The mitochondrial genome of Priapulus caudatus Lamarck (Priapulida: Priapulidae)

We sequenced and annotated the complete mitochondrial (mt) genome of the priapulid Priapulus caudatus in order to provide a source of phylogenetic characters including an assessment of gene order arrangement. The genome was 14,919 bp in its entirety with few, short non-coding regions. A number of protein-coding and tRNA genes overlapped, making the genome relatively compact. The gene order was: cox1, cox2, trnK, trnD, atp8, atp6, cox3, trnG, nad3, trnA, trnR, trnN, rrnS, trnV, rrnL, trnL(yaa), trnL(nag), nad1, -trnS(nga), -cob, -nad6, trnP, -trnT, nad4L, nad4, trnH, nad5, trnF, -trnE, -trnS(nct), trnI, -trnQ, trnM, nad2, trnW, -trnC, -trnY; where '-' indicates genes transcribed on the opposite strand. The gene order, although unique amongst Metazoa, shared the greatest number of gene boundaries and the longest contiguous fragments with the chelicerate Limulus polyphemus. The mt genomes of these taxa differed only by a single inversion of 18 contiguous genes bounded by rrnS and trnS(nct). Other arthropods and nematodes shared fewer gene boundaries but considerably more than the most similar non-ecdysozoan.     

Burrowing dynamics and energy cost of transport in the soft-bodied marine invertebrates Polyphysia crassa and Priapulus caudatus

The dynamics and mechanical forces generated during burrowing in Polyphysia crassa (Annelida: Polychaeta) and Priapulus caudatus (Priapulida) were investigated. Both animals live in soft marine muds and burrow by utilizing a direct peristaltic wave alternating with a high internal pressure event which thrusts the anterior part of the body into the substratum. Forces generated during the various phases of a typical burrowing cycle were measured in animals moving beneath the natural substratum at 5±3 °C using electronic transducers and recorder. During 'head' advance Polyphysia generated 0.027 N, and during 'tail' advance 0.020 N, with peak internal pressures averaging 0.95 kPa (= 0.095 N/cm²). Force by Priapulus during head advance and tail advance was 0.081 N and 0.121 N, respectively, with peak internal pressures averaging 2.47 kPa (= 0.247 N/cm²). Polyphysia moves more slowly (0.24 cm/min) than does Priapulus (0.76 cm/min) and expends more energy on mass moved per unit distance. These force measurements during a burrowing cycle were used in place of respirometry as a basis for computation of net cost of transport (NCT) for each animal. NCT for Polyphysia was 635 J kg^-1 m^-1 and for Priapulus was 314Jkg^-1m^-l. Cost of transport for all burrowing animals thus far investigated is high compared to swimming, running and flying. For soft-bodied invertebrates that live an entirely buried existence this high cost must be interpreted in the broader context of the adaptive value of infaunal life, especially protection against predation, and not as simply a means of moving about.     

Ultrastructure of erythrocytes and leucocytes of Priapulus caudatus (de Lamarck) (Priapulida)

The marine priapulid Priapulus caudatus has a voluminous body cavity filled with a blood-like fluid containing erythrocytes and leucocytes (amoebocytes). The hematocrit of animals weighing 0.5–14 gm was 2–10%. The erythrocytes contain a hemerythrin blood pigment. The structure of the coelomocytes was studied by light and electron microscopy. The erythrocytes are nucleated and contain marginal bands, vacuoles and occasionally crystals. The cytoplasm has few organelles. The leucocytes are amoeboid motile cells, the cytoplasm of which contains numerous organelles. The most conspicuous of these are oval particles, probably representing developmental stages of lysosomes. Most of these organelles contain tubules stretching from one pole to another. In the hind part of the animal, certain tissues, primarily the posterior warts contain large numbers of coelomocytes. The histological picture is complicated, showing some resemblance to the lymphoepithelial tissues of vertebrates.     

Amoebocytes in the Lining of the Body Cavity and Mesenteries of Priapulus caudatus (Priapulida)

Nuclei associated with membranes lining the body cavity in Priapulus caudatus are shown by electron microscopy to be the nuclei of amoebocytes. The membranes themselves are not cellular and do not constitute a peritoneum. Mesenteries, present in the body cavity, also are not cellular; amoebocytes occasionally are found within them. These findings weaken the idea that priapulids are coelomates, although unequivocal proof that priapulids are pseudocoelomates would require a fine structural study of the early developmental stages.     

Distribution patterns and biogeographic analysis of Austral Polychaeta (Annelida)

Aim We investigate the biogeography of Austral Polychaeta (Annelida) using members of the families Eunicidae, Lumbrineridae, Oenonidae, Onuphidae, Serpulidae and Spionidae and Parsimony Analysis of Endemicity (PAE). We determine whether observed polychaete distribution patterns correspond to traditional shallow-water marine areas of endemism, estimate patterns of endemism and relationships between areas of endemism, and infer the biological processes that have caused these patterns. Location The study is concerned with extant polychaete taxa occupying shallow-water areas derived from the breakup of the Gondwana landmass (i.e. Austral areas). Methods Similarity was assessed using a significance test with Jaccard's indices. Areas not significantly different at 0.99 were combined prior to the PAE. Widespread species and genera (155 taxa) were scored for presence/absence for each area of endemism. PAE was used to derive hypotheses of area relationships. Hierarchical patterns in the PAE trees were identified by testing for congruence with patterns derived from cladistic biogeographic studies of other Gondwanan taxa and with geological evidence. Results The polychaete faunas of four area-pairs were not significantly different and the areas amalgamated: South-west Africa and South Africa, New Zealand South Island and Chatham Islands, Macquarie Island and Antipodean Islands, and West Antarctica and South Georgia. Areas with the highest levels of species endemism were southern Australia (67.0%), South-east South America (53.2%) and South Africa (40.4%). About 60% of species and 7.5% of genera occupied a single area of endemism. The remainder were informative in the PAE. Under a no long-distance dispersal assumption a single minimal-length PAE tree resulted (l=367; ci=0.42); under dispersal allowed, three minimal-length trees resulted (l=278; ci=0.56). In relation to the sister grouping of the New Zealand areas and Australia we find congruence between our minimal-length trees and those derived from a biogeographic study of land plants, and with area relationships predicted by the Expanding Earth Model. Main conclusions The polychaete distribution patterns in this study differ slightly from the classical areas of endemism, most notably in being broader, thereby bringing into question the value of using single provincial system for marine biogeographic studies. The Greater New Zealand region is found to be ‘monophyletic’ with respect to polychaetes, that is comprising a genuine biogeographical entity, and most closely related to the polychaete fauna of southern Australia. This finding is consistent with studies of land plants and with the Expanding Earth model, but disagrees with conventional geology and biogeographic hypothesis involving a ‘polyphyletic’ New Zealand. Both vicariance and concerted range expansion (=biotic dispersion) appear to have played important roles in shaping present-day distribution patterns of Austral polychaetes. Shallow-water ridge systems between the Australian and Greater New Zealand continental landmasses during the Tertiary are thought to have facilitated biotic dispersion.     

Allozymes characterize sibling species of bipolar Priapulida (Priapulus,Priapulopsis)

Electrophoretic polymorphism of glucosephosphate isomerase (Gpi^*) and phosphoglucomutase (Pgm ^*) polymorphisms were assayed in the bipolardisjunct species pairsPriapulus caudatus/P. tuberculatospinosus andPriapulopsis bicaudatus/Priapulopsis australis (phylum Priapulida). Numbers of genotypes fromGpi ^* alleles and 14Pgm ^* alleles generally did not show deviations from Hardy-Weinberg expectations in composite populations sampled over geographic distances up to 500 km linearly measured. This genetic pattern suggests efficient population cohesion in a phylum where pelagic larvae have not been observed. Sibling species in generaPriapulus orPriapulopsis from northern and southern polar seas did not share identical allozyme alleles.     

Evolution of interstitial Polychaeta (Annelida)

An update of the systematics is given for the eight most important interstitial polychaete families: Diurodrilidae, Nerillidae, Protodrilidae, Protodriloididae, Saccocirridae, Parergodrilidae, Polygordidae and Psammodrilidae. Additional information and new observations are presented for the Diurodrilidae, Nerillidae and Psammodrilidae. Three new supplementary evolutionary hypotheses for these families are here suggested: (I) basal position of Diurodrilidae in Polychaeta, (2) evolution of Nerillidae in mud, and (3) evolution from meio- to macrofaunal forms of Psammodrilidae.     

Ultrastructure of a Coccidium (Apicomplexa: Sporozoea: Coccidia) in Priapulus caudatus (Priapulida)1

Stages in the life cycle of a coccidium are described from the intestine of Priapulus caudatus Lamarck, 1816. Meronts, merozoites, microgamonts, microgametes, and walled and unwalled macrogametes were seen in intestinal cells. Meronts were about 8 μm long and 3–7 μm wide and produced up to seven merozoites. Free merozoites were about 9 μm long and 4 μm wide and contained about 43 subpellicular microtubules that terminated in the outer polar ring. Microgamonts were up to 23 μm long and 7 μm wide and usually were delimited by a single membrane. Microgametes were about 5 μm long, exclusive of the two flagella, about 2 μm wide, and contained a nucleus that was not uniformly dense. Macrogametes, about 6 μm in diameter, had a nucleus largely without dense chromatin. The oocyst wall formed around intracellular macrogametes to a thickness of 0.2–0.5 μm as thin, osmiophilic elements that became arranged in reticular and tubular layers. Wall-forming bodies were not seen, but fine filaments may participate in wall formation, as these were found between the outer membrane of the pellicle and the nearest wall elements. Microgametes and walled macrogametes were delivered to the lumen of the host intestine during apocrine secretion or excretion by the intestinal cells. Fertilization may occur in the intestinal lumen. Unsporulated ovoid oocysts, 18–27 μm long and 10–14 μm wide, with a 3 μm micropyle and a wall 0.6–0.7 μm thick, were passed from the host.     

Microscopic anatomy and ultrastructure of the introvert of Priapulus caudatus and P. tuberculatospinosus (Priapulida)

Introverts of Priapulus caudatus and P. tuberculatospinosus bear 25 rows of scalids, as well as 8 spines and scattered papillae in the region the circumoral lip. These, as well as the first ring of pharyngeal teeth in P. tuberculatospinosus, are sensory organs. Although superficially they differ between species, they are all characterized by apical and/or subapical openings which are located on tiny cuticular tubules. All sensory organs contain cilia bearing bipolar receptor cells. The 8 sensory spines situated between the circumoral area and the beginning of the scalids are ultrastructurally similar to the scalids. The introvert and pharyngeal body walls, and associated muscles are described. © 1994 Wiley-Liss, Inc.     

Morphology of the nervous system of Polychaeta (Annelida)

The article summarizes our up to date knowledge about the morphology of the annelid, especially the polychaete, central and peripheral nervous system. Since the cephalic nervous system was in the focus of controversial discussions for decades, the structure of its neuropile, associated ganglia and nerves is reviewed in detail. The enormous variation of the ventral nerve cord and peripheral nerves is presented as well as a theory how this might have evolved. A ground pattern of the polychaete nervous system is suggested, based on developmental and regeneration studies.     

Phylogenetic affinities of the Flabelligeridae (Annelida, Polychaeta)

Cirratuliformia includes Acrocirridae, Cirratulidae, Ctenodrilidae, Flabelligeridae, Flotidae and Sternaspidae. The phylogenetic affinities have not been settled due to a limited availability of type or non-type material and the relationship between acrocirrids and flabelligerids have been problematical. In our study, the type material of all type species for all flabelligerid, and most acrocirrid genera have been studied and the morphological features have been used in a phylogenetic analysis. The results indicate that Acrocirridae, Ctenodrilidae, Fauveliopsidae, Flabelligeridae and Flotidae are monophyletic and that Sternaspidae falls within Cirratulidae; however, the latter conclusion might be reversed through increased taxon-sampling. The flabelligerid genera Brada, Flabelligera, Pherusa and Stylarioides each consists of several monophyletic groups and may be split. Conversely, Bradiella includes Diversibranchius, and the pelagic Buskiella includes Flota. The generic affinities of Poeobius remain uncertain, collecting better materials may resolve this issue.     

Coelomocytes of Nephtys coeca (Annelida,Polychaeta)

Coelomocytes of Nephtys coeca were studied by transmission electron microscopy. The majority of the coelomocytes were found to be structurally identical with the muscle cells of the body wall. Animals kept under unfavourable conditions tended to have an increased number of coelomocytes and a decreased thickness of the body wall. The muscular coelomocytes, probably released from the body wall, showed various degrees of decomposition, indicating a process of autophagy.     

Marine biodiversity of Costa Rica: class Polychaeta (Annelida)

Three hundred and thirty two species of polychaetes belonging to 50 families are reported for Costa Rica based on previously published reports. In a few families, available material was re-examined and appropriate changes in species identifications were made. Only three species, (Glycera oxycephala Ehlers, 1887; Hemipodia pustulata (Friedrich 1956); and Scolepis (Scolepis) squamata (Miller, 1806)) have been reported from the Caribbean coast of Costa Rica, and these three species have also been reported from the Pacific Coast. The families with more than 15 species are the Spionidae (26), Cirratulidae (24), Nereididae (21), Lumbrineridae (19) and Paraonidae (16). Despite extensive sampling on the Pacific coast over the last 25 years, the Costa Rican polychaete fauna, especially that of the Caribbean coast, remains poorly known.     

Clonal domains in postlarval Platynereis dumerilii (Annelida: Polychaeta)

Following an enzymatic procedure for softening the egg envelope, blastomeres in the embryo of the polychaete Platynereis dumerilii were injected with TRITC-dextran. Injection was successful in the following blastomeres: AB, CD, A, B, C, D, 1a-1d, 1A-1D, 4d, and 4d(1). The distribution of fluorescent label was recorded by confocal laser scanning microscopy of young, three-segmented worms after 3 or 4 days of development, in some cases also in 1-day-old trochophore larvae. Results were documented by single optical sections, by stacking a limited number or a complete set of optical sections, and by computer-generated surface views of both the labeled tissue domains and the body contours from complete image stacks of whole worms. With respect to their descent from the embryonic cell pattern, five major compartments can be distinguished which together compose the body of the young worm: 1) The epispheric, epidermal, and neural region of the head, composed of four domains arranged as quasi-radial sectors derived from micromeres 1a, 1b (left and right ventral), and 1c and 1d (right and left dorsal). 2) A posttrochal epidermal region of the head originating from micromeres 2a(1)-2c(1) and constituting the ventral and lateral posttrochal epidermis of the head. 3) A stomodeal-ectomesodermal region of the head, including the stomodeum (micromeres 2a(2) and 2c(2)), its mesodermal envelope, and head mesoderm (micromeres 3a-3d). 4) A solid cone composed of the four terminal macromeres 4A-4D, forming the core of the trunk as the endoderm anlage. 5) An epidermal and mesodermal coating of the trunk originating from the dorsal micromeres 2d and 4d. The region of the so-called (first, anterior) peristomial cirri at the posterior flanks of the head is also composed of 2d- and 4d-derived trunk tissue, thus corroborating the postulated descent of this region and its appendages from a cephalized anteriormost trunk segment and its parapodia. The cell-lineage domains of the first and third micromere tiers are arranged left or right of the sagittal plane, while two micromeres of the second quartet are in a lateral and, initially, two in a median position (2b ventral and 2d dorsal). The offspring of micromere 2d expand from a dorsal position toward the ventral midline and those of cell 4d from a posterior-dorsal site toward the anterior, initially forming two lateral bands. In the epispheric part of the head, part of the neurectodermal tissue derived from micromeres 1a and 1b interweaves in a medio-sagittal bar, and part of the first micromere offspring of all four quadrants (1a-1d) combine in forming a central brain neuropil. Each of the latter sends neurites through both of the circumesophageal connectives. Paired muscle tracts extend through the head toward the base of the antennae and are probably derived from micromeres 3a and 3b. A mesodermal envelope of the stomodeum is probably built by the 3c and 3d micromeres. The formation of symmetry and the nature of the body axes in the embryo and adult of Platynereis dumerilii are discussed. J. Morphol.     

Morphological variation in Polydora ciliata complex (Polychaeta: Annelida)

Intraspecific variation in Polydora ciliata (Johnston) was assessed on the basis of evidence from morphological characters, which were studied by scanning electron microscopy. Differences were observed between populations of P. ciliata with respect to the setae of the fifth modified segment and in the caruncle. Two other species, namely P. ligni Webster and P. limicola (Annenkova), were also studied in order to assess interspecific variation. The two forms of P. ciliata , that is boring and non-boring, were found to be different morphologically.     

Phylogenetic relationships within Nephtyidae (Polychaeta,Annelida)

Ravara, A., Wiklund, H., Cunha, M. R. & Pleijel, F. (2010). Phylogenetic relationships within Nephtyidae (Polychaeta, Annelida). —Zoologica Scripta, 39, 394–405. We present the first phylogeny of nephtyids, a common, soft‐bottom living polychaete family comprising five genera and over 100 species. Characters used to distinguish nephtyid genera are a matter of controversy and considerable confusion remains as to the generic delineations. The phylogeny is estimated with molecular data from the mitochondrial genes cytochrome oxidase I and 16S rDNA, the nuclear genes 18S rDNA and 28S rDNA and morphological data. The results reveal two well‐supported major clades, corresponding in part to the two main genera of the family, Aglaophamus and Nephtys. The species Nephtys pulchra and Nephtys australiensis are transferred to Aglaophamus, and new diagnoses for the genera are provided. Dentinephtys is synonymized with Nephtys, and Nephtys cornuta is sister to the remaining nephtyids and is referred to the new genus Bipalponephtys, together with Nephtys danida and Micronephthys neotena. Micronephthys is sister to Nephtys and Inermonephtys is of uncertain position.