Pattern of body-wall muscle differentiation during embryonic development of Enchytraeus coronatus (Annelida: Oligochaeta; Enchytraeidae)

The plesiomorphic arrangement of body-wall musculature within the annelids is still under discussion. While polychaete groups show a great variety of patterns in their somatic muscles, the musculature of soil-living oligochaetes was thought to represent the characteristic pattern in annelids. Oligochaete body-wall muscles consist of an outer continuous layer of circular and an inner continuous layer of longitudinal muscles, forming a closed tube. Since designs of adult body musculature are influenced by evolutionary changes, additional patterns found during embryogenesis can give further information about possible plesiomorphic features. In oligochaetes, detailed cell-lineage analyses document the origin of the mesoderm and consequently the muscles, but later processes of muscle formation remain unclear. In the present work, body-wall muscle differentiation was monitored during embryogenesis of thesoil-living oligochaete Enchytraeus coronatus (Annelida) by phalloidin staining. Primary circular muscles form in a discrete anterior-to-posterior segmental pattern, whereas emerging longitudinal muscles are restricted to one ventral and one dorsal pair of primary strands, which continuously elongate towards posterior. These primary muscles establish an initial muscle-template. Secondary circular and longitudinal muscles subsequently differentiate in the previous spaces later in development. The prominent ventral primary longitudinal muscle strands on both sides eventually meet at the ventral midline due to neurulation, which moves the ventral nerve cord into a coelomic position, closing the muscle layers into a complete tube. This early embryonic pattern in E. coronatus resembles the adult body-wall muscle arrangements in several polychaete groups as well as muscle differentiation during embryonic development of the polychaete Capitella sp. I.     

Life history of the oligochaete Enchytraeus coronatus (Annelida, Enchytraeidae) in agar culture

Abstract. Life-history attributes and the reproductive biology of Enchytraeus coronatus (Oligochaeta, Enchytraeidae) were studied in agar cultures, as a basis for using this species in toxicity tests and other experimental studies. The use of agar allows daily examination of the behavior, reproduction, and mortality of the worms. At 20°C, E. coronatus had a generation time of 21–22 days and an embryonic period of 7–10 days. Hatch rate was high (>80%) and worms became sexually mature 8–10 days after hatching. Cocoon production and number of eggs per cocoon were positively correlated. Mortality in the first stages of the life cycle (embryonic period and first week after hatching) was about 38%. Three different periods of cocoon production were distinguished: a period characterized by a regular increase in the number of cocoons produced (weeks 4–9), a period of production at a constant high rate of ∼5 cocoons per week (weeks 10–20), and a period of decline in production rate (weeks 21–32), down to a mean value of 1.6 cocoons per adult in the last week. This study showed that synchronized agar cultures of cohorts can provide adults for long-term experimental tests, with the age for optimum reproduction being 8–20 weeks.     

Cell lineage and fate map of the primary somatoblast of the polychaete annelid Capitella teleta

Like most polychaete annelids, Capitella teleta (formerly Capitella sp. I) exhibits a highly stereotypic program of early development known as spiral cleavage. Animals with spiral cleavage have diverse body plans, and homologous embryonic cells can be readily identified among distantly related animals. Spiralian embryos are particularly amenable to studies of fate-mapping, and larval fates of identified cells are conserved among diverse taxa. One cell of particular importance in spiralian development is 2d, or the primary somatoblast, which generates ectoderm of the body posterior to the prototroch. We are interested in the evolution of the primary somatoblast, and thus far, the 2d sublineage has only been analyzed in a few species. In Capitella teleta, 2d generates ectoderm of the segmented trunk and post-segmental pygidium. In this study, development of the 2d lineage was characterized in detail through intracellular injections of DiI, and time-lapse as well as confocal microscopy to analyze cleavage patterns and the fates of larval cells. Analysis of cleavage patterns reveals that the first bilateral division in the 2d sublineage occurs with the division of 2d112, the same 2d daughter cell that first divides bilaterally in the polychaete Platynereis dumerilii. Larval fates of blastomeres 2d1, 2d2, 2d11, 2d12, 2d112, 2d1121, and 2d1122 were determined. All cells show stereotypic descendant clones that are consistent with segregation within sublineages. In the first few divisions of the 2d sublineage, larval-specific structures (neurotroch and telotroch) and pygidial ectoderm are segregated from segmental ectoderm and ventral nerve cord. The daughters of the first bilateral division, 2d1121 and 2d1122, generate the right and left halves of the segmental ectoderm and ventral nerve cord respectively, although the clones are consistently asymmetric across the dorsal midline. The pattern of cleavage divisions and the fates of the 2d daughters in Capitella teleta are compared to those in other spiralians with special attention to annelids.     

Expression of <Emphasis Type="Italic">hunchback</Emphasis> protein in a subset of ectodermal teloblasts of the oligochaete annelid <Emphasis Type="Italic">Tubifex</Emphasis>

The early expression patterns of hunchback protein (T-hb protein) were examined in the oligochaete Tubifex, using an antibody raised to the LZF2 protein in leech. This antibody recognizes a 60-kDa polypeptide in the Tubifex embryo. Before teloblastogenesis, T-hb protein is expressed in every cleavage-stage blastomere. At the completion of teloblastogenesis, the only cells expressing T-hb are a fraction of the micromere-derived epithelial cells. During gastrulation, nuclear T-hb is seen in spreading micromere-derived epithelial cells and also in a subset of ectodermal teloblasts. Comparisons of these results with those from other annelids suggest that hb expression in the early cleavage blastomeres and the micromere-derived epithelium are features highly conserved among annelids. In contrast, hb expression in teloblasts appears to be an innovation evolved in the oligochaetes.     

Establishment of the dorsoventral axis in nemertean embryos: Evolutionary considerations of spiralian development

The Nemertea represent one of a number of invertebrate phyla that display a highly conserved pattern of cell division known as spiral cleavage. The fates of the early blastomeres are known for representatives of some spiralian phyla (i.e., molluscs and annelids) and in these species there appears to be a high degree of conservation in the ultimate fates of particular embryonic cells. The first two cleavage planes bear an invariant relationship to the symmetry properties of the future larval and adult body plan. To investigate whether these properties of spiralian embryo-genesis are shared (conserved) amongst members of other spiralian phyla, individual blastomeres in two- and four-cell embryos of the nemertean, Nemertopsis bivittata, were microinjected with bi-otinylated dextran lineage tracers. N. bivittata is a direct-developing hoplonemertean that forms a nonfeeding larva. When individual blastomeres are injected at the two-cell stage, two sets of complementary labeling patterns (a total of four different patterns) were observed in the ectoderm of the larvae. When cells were injected at the four-cell stage, four different patterns were observed that represented subsets of the four patterns observed in the previous experiment. Unlike the case in the annelids and molluscs, in which the first cleavage plane bears a strict 45° angular relationship to the future dorsoventral axis, the first cleavage plane in N. bivittata can bear one of two different relationships relative to the larval/adult dorsoventral axis. In half the cases examined, the first cleavage plane corresponded roughly to the plane of bilateral symmetry, and in the rest, it lay along a frontal plane. A similar result was observed for the embryos of the indirect-developing heteronemertean, Cerebratutus lacteus. These results indicate that the fates of the four cell quadrants in nemerteans are not directly homologous to those in other spira-lians, such as the annelids and molluscs. For instance, no single cell quadrant appears to contribute a greater share to the formation of ectoderm, as is the case in the formation of the post-trochal region by the D-cell quadrant in annelids and mol-luscs. Rather, two adjacent cell quadrants contribute nearly equally to the formation of dorsal or ventral ectoderm in the larvae. Possible explanations for the determination of dorsoventrality in nemerte-ans, as well as implications of these findings regarding the evolution of spiralian development, are discussed. © 1994 Wiley-Liss, Inc.     

Trochophora larvae: cell-lineages, ciliary bands, and body regions. 1. Annelida and Mollusca

The trochophora concept and the literature on cleavage patterns and differentiation of ectodermal structures in annelids ("polychaetes") and molluscs are reviewed. The early development shows some variation within both phyla, and the cephalopods have a highly modified development. Nevertheless, there are conspicuous similarities between the early development of the two phyla, related to the highly conserved spiral cleavage pattern. Apical and cerebral ganglia have almost identical origin in the two phyla, and the cell-lineage of the prototroch is identical, except for minor variations between species. The cell-lineage of the metatrochs is almost unknown, but the telotroch of annelids and the "telotroch" of the gastropod Patella originate from the 2d-cell, as does the gastrotroch in the few species which have been studied. The segmented annelid body, i.e. the region behind the peristome, develops through addition of new ectoderm from a ring of 2d-cells just in front of the telotroch. This whole region is thus derived from 2d-cells. Conversely, the mollusc body is covered by descendants of cells from both the C and D quadrants and a growth zone is not apparent. This supports the notion that the molluscs are not segmented like the annelids, and that the repeated structures seen in polyplacophorans and monoplacophorans do not represent a segmentation homologous to that of the annelids.     

Cleavage patterns, cell-lineages and cell specification are clues to phyletic lineages in Spiralia

Embryos of molluscs, annelids, nemerteans and platyhelminthes show remarkable intra- and interphyletic resemblances and differences in mesentoblast, dorso-ventral axis and trochoblast specification. These variations have been used to investigate their evolutionary relationship. In molluscs and annelids a heterochronic shift parallels evolutionary relations based on adult characters. Nemerteans and platyhelminthes lack trochal cells and differ in the specification of the mesodermal precursor cell. Nemerteans also differ fundamentally with respect to axis specification related to the first cleavage. Therefore, close phylogenetic relations exist between molluscs and annelids, whereas nemerteans and platyhelminthes are only remotely related with each other and with molluscs and annelids.     

The Articulata hypothesis – or what is a segment?

The long held view that annelids and arthropods are closely related (Articulata) has been challenged recently by phylogenetic analyses using molecular data. The outcome of these studies is a clade of moulting animals (Ecdysozoa) comprising arthropods and some taxa of the nemathelminth worms. Monophyly of the Ecdysozoa has not yet been shown convincingly on morphological evidence, but is strongly supported by molecular data. The implication of the Ecdysozoa hypothesis is that the type of segmentation found in annelids and arthropods must be either convergent or an ancestral feature of protostomes or even bilaterians. The present review discusses aspects of segmentation in annelids and arthropods at the genetic, cellular, morphogenetic and morphological levels. Based on numerous similarities not shared with other bilaterian taxa it is suggested that segmentation of annelids and arthropods is homologous and apomorphic for a monophyletic Articulata. However, the challenge provided by the molecular analyses should stimulate research programmes gaining more data such as on additional genes, cleavage patterns, molecular developmental biology, and the comparison of nervous systems at the level of single neurons.     

Conserved mechanism of dorsoventral axis determination in equal-cleaving spiralians

Many members of the spiralian phyla (i.e., annelids, echiurans, vestimentiferans, molluscs, sipunculids, nemerteans, polyclad turbellarians, gnathostomulids, mesozoans) exhibit early, equal cleavage divisions. In the case of the equal-cleaving molluscs, animal-vegetal inductive interactions between the derivatives of the first quartet micromeres and the vegetal macromeres specify which macromere becomes the 3D cell during the interval between fifth and sixth cleavage. The 3D macromere serves as a dorsal organizer and gives rise to the 4d mesentoblast. Even though it has been argued that this situation represents the ancestral condition among the Spiralia, these inductive events have only been documented in equal-cleaving molluscs. Embryos of the nemertean Cerebratulus lacteus also undergo equal, spiral cleavage, and the fate map of these embryos is similar to that of other spiralians. The role of animal first quartet micromeres in the establishment of the dorsal (D) cell quadrant was examined in C. lacteus by removing specific combinations of micromeres at the eight-cell stage. To follow the development of various cell quadrants, one quadrant was labeled with DiI at the four-cell stage, and specific first quartet micromeres were removed from discrete positions relative to the location of the labeled quadrant. The results indicate that the first quartet is required for normal development, as removal of all four micromeres prevented dorsoventral axis formation. In most cases, when either one or two adjacent first quartet micromeres were removed from one side of the embryo, the cell quadrant on the opposite side, with its macromere centered under the greatest number of the remaining animal micromeres, ultimately became the D quadrant. Twins containing duplicated dorsoventral axes were generated by removal of two opposing first quartet micromeres. Thus, any cell quadrant can become the D quadrant, and the dorsoventral axis is established after the eight-cell stage. While it is not yet clear exactly when key inductive interactions take place that establish the D quadrant in C. lacteus, contacts between the progeny of animal micromeres and vegetal macromeres are established during the interval between the fifth and sixth round of cleavage divisions (i.e., 32- to 64-cell stages). These findings argue that this mechanism of cell and axis determination has been conserved among equal-cleaving spiralians.     

Unifying Concepts Learned from Methyl Farnesoate for Invertebrate Reproduction and Post-Embryonic Development

Since the discovery that methyl farnesoate (MF), the unepoxidatedform of the insect juvenile hormone (JHIII), is produced bymandibular organs of numerous crustaceans, extensive evidencehas accumulated that this compound appears to perform similarfunctions in the Crustacea as JH performs in insects. A majorfunction of MF appears to be in enhancing reproductive maturation.This was first shown by indirect experimentation with eyestalkablation, which augmented MF production. Subsequently, directtreatments of several species of crustacea with MF showed thatreproductive maturation was enhanced. A second function of MF, similar to that of the JH of insects,is in the maintenance of juvenile morphology. This is especiallytrue in the late larval transformations into juveniles, whereMF plays an inhibitory role, as well as during the transformationof juveniles into adults. These results were inferred from eyestalkremoval experiments. In the case of the larval-juvenile transition,inhibitory results were also obtained with MF by direct hormonetreatments. However, the transition from very early larval stages,such as one nauplius stage proceeding to the next, which inmany cases also involves morphogenetic changes, may be occurringin the presence of MF. Indeed, MF appears to be stimulatoryto early postembryonic larval stages of Crustacea. Again, thisfunction of MF in Crustacea appears to be similar to functionsof JH in early postembryonic insects. However, it should bepointed out that there are many more "early" stages in Crustaceathan there are in insects, and very few of these cases havebeen investigated. When considering the animal kingdom and larval metamorphosis,the question may be raised whether there are other members ofthe JH family regulating metamorphosis and reproduction. Oneplausible example appears to be among certain annelids. Thetrochophores of Capitella respond to various juvenoids, butare most responsive, within one hour, to MF and eicosatrienoicacid. This latter compound is present also in adult annelids,where it has been named "Sperm Maturation Factor," since itseems to function in the maturation of sperm in Arenicola. Therefore,eicosanoids perform in annelids two functions performed in insectsby JHs. In conclusion, it seems that there are morphogenesis promotingresponses to JHs in early larval development in crustaceans,annelids, and possibly other forms, which differ from thoseMF effects in later larvae of Crustacea where MF retards morphogenesis.Such early responses as noted here have recently also been describedfor insects. Furthermore, it is clear that the polyunsaturated8,11,14-eicosatrienoic and aracidonic acids seem to be juvenoids,and appear to function as such in annelids, and may also befunctionally active in insects and crustaceans. It seems reasonableto conclude therefore that new and novel juvenoids exist, whileothers still await discovery.     

Phylogenetic history of sifakas (Propithecus: lemuriformes) derived from mtDNA sequences

The sifakas (Propithecus) include three species containing up to 10 described subspecies, whose evolutionary relationships remain contentious. In particular, it is unclear whether P. verreauxi deckeni and P.v. coronatus populations are differentiated at the subspecific level. Furthermore, the taxonomic status of the recently discovered P. tattersalli and its phylogenetic position also require further examination. About 2,400 bp of mitochondrial DNA sequence data from part of the COIII gene, together with complete genes for ND3, ND4L, ND4, and five tRNAs, were used to clarify relationships among Propithecus species and subspecies. All analyses group Avahi as the sister group to all sifakas. P. diadema is placed as a sister group to all other Propithecus. Among the remaining sifakas, one subclade is formed by Puv. coquereli and P. tattersalli, while P.v. verreauxi, P.v. deckeni, and P.v. coronatus form the second subclade. All analyses fail to resolve P.vu. coronatus and P.v. deckeni into separate monophyletic lineages. Based on pairwise distance comparisons and tree topology, we conclude that P. tattersalli does not represent a distinct species and that P.v. deckeni and P.v. coronatus do not deserve subspecific rank. On the other hand, our analyses indicate that P.v. coquereli may well represent a separate species.     

Experimental evidence for the origins of determinative development in the polyclad turbellarians

Cell-deletion experiments were carried out on the embryo of the polyclad turbellarian Hoploplana inquilina to examine further the nature of development in primitive spiralians. The polyclads are of particular interest because they provide a link between the regulative development of acoels and the determinative development of annelids and molluscs. Single blastomeres were deleted at the two- and four-cell stages by puncture through the eggshell membrane with tungsten needles. All deletions resulted in abnormal larvae with consistent characteristics representing half or three-quarter Müller's larvae. The number of larval eyes was a particularly useful character in revealing mosaicism. This study establishes the polyclad embryo as determinative, but with important cell interactions also occurring during early development, and provides evidence that mosaicism became associated with spiral cleavage in the quartet form during the evolution of the Turbellaria.     

The polychaete Platynereis dumerilii (Annelida): a laboratory animal with spiralian cleavage, lifelong segment proliferation and a mixed benthic/pelagic life cycle

Platynereis dumerilii, a marine polychaetous annelid with indirect development, can be continuously bred in the laboratory. Here, we describe its spectacular reproduction and development and address a number of open research problems. Oogenesis is easily studied because the oocytes grow while floating in the coelom. Unlike the embryos of other model spiralians, the Platynereis embryo is transparent giving insight into the dynamic structures and processes inside the cells that accompany the prevailing anisotropic cleavages. Functional studies on cell specification and differential gene expression in embryos, larvae, and later stages are underway. Lifelong proliferation of uniform trunk segments qualifies Platynereis as a model for the study of gene expression and of the functional circuitry of this process. Platynereis can also become a stepping stone in the comparison of segmentation between annelids and arthropods because it comes closer to the putative ancestral morphology and style of development than other model annelids.     

Phylogenetic relationships of annelids,molluscs, and arthropods evidenced from molecules and morphology

Annelids and arthropods have long been considered each other's closest relatives, as evidenced by similarities in their segmented body plans. An alternative view, more recently advocated by investigators who have examined partial 18S ribosomal RNA data, proposes that annelids, molluscs, and certain other minor phyla with trochophore larva stages share a more recent common ancestor with one another than any do with arthropods. The two hypotheses are mutually exclusive in explaining spiralian relationships. Cladistic analysis of morphological data does not reveal phylogentic relationships among major spiralian taxa but does suggest monophyly for both the annelids and molluscs. Distance and maximum-likelihood analyses of 18S rRNA gene sequences from major spiralian taxa suggest a sister relationship between annelids and molluscs and provide a clear resolution within the major groups of the spiralians. The parsimonious tree based on molecular data, however, indicates a sister relationship of the Annelida and Bivalvia, and an earlier divergence of the Gastropoda than the Annelida–Bivalvia clade. To test further hypotheses on the phylogenetic relationships among annelids, molluscs, and arthropods, and the ingroup relationships within the major spiralian taxa, we combine the molecular and morphological data sets and subject the combined data matrix to parsimony analysis. The resulting tree suggests that the molluscs and annelids form a monophyletic lineage and unites the molluscan taxa to a monophyletic group. Therefore, the result supports the Eutrochozoa hypothesis and the monophyly of molluscs, and indicates early acquisition of segmented body plans in arthropods. Received: 25 September 1995 / Accepted: 15 March 1996     

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.     

Postembryonic development of dorsoventral and longitudinal musculature in Pycnophyes kielensis (Kinorhyncha, Homalorhagida)

We investigated the development of dorsoventral and longitudinalmusculature in all postembryonic stages of the kinorhynch Pycnophyeskielensis. Although the earliest stages have only 8 externallyseparated trunk segments, they already possess dorsoventralmuscles for 10 (prospective) trunk segments. The last, 11th,pair is added in the third juvenile stage. Longitudinal musculature,in contrast, is slower to develop and reaches its full lengthonly in the adult. In several juvenile individuals, single fibersproject from the longitudinal musculature into the followingsegments. In all juvenile stages, longitudinal muscles are continuousbetween segments, whereas in adults they are segmentally separatedfrom each other. Such late occurrence of a segmental patternin the longitudinal musculature is in contrast to patterns ofmuscle development in arthropods and annelids.     

D quadrant specification in the leech Helobdella: Actomyosin contractility controls the unequal cleavage of the CD blastomere

The unequal division of the CD blastomere at second cleavage is critical in establishing the second embryonic axis in the leech Helobdella, as in other unequally cleaving spiralians. When CD divides, the larger D and smaller C blastomeres arise invariantly on the left and right sides of the embryo, respectively. Here we show that stereotyped cellular dynamics, including the formation of an intercellular blastocoel, culminate in a morphological left-right asymmetry in the 2-cell embryo, which precedes cytokinesis and predicts the chirality of the second cleavage. In contrast to the unequal first cleavage, the unequal second cleavage does not result from down-regulation of one centrosome, nor from an asymmetry within the spindle itself. Instead, the unequal cleavage of the CD cell entails a symmetric mitotic apparatus moving and anisotropically growing rightward in an actomyosin-dependent process. Our data reveal that mechanisms controlling the establishment of the D quadrant differ fundamentally even among the monophyletic clitellate annelids. Thus, while the homologous spiral cleavage pattern is highly conserved in this clade, it has diverged significantly at the level of cell biological mechanisms. This combination of operational conservation and mechanistic divergence begins to explain how the spiral cleavage program has remained so refractory to change while, paradoxically, accommodating numerous modifications throughout evolution.     

Phoretic Behaviour of Bromeliad Annelids (<Emphasis Type="Italic">Dero</Emphasis>) and Ostracods (<Emphasis Type="Italic">Elpidium</Emphasis>) using Frogs and Lizards as Dispersal Vectors

The phoretic behaviour of ostracods (Elpidium bromeliarum) andannelids (Dero superterrenus) that inhabit tank bromeliads was studied. Our previous field observations had shown that bromeliad ostracods can be found attached to the skin of amphibians and reptiles that move among bromeliads, probably allowing the ostracods to colonise new tanks. In this paper, we present the first record of bromeliad annelids found attached to frogs moving among bromeliads in the field. We have also enlarged the database on bromeliad ostracods engaged in phoretic association with terrestrial vertebrates in three locations in southeastern Brazil. In our laboratory experiments bromeliad annelids show a strong significant tendency to climb onto papers that had been in contact with frog skin when compared with control papers, indicating a kind of chemically oriented behaviour. Bromeliad ostracods, on the other hand, attached themselves to treated and untreated papers with same frequency. When brought into contact with various species of frogs and lizards, the bromeliad annelids and ostracods both presented preference to attach themselves to frogs, but the annelids showed a stronger preference to attach to frogs and to avoid attachment to lizards. Another experiment demonstrated that bromeliad annelids are much more prone to dehydration than are ostracods. We suggest that the chemically oriented behaviour presented by bromeliad annelids toward frogs could diminish the risk of death by dehydration during the transport among bromeliads due to the moist characteristic of frog skins.     

Regeneration of posterior segments and terminal structures in the bearded fireworm,Hermodice carunculata (Annelida: Amphinomidae)

Like many other annelids, bearded fireworms, Hermodice carunculata, are capable of regenerating posterior body segments and terminal structures lost to amputation. Although previous research has examined anterior regeneration in other fireworm species, posterior regenerative ability in fireworms remains poorly studied. As the morphology of the anal lobe (a small, fleshy terminal structure of unknown function) has been used to distinguish East and West Atlantic H. carunculata populations, there is a more imminent need to understand the morphology and organization of tissues in specimens undergoing posterior regeneration, and the timeframe in which significant developmental changes occur. To further investigate this phenomenon, we amputated the posterior segments of living H. carunculata specimens collected from the Gulf of Mexico and monitored posterior regeneration over a 6‐month study period. Although many aspects of posterior regeneration in H. carunculata are consistent with the findings of other annelid regeneration studies, histological analysis revealed that once formed, anal lobe morphology remains relatively unchanged at all stages of posterior regeneration; East Atlantic morphotypes were not observed in the West Atlantic specimens studied here. Additionally, we found that the ventral nerve chord, which is partially responsible for the regeneration of lost body parts in polychaete annelids, terminates within the anal lobe, suggesting that this structure may play a role in the formation of new segments. J. Morphol. 275:1103–1112, 2014. © 2014 Wiley Periodicals, Inc.     

The complete mitochondrial genome sequence of Whitmania pigra (Annelida, Hirudinea): the first representative from the class Hirudinea

The mitochondrial genome is a significant tool for investigating the evolutionary history of metazoan animals. The currently available mitochondrial genome data in GenBank is limited to understand the detail evolutionary relationship among the metazoan animals, especially in the phylum Annelida. Here we present the mitochondrial gene organization, gene order and codon usage of the leech Whitmania pigra (Annelida), which is the first representative from the class Hirudinea. It is a circular molecule of 14,426bp, and encodes 13 protein-coding genes, 2 ribosomal RNA genes, and 22 transfer RNA genes. All 37 genes of W. pigra mitochondrial genome are transcribed from the same strand, which is identical to studied annelids, two echiurans, two sipunculans and many other lophotrochozoans. Five conserved gene clusters can be found in mitochondrial genomes of nine studied annelids, including (1) cox1-N-cox2; (2) cox3-Q-nad6-cob-W-atp6; (3) H-nad5-F-E-P-T-nad4L-nad4; (4) srRNA-V-lrRNA; and (5) nad3-S(1)-nad2. Compared with that of other studied annelids, translocations of transfer RNAs were found in the gene arrangement of W. pigra mitochondrial genome. Phylogenetic analysis strongly support that the species from Hirudinina and Oligochaeta form a monophyletic group Clitellata (BPM=100, BPP=100), which is consistent with previous research based on morphological and other molecular data. Both gene order data and amino acid sequences reveal that echiurans are derived annelids and sipunculans should be clustered with annelids and echiurans.