus regarding the application of genetic data in taxonomy, we here provide examples of, and suggestions for, the treatment of meiofaunal species that are distinguished exclusively by molecular data, e.g. by genetic distance values, cluster analyses, diagnostic (= autapomorphic) DNA fragments from DNA fingerprinting procedures (RAPD) and/or DNA sequence differences (e.g. of ITS 2). Although no holotype material may be available because the molecular procedures require the preparation of entire specimens, practical taxonomic problems can be overcome and the recommendations of the Zoological Code of Nomenclature satisfied, by adopting the following procedures: (1) deposition of band-patterns of an individual obtained with the primers used to find diagnostic markers; (2) deposition of DNA in ethanol of one syntype individual; (3) deposition of fixed specimens (syntypes) from the locus typicus. Electronic Publication     

Immunohistochemical analysis of nervous system regeneration in chimeric individuals of Dorvillea bermudensis (Polychaeta, Dorvilleidae)

In regeneration experiments, 0.5% of the two- or five-segmented fragments of the polychaete Dorvillea bermudensis were found unexpectedly transplanted: two fragments of each that were lying close together during the initial period, fused and regenerated a chimeric individual. Of the three theoretical possibilities (i.e. fusion of (i). two posterior ends; (ii). one anterior and one posterior end; (iii). or two anterior ends) only the last two were realized. The similarly oriented fragments regenerated a normal animal while anterior-anterior fused ones produced two heads or a double head. Whether the ventral cords of the fragments are located vis-à-vis or adjacent, influences the course of regeneration as well. Immunohistochemical methods (anti-acetylated alpha-tubulin) in conjunction with confocal laser scanning microscopy were used to investigate the wiring pattern of the nervous systems of the grafts. In all cases, at least two supraesophageal ganglia were formed and palps, antennae and nuchal organs were innervated by the correct nerves but, in special cases, were innervated vice versa from the other brain. From these results it can be concluded that fusion of a regenerating connective with another connective results in formation of a new brain, irrespective of whether it belongs to the same nerve cord or not.     

Molecular phylogeny of lugworms (Annelida, Arenicolidae) inferred from three genes

Arenicolids comprise a group of four genera in which about 30 nominal species are described. Whereas the biology of many arenicolids is well known, the phylogenetic relationships of these worms are inadequately studied. A close relationship of Arenicolidae and Maldanidae is generally accepted. The phylogenetic relationships of arenicolid taxa were reconstructed based on sequence data of the mitochondrial 16S rRNA gene, the nuclear 18S rRNA gene, and a small fraction of the nuclear 28S rRNA gene. Members of all described arenicolid genera are included in the data set. Phylogenetic analyses were conducted using Maximum Likelihood, Bayesian inference, and Maximum Parsimony. The monophyly of the Maldanidae, as well as of the Arenicolidae is supported by all conducted analyses. Two well supported major clades are highest ranked sister taxa in the Arenicolidae: one containing all Abarenicola species and one containing Arenicola, Arenicolides, and Branchiomaldane. Evidence is given for a closer relationship between the two investigated Branchiomaldane species and Arenicolides ecaudata in the combined analysis. In the light of the molecular data the best explanation for structural and morphological observations is that Branchiomaldane evolved by progenesis.     

Host-specific performance and host use in the kleptoparasitic marine snail<Emphasis Type="Italic"> Trichotropis cancellata</Emphasis>

Generalist parasites may disproportionately use certain hosts because of different benefits associated with each host species. I measured the growth rate of the marine snail Trichotropis cancellata, a facultative kleptoparasite that can suspension feed and steal food, on different hosts to determine the relative nutritional benefits of each host. The variation in tentacle (feeding structure) area among the hosts studied had the potential to provide parasitic snails with different amounts of nutrition for growth. In field experiments, suspension-feeding snails isolated from potential hosts grew at a similar rate to snails on brachiopods and significantly more slowly than snails on the following polychaete worms: Serpula columbiana (Serpulidae), Pseudopotamilla ocellata (Sabellidae), Schizobranchia insignis (Sabellidae), and Eudistylia vancouveri (Sabellidae). However, choice among worm hosts affected snail growth rates only in the fall, when phytoplankton levels are low. At this time, snails parasitizing the sabellids Schizobranchia and Eudistylia grew more quickly than snails on Serpula. In the spring and summer, with high levels of phytoplankton, Trichotropis grew at similar rates on all worm species tested. Trichotropis spent approximately the same time stealing food from each worm host species, >50% of the time the worms had their tentacles extended (the difference among hosts was not significant). This finding demonstrates that the similarity of snail growth rates on different worm species is not due to the snails compensating for poor hosts (worms that provide food at a slower rate) by spending more time stealing food. Snails in choice experiments preferred live Serpula to empty Serpula tubes, indicating that at least some of the cue(s) snails use to identify hosts are derived from living host tissues. In choice racks containing live Serpula and live Schizobranchia, snails did not choose one host worm significantly more often than the other. Because Trichotropis grows faster on sabellids than serpulids in the fall, I predicted that snails in nature would infect sabellids more often than other species. However, snails were usually distributed randomly among host species. In the few cases where the snails showed a significant preference among host species, proportionally more snails were found on serpulids than on sabellids or sabellarids. This study is the first to quantify under natural conditions the growth benefits of a kleptoparasite across the range of possible hosts, and implies that factors other than growth rate influence host choice specificity in the marine kleptoparasite T. cancellata.     

Polychaetes associated with the sciaphilic alga community in the northern Aegean Sea: spatial and temporal variability

Polychaete biodiversity has received little attention despite its importance in biomonitoring. This study describes polychaete diversity, and its spatial and temporal variability in infralittoral, hard substrate assemblages. Seven stations were chosen in the central area of the northern Aegean Sea. At each station, one to three depth levels were set (15, 30 and 40 m). Five replicates were collected by scuba diving with a quadrat sampler (400 cm²) from each station and depth level during summer for the spatial analysis, and seasonally for the study of temporal changes. Common biocoenotic methods were employed (estimation of numerical abundance, mean dominance, frequency, Margalefs richness, Shannon-Weaver index and Pielous evenness). A total of 5,494 individuals, belonging to 79 species, were counted and classified. Diversity indices were always high. Clustering and multidimensional scaling techniques indicated a high heterogeneity of the stations, although these were all characterized by the sciaphilic alga community. A clear seasonal pattern was not detectable. Summer and autumn samples discriminate, while winter and spring form an even group. The abundance/biomass comparison indicated a dominance of k-strategy patterns, characteristic of stable communities.Communicated by H.-D. Franke     

Comparative anatomy of excretory organs in vestimentiferan tube worms (Pogonophora, Obturata)

In the past, the excretory systems of only few vestimentiferan species have been examined in detail. This study presents comparative data on eight species on the basis of histological serial sections. Ridgeia piscesae was studied by transmission electron microscopy. All species examined possess a central excretory organ consisting of numerous small branching and intertwined excretory tubules. These are connected to voluminous glandular excretory ducts that lead to the exterior by single or paired excretory pores located at the anterior end of the vestimentum. A comparative analysis shows differences among the species with regard to several features, such as the number of excretory pores, presence/absence of excretory grooves and papillae, position of the excretory organ relative to the brain, and the shape of the excretory ducts. Neither podocytes nor coelomoducts could be detected; therefore, there is no indication of the presence of metanephridia. The vestimentiferan excretory system shows some similarities with the design in Athecanephria (Pogonophora, Perviata) and a general resemblance to the design in sabellid polychaetes, even though in the latter metanephridia are clearly present.     

Genetic differences between two allopatric sibling species of the genus Polydora (Polychaeta: Spionidae) from the West Pacific

Polydora vulgaris Mohammad, 1972, a commensal borer of the oysters Pinctada margaritifera and Hyotissa hyotis from the South China Sea, was investigated by means of starch gel electrophoresis. Polydora vulgaris and the allopatric sibling Polydora glycymerica Radashevsky, 1993, a commensal borer of the bivalve Glycymeris yessoensis from the Sea of Japan, were compared with respect to their allozymic variation and number of isozyme loci. Interspecific differences in the number of gene loci coding for three enzymes: alanopine dehydrogenase, glucose-6-phosphate isomerase and -iditol dehydrogenase were revealed suggesting that we are dealing with two valid species. Two different modes of origin duplicate loci in polydorids are dicussed—polyploidization and regional gene duplication. The use of gene number as a character for discriminating between morphologically indistinguishable allopatric polydorid taxa is outlined.     

Acrosome formation and the centriolar complex in the spermatozoa of Sabella penicillum (Polychaeta)

Summary The acrosomal vesicle of Sabella penicillum spermatids consists of an electrondense core and a more transparent surrounding zone. During subsequent differentiation the vesicle membrane forms several invaginations in the juxtanuclear area. These invaginations later establish contact with the core. In the mature spermatozoon the spaces between the invaginations appear as electron-dense tubules; this is probably due to a shift of material from core to periphery. The ultrastructure of the centriolar complex is described in detail.Work supported by the Norwegian Research Council for Science and Humanities (NAVF; Grant Nr. D 61.44) and The Austrian Fonds zur Förderung der wissenschaftlichen Forschung Projekt 2183 and N 39.     

Hox gene expression in larval development of the polychaetes Nereis virens and Platynereis dumerilii (Annelida, Lophotrochozoa)

The bilaterian animals are divided into three great branches: the Deuterostomia, Ecdysozoa, and Lophotrochozoa. The evolution of developmental mechanisms is less studied in the Lophotrochozoa than in the other two clades. We have studied the expression of Hox genes during larval development of two lophotrochozoans, the polychaete annelids Nereis virens and Platynereis dumerilii. As reported previously, the Hox cluster of N. virens consists of at least 11 genes (de Rosa R, Grenier JK, Andreeva T, Cook CE, Adoutte A, Akam M, Carroll SB, Balavoine G, Nature, 399:772–776, 1999; Andreeva TF, Cook C, Korchagina NM, Akam M, Dondua AK, Ontogenez 32:225–233, 2001); we have also cloned nine Hox genes of P. dumerilii. Hox genes are mainly expressed in the descendants of the 2d blastomere, which form the integument of segments, ventral neural ganglia, pre-pygidial growth zone, and the pygidial lobe. Patterns of expression are similar for orthologous genes of both nereids. In Nereis, Hox2, and Hox3 are activated before the blastopore closure, while Hox1 and Hox4 are activated just after this. Hox5 and Post2 are first active during the metatrochophore stage, and Hox7, Lox4, and Lox2 at the late nectochaete stage only. During larval stages, Hox genes are expressed in staggered domains in the developing segments and pygidial lobe. The pattern of expression of Hox cluster genes suggests their involvement in the vectorial regionalization of the larval body along the antero-posterior axis. Hox gene expression in nereids conforms to the canonical patterns postulated for the two other evolutionary branches of the Bilateria, the Ecdysozoa and the Deuterostomia, thus supporting the evolutionary conservatism of the function of Hox genes in development. Milana Kulakova, Nadezhda Bakalenko and Elena Novikova contributed equally to this work.