Chaetal arrangement provides no support for a close relationship of Sabellidae and Sabellariidae (Annelida)

Sabellid and sabellariid polychaetes are regarded as sister groups in a number of recent phylogenetic analyses. This is based mainly on a shared specific arrangement of chaetae referred to as chaetal inversion. Remarkably, the uncini have a notopodial position in the abdomen, whereas capillary chaetae occur in the neuropodia in both taxa in contrast to the situation in putative relatives. However, in sabellids uncini and capillary chaetae change their position completely at the border between thorax and abdomen, whereas uncini are missing in the parathorax of Sabellariidae. Due to this difference the significance of the chaetal inversion for systematics has been subject to discussion for years. Serial semithin sections of parapodia of the Sabellidae Sabella pavonina, Branchiomma bombyx, Fabricia stellaris, and of the Sabellariidae Sabellaria alveolata were studied in order to obtain detailed information on their chaetal arrangement and sites of chaetal origin. SEM investigations and computer-aided 3D-reconstructions provide deep insight into the spatial organization of the rami. Though differing externally, the principal chaetal arrangement and the location of the formative sites turned out to be almost identical within the species of Sabellidae. Most chaetae are aligned in straight transverse rows with a dorsal site of origin within neuropodia and a ventral one in notopodia as is common in sedentary polychaetes. Semicircular and spiral arrangements are revealed to be modified transverse rows. Only in thoracic notopodia does an additional dorsocaudal formative site form distinct rows. The chaetal inversion in Sabellidae is additionally characterized by an abrupt change of capillary chaetae and uncini along with a sudden change of the parapodial morphology at the border between thorax and abdomen. All chaetae of S. alveolata are aligned in transverse rows with the same location of the formative sites as in sabellids and other sedentary polychaetes. However, in contrast to sabellids the chaetae are not inverted across a parathoracic abdominal border. Moreover, there is no inversion of the parapodial structure from parathorax to abdomen and the neuropodial chaetal composition changes gradually from parathorax to abdomen. The chaetal arrangement in Sabellariidae thus cannot be described as inverted along the body-axis as in Sabellidae. Evolutionary steps implied by the assumption of an inverted chaetal pattern in a supposed common ancestor are discussed. It is concluded that the specific chaetal arrangement of Sabellidae and Sabellariidae arose independently and therefore provides no support for a sistergroup relationship of sabellids and sabellariids.     

Chaetal arrangement in Orbiniidae (Annelida,Polychaeta) and its significance for systematics

The systematic position of Orbiniidae within Polychaeta is still uncertain. In order to provide additional comparative data, we investigated the chaetal arrangement in this family, which is considered valuable for polychaete systematics. Specimens of Scoloplos armiger, Orbinia latreillii, and Pettibonella multiuncinata were examined by SEM and serial sections analysed by computer aided 3D-reconstructions. The obtained data suggest that the chaetal arrangement of Orbiniidae resembles that of other sedentary polychaetes in only a few respects. Transverse rows are only present in the main, anterior part of the chaetal patches of thoracic neuropods. The position of the formative site indicates homology with the transverse rows of several sedentary polychaete taxa. The chaetal patches thus differ significantly from those known in Apistobranchidae. Independent rows with an own caudal formative site, which run along the caudoventral edge of the chaetal patches, resemble the neuropodial ventral longitudinal rows known in Spionidae and related taxa. The abdominal neuropodia of S. armiger and O. latreillii bear longitudinal rows of chaetae. These are reorientated during ontogenetic chaetiger transformation and become the transverse rows of the thoracic chaetal patches. 3D reconstruction of S. armiger revealed that the notopodial chaetal bundles are organized in rows as well. Notopodia and abdominal neuropodia bear deep reaching supportive chaetae. They are the first chaetae formed during neuropodial development and reside dorsally to the longitudinal row of capillary chaetae. Neither position nor structure indicates homology with the supportive chaetae of other sedentary polychaetes. Spionidae and related taxa are thus the only sedentary polychaetes, which specifically resemble Orbiniidae in certain aspects of their chaetal arrangement. Dedicated to Prof. Dr. Wilfried Westheide on the occasion of his 70th birthday.     

Homology and Evolution of the Chaetae in Echiura (Annelida)

Echiura is traditionally regarded as a small phylum of unsegmented spiralian worms. Molecular analyses, however, provide unquestionable evidence that Echiura are derived annelids that lost segmentation. Like annelids, echiurans possess chaetae, a single ventral pair in all species and one or two additional caudal hemi-circles of chaetae in two subgroups, but their evolutionary origin and affiliation to annelid chaetae are unresolved. Since annelids possess segmental pairs of dorsal (notopodial) and ventral (neuropodial) chaetae that are arranged in a row, the ventral chaetae in Echiura either represent a single or a paired neuropodial group of chaetae, while the caudal circle may represent fused rows of chaetae. In annelids, chaetogenesis is generally restricted to the ventral part of the notopodial chaetal sac and to the dorsal part of the neuropodial chaetal sac. We used the exact position of the chaetal formation site in the echiuran species, Thalassema thalassemum (Pallas, 1766) and Echiurus echiurus (Pallas, 1767), to test different hypotheses of the evolution of echiurid chaetae. As in annelids, a single chaetoblast is responsible for chaetogenesis in both species. Each chaeta of the ventral pair arises from its own chaetal sac and possesses a lateral formation site, evidencing that the pair of ventral chaetae in Echiura is homologous to a pair of neuropodia that fused on the ventral side, while the notopodia were reduced. Both caudal hemi-circles of chaetae in Echiurus echiurus are composed of several individual chaetal sacs, each with its own formative site. This finding argues against a homology of these hemi-circles of chaetae and annelids’ rows of chaetae and leads to the hypothesis that the caudal chaetal rings evolved once within the Echiura by multiplication of ventral chaetae.     

Development of the deep‐sea viviparous quill worm Leptoecia vivipara (Hyalinoeciinae,Onuphidae, Annelida)

Development of Leptoecia vivipara, a brooding deep‐sea onuphid polychaete with a circum‐Antarctic distribution, was studied using light microscopy, scanning and transmission electron microscopy, and histology. All specimens examined were brooding viviparous females or juveniles; no male gametes were detected. Anterior segments of juvenile and adult worms bore paired compact ovaries with clusters of vitellogenic oocytes. In adults, the mid‐body region formed a chamber containing up to 12 offspring at different stages of development, from oocyte to 13 chaetigers. Mature oocytes freely floated in the coelomic fluid, while embryos and juveniles were enclosed in peritoneal envelopes. Chaetal replacement in juveniles and the morphology of the provisional maxillae are described. Leptoecia vivipara is argued to be a progenetic species with juvenile‐like external morphology and accelerated sexual maturation. These traits may have arisen as adaptations to epibenthic life in a high‐latitude deep‐sea environment affected by seasonal pulses of organic matter.     

Parapodial glandular organs in Spiophanes (Polychaeta: Spionidae)—studies on their functional anatomy and ultrastructure

Parapodial glandular organs (PGOs) of Spiophanes (Polychaeta: Spionidae) were studied using light and electron microscopy. These organs are found in parapodia of the mid body region, starting on chaetiger 5 and terminating with the appearance of neuropodial hooks (chaetiger 14 or 15 in adult individuals). Large PGOs in anterior chaetigers display different species‐specific types of openings whereas small PGOs in posterior parapodia of the mid body region always open in a simple vertical slit. Each PGO is composed of three main complexes: (1) the glandular sac with several distinct epithelia of secretory cells and secretory cell complexes and the reservoir filled with fibrous material, (2) the gland‐associated chaetal complex (including the region of chaetoblasts and follicle cells, follicular canals, two chaetal collector canals, the combined conducting canal, the chaetal spreader including the opening of the glandular organ with associated type‐1 secretory cells, and the gland‐associated chaetae), and (3) a bilayered musculature surrounding the gland. A considerable number of different cell types are involved in the secretory activity, in the guidance of the gland‐associated chaetae, and in the final expulsion of the fibrous secretion at the opening slit. Among these different cell types the type‐5 secretory cells of the proximal glandular complex with their cup‐shaped microvilli emanating thick microfibrils into the lumen of the glandular sac are most conspicuous. Secretory cells with cup‐shaped microvilli being involved in the production of β‐chitin microfibrils have so far only been reported from some representatives of the deep‐sea inhabiting Siboglinidae (Polychaeta). We suggest that the gland‐associated chaetae emerging from inside the PGOs of Spiophanes are typical annelid chaetae formed by chaetoblasts and follicle cells. Functional morphology implies the crucial role of PGOs in tube construction. Furthermore, the PGOs are discussed in consideration of phylogenetic aspects. J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

Intra- and interspecific ultrastructural character variation: The chaetation of the Microphthalmus listensis species group (Polychaeta,Hesionidae)

Summary The different chaetal types of three eidonomically very similar, closely related polychaete species (Microphthalmus carolinensis, M. nahantensis and M. listensis) were ultrastructurally investigated and compared with one another. Characteristic substructures are regularly arranged channels, the course and number of which were analysed in cross-sections. The intraspecific variability of the number of these channels is low in the individual chaetal type. Between the species, however, distinct differences exist, facilitating separation and identification of the species at the level of ultrastructure in certain chaetae.     

Locomotion and fine structure of parapodia in <Emphasis Type="Italic">Myzostoma cirriferum</Emphasis> (Myzostomida)

Most myzostomids are ectocommensals of crinoids on which they move freely. Their locomotion is ensured by five pairs of parapodia located laterally below their trunk. Each parapodium in Myzostoma cirriferum is a conical structure that includes a hook-like chaeta, replacement chaetae and an aciculum. Structure and ultrastructure of the myzostomid chaetae are similar to those of polychaetes: they are formed by a chaetoblast, which gives rise to microvilli where chaetal material is assembled on the outer surface. Myzostoma cirriferum walks on its host. It moves the anterior part, the posterior part or the lateral parts forwards but is able to rotate of 180° on itself. Its locomotion entirely depends on parapodial motions and not on trunk movements. Three pairs of muscles are involved in parapodial motions: parapodium flexor and parapodium extensor, aciculum protractor and aciculum retractor, and hook protractor with conjunctor. A functional model is proposed for explaining the global motion of a parapodium in M. cirriferum that may be extended to all ectocommensal myzostomids.     

Getting to the root of fireworms' stinging chaetae—chaetal arrangement and ultrastructure of Eurythoe complanata (Pallas, 1766) (Amphinomida)

Amphinomid species are since long known to cause urtication upon contact with the human skin. Since it has been reported that amphinomid chaetae are hollow, it has repeatedly been suggested that poison is injected upon epidermal contact. To test predictions for the structural correlate of such a stinging device we studied the structure and formation of chaetae in the fireworm Eurythoe complanata (Amphinomida). Neither the structure of the chaetae nor their formation and their position within the parapodium provide evidence for their function as hollow needles to inject poison. The chaetae even turned out to be not hollow, but containing calcareous depositions. The latter most likely cause artificial ruptures of delicate chitin lamellae in the inner of the chaeta when treated with acidic fixatives. Inorganic calcium compounds harden the chaetae and make them brittle so that they break easily. Additional information on the structure of the chaetal sac, the site of formation and the acicula do not contradict the position of the Amphinomida within Annelida as revealed by phylogenomic studies.     

On the absence of circular muscle elements in the body wall of Dysponetus pygmaeus (Chrysopetalidae, 'Polychaeta', Annelida)

The annelid body wall generally comprises an outer layer of circular muscle fibres and an inner layer of longitudinal muscle fibres as well as parapodial and chaetal muscles. An investigation of Dysponetuspygmaeus (Chrysopetalidae) with confocal laser scanning microscopy showed that circular muscles are entirely absent. Further studies indicate that this feature is characteristic for all Chrysopetalidae. A scrutiny of the literature showed a similar situation in many other polychaetes. This lack of circular muscle fibres may either be due to convergence or represent a plesiomorphic character. Since circular muscles are very likely important for burrowing forms but not necessary for animals which proceed by movements of their parapodial appendages or cilia, this problem is also related to the question of whether the ancestral polychaete was epi‐ or endobenthic.     

Ultrastructure of opisthosomal chaetae in Vestimentifera (Pogonophora, Obturata) and implications for phylogeny

Schulze, A. 2000. Ultrastructure of opisthosomal chaetae in Vestimentifera (Pogonophora, Obturata) and implications for phylogeny. — Acta Zoologica (Stockholm) 82 : 127–135
The posterior segmented body region of Vestimentifera bears rows of uncini that function to anchor the animal within its tube. SEM studies of five vestimentiferan species reveal intraspecific and interspecific variation in the number of chaetigerous segments and the arrangement of uncini within a given segment. The portion of an uncinus that extends beyond the epidermis comprises two opposing groups of teeth that probably correspond to the capitium and subrostral process of polychaete uncini, and a distinct protuberance between them, interpreted as a rostrum. In Ridgeia piscesae , the uncini are formed by chaetal follicles, consisting of a chaetoblast, a follicle cell and an epidermis cell. The chaetal shaft is elongate and composed of up to 40 hollow cylinders that are invaded at their base by microvilli from the apical part of the chaetoblast. Opisthosomal chaetae in perviate Pogonophora are usually restricted to four per segment and are of a rod-shaped type. It is hypothesized that the rod-shaped chaetae represent reduced hooked chaetae probably derived from a condition such as found in Monilifera. Uncini of Pogonophora, Sabellida, Terebellida and Oweniida are considered homologous but details of chaetal design may be due to functional adaptations and thus do not represent reliable characters for phylogenetic studies on higher taxonomic levels than genera or potentially families.     

Polychaete meets octopus: symbiotic relationship between Spathochaeta octopodis gen. et sp. nov. (Annelida: Chrysopetalidae) and Octopus sp. (Mollusca: Octopodidae)

Marine annelids in the subfamily Calamyzinae (family Chrysopetalidae) are either symbiotic or free-living forms that have been mainly reported from deep-sea chemosynthetic systems. Symbiotic calamyzines mainly live in the mantle cavity of bivalves in hydrothermal vents or cold seeps, but one species is also found to be inserted into the epidermis of polychaetes. We found a single specimen of calamyzine polychaete on the body surface of Octopus sp. collected in the Sea of Kumano (Japan), which represents the first known record of symbiotic association between polychaetes and octopuses. We described the specimen as Spathochaeta octopodis gen. et sp. nov. Spathochaeta gen. nov. can be discriminated from other genera in Calamyzinae by the presence of spatula-shaped notochaetae and dorsal chaetal lobes. We also provided the phylogenetic position of S. octopodis gen. et sp. nov. within Chrysopetalidae based on four gene markers (COI, 16S, 18S, H3). www.zoobank.org/urn:lsid:zoobank.org:pub:A8FB15C1-31A7-4487-966B-13F10E19A373.     

Chaetal arrangement and chaetogenesis of hooded hooks in Lumbrineris (Scoletoma) fragilis and Lumbrineris tetraura (Eunicida,Annelida)

As the structure and arrangement of chaetae are highly specific for annelid species and higher taxonomic entities, we assume that rather conservative information guarantees formation of specific chaetae. Each chaeta of an annelid is formed within an ectodermal invagination, and the modulation of the apical microvilli pattern of the basalmost cell of this invagination determines the structure of the chaeta. Any hypothesis of the homology of chaetae could thus be tested by examining the process of chaetal formation. Investigations into the ultrastructure and formation of hooded hooks in different capitellids and spionids revealed that these chaetae can be homologized. The hood of each of their hooded hooks is formed by elongation of two rings of microvilli peripheral to the chaetal anlage, which give rise to the inner and outer layers of the hood. The hood layers are well separated and surround an empty space. Superficially similar hooded hooks are described for certain Eunicida. Presently available cladistic analyses suggest that the hooded hooks of eunicidans evolved independently of those in Capitellidae and Spionidae. Compared with the latter two families, we therefore expected to find differences in chaetogenesis of the hooded hooks in the eunicids Lumbrineris (Scoletoma) fragilis and Lumbrineris tetraura (Lumbrineridae). This was the case. In these eunicidans, the hood was formed by the bisected apical wall of the chaetoblast right after the mid‐apical section of the chaeta had been sunk deeply into the chaetoblast during its formation. The apical wall generated a brush of microvilli that preformed the hood. Because the microvilli of the hood showed some accelerated differentiation, they soon merged with those of the slowly growing setal shaft to form the broad manubrium of the hooded hook in lumbrinerids. Our study confirms the predicted differences in chaetogenesis of the superficially similar hooded hooks of capitellids and spionids compared with those of eunicids.     

Textures and traction: how tube‐dwelling polychaetes get a leg up

By controlling the traction between its body and the tube wall, a tube‐dwelling polychaete can move efficiently from one end of its tube to the other, brace its body during normal functions (e.g., ventilation and feeding), and anchor within its tube avoiding removal by predators. To examine the potential physical interaction between worms and the tubes they live in, scanning electron microscopy was used to reveal and quantify the morphology of worm bodies and the tubes they produce for species representing 13 families of tube‐dwelling polychaetes. In the tubes of most species there were macroscopic or nearly macroscopic (~10 μm–1 mm) bumps or ridges that protruded slightly into the lumen of the tube; these could provide purchase as a worm moves or anchors. At this scale (~10 μm‐1 mm), the surfaces of the chaetal heads that interact with the tube wall were typically small enough to fit within spaces between these bumps (created by the inward projection of exogenous materials incorporated into the tube wall) or ridges (made by secretions on the interior surface of the tube). At a finer scale (0.01–10 μm), there was a second overlap in size, usually between the dentition on the surfaces of chaetae that interact with the tube walls and the texture provided by the secreted strands or microscopic inclusions of the inner linings. These linings had a surprising diversity of micro‐textures. The most common micro‐texture was a “fabric” of secreted threads, but there were also orderly micro‐ridges, wrinkles, and rugose surfaces provided by microorganisms incorporated into the inner tube lining. Understanding the fine structures of tubes in conjunction with the morphologies of the worms that build them gives insight into how tubes are constructed and how worms live within them.     

Biology of Pseudopotamilla reniformis (Müller 1771) in the White Sea,with description of asexual reproduction

The tube-dwelling polychaete Pseudopotamilla reniformis (Sabellidae) forms dense and complex aggregations of flexible tubes on hard substrates in the subtidal zone of the White Sea. No sexual reproduction was observed in this study and recruitment appeared to be due to asexual reproduction by architomy in winter, from October to March. The posterior part of the abdomen undergoes spontaneous fission into from 2 to 4 fragments and depending on their position, the fragments regenerate their anterior ends or both anterior and posterior ends. Regeneration in P. reniformis takes place via a combination of epimorphosis (replacement of missing parts by cell proliferation and the growth of new tissue) and morphallaxis (the remodelling of pre-existing structures without cell proliferation). The morphogenetic events during regenerative restoration include de novo formation of branchial crown, formation of thoracic segments and restoration of the posterior end. Asexual reproduction appears to play a crucial role for formation of P. reniformis aggregations and is very important for the population in the White Sea, at the margin of the species’ range.     

Chaetae and mechanical function: tools no Metazoan class should be without

To address the functional contributions of capillary chaetae in the maldanid polychaete Clymenella torquata, we compared irrigation efficiency and tube structure for animals with intact and trimmed capillary chaetae. We measured pumping rates for worms before and after they were anaesthetized and subjected either to capillary trimming or mock trimming, i.e. handling without trimming. Worms with trimmed chaetae were significantly less effective at moving water through their tubes than those with intact chaetae. There were no significant differences in the ability of control worms to move water within their tubes. No significant changes in rates of peristalsis were observed among experimental or control groups. These data strongly suggest that body musculature and capillary chaetae work in concert to hold worms in position within tubes during peristaltic pumping. When chaetae are shortened, the body musculature must contract to a greater degree, increasing the functional diameter of the worm to achieve the necessary traction with the tube wall, resulting in less efficient irrigation. We also compared the inner diameters of original field tubes to tubes built by control worms or worms after capillary trimming. The inner diameters of new tubes built by worms with shortened chaetae were larger than their original tubes, while those of both control groups were not. One possible explanation is that the chaetae have a sensory role and shortened chaetae send the false message that the nascent tube walls are farther away than they are, the body contracts in compensation and the tube is widened, however this idea has not been tested.     

F-actin framework in Spirorbis cf. spirorbis (Annelida: Serpulidae): phalloidin staining investigated and reconstructed by cLSM

Abstract. Serpulidae encompasses polychaete species whose members have fused anterior ends bearing a tentacular crown, a heteronomous segmented body with a thorax and abdomen, and “chaetal inversion” between the two tagmata. The sessile filter‐feeding organisms live in self‐built, coiled, calcareous tubes on algae. The F‐actin muscular subset of Spirorbis cf. spirorbis was stained with phalloidin and three‐dimensionally reconstructed by means of cLSM, aiming to investigate (1) how the tentacular crown is organized and moved, (2) whether the internal structures, e.g., musculature, follow the thorax–abdomen inversion, and (3) whether circular muscles are present in serpulids. The third aim is by reason of recent investigations suggesting that lack of circular muscle fibers may be a common situation rather than a rare variation in polychaetes. In this manner, this article is part of a comparative evaluation of polychaete muscle systems. We found that longitudinal muscles of the body wall project into the tentacular crown, and that radioli and pinnulae possess three muscle types each, facilitating their great mobility. Operculum, collar, and a pair of unidentified organs possess distinct F‐actin filaments. The trunk is mainly moved by five longitudinal muscle strands, most obvious in the abdomen: two dorsal, two ventral, and an unpaired ventromedian one, out of which the dorsal ones are the strongest. In anterior regions, the two dorsal strands form a single continuous layer; the separated strands lessen posteriorly. Solitary transverse fibers are located ventrally in the middle of each segment, stretching between longitudinal muscles and coelomic lining laterally, where they end. Peripheral and central dorsoventral muscles, two pairs per segment each, are present. Circular fibers as well as bracing muscles were not detected. The results indicate that the musculature does not follow the thorax–abdomen inversion and Serpulidae represents the 15th polychaete taxon in which circular fibers are totally missing.     

Muscular system in polychaetes (Annelida)

The structure of the polychaete muscular system is reviewed. The muscular system comprises the muscles of the body wall, the musculature of the parapodial complex and the muscle system of the dissepiments and mesenteries. Various types of organisation of the longitudinal and circular components of the muscular body wall are distinguished. In Opheliidae, Polygordiidae, Protodrilidae, Spionidae, Oweniidae, Aphroditidae, Acoetidae (=Polyodontidae), Polynoidae, Sigalonidae, Phyllodocidae, Nephtyidae, Pisionidae, and Nerillidae circular muscles are lacking. It is hypothesised that the absence of circular muscles represents the plesiomorphic state in Annelida. This view contradicts the widely accepted idea of an earthworm-like musculature of the body wall comprising an outer layer of circular and an inner layer of longitudinal fibres. A classification of the various types of parapodial muscle construction has been developed. Massive and less manoeuvrable parapodia composed of many components like those of Aphrodita are regarded to represent the plesiomorphic state in recent polychaetes. An analysis of the diversity of the muscular structure supports the hypothesis that the primary mode of life in polychaetes was epibenthic and the parapodial chaetae had a protective function.     

Characterization of six polymorphic microsatellites for the polychaete tubeworm Hydroides elegans and cross‐species amplification in the congener Hydroides hexagonus

We isolated and characterized six polymorphic microsatellite loci for the polychaete tubeworm, Hydroides elegans. Two additional loci were not reliably scorable and estimates of heterozygosity were obtained for the other six. In addition, cross‐species amplification was successful for two loci using the congener H. hexagonus. Given that few microsatellite loci are available for polychaetes, these markers will be useful in assessing dispersal and gene flow in H. elegans and probably also other polychaetes.     

On the associations between Haplosyllis (Polychaeta, Syllidae) and gorgonians (Cnidaria, Octocorallaria), with the description of a new species

The present paper includes a morphological, ecological and biological updating of the three gorgonian associated species of Haplosyllis (Polychaeta, Syllidae) known to date: H. chamaeleon (symbiont with Paramuricea clavata in the Mediterranean), H. anthogorgicola (symbiont with Anthogorgia bocki in the Japanese seas) and H. villogorgicola , a new species living symbiotically with Villogorgia bebrycoides which is only known from Tenerife (Canary Islands, Eastern Central Atlantic). The new species is described on the basis of ecological, morphological, morphometric and statistical analysis of relevant characteristics. Each host colony harboured about 15 pale-yellowish worms, whose cryptic colouration mimicked that of the host. They occurred either on the host branches or partly hidden inside cavities formed by the fusion of two branches. The new species is characterized by the presence of simple chaetae with clearly bidentate tips all along the body, the presence of gland pore aggregates distributed in two lateral rows and two ventral patches on each palp and the absence of ciliary tufts on the pharyngeal papillae. H. villogorgicola sp. nov. is closely related to H. chamaeleon . Thus, it is compared with two populations of this species collected in the north-west and south-west Mediterranean. Stolons of H. chamaeleon are re-described as tetracerous and a peculiar posterior end regeneration process occurring in adult worms during the stolon formation is described. H. anthogorgicola is also re-described, with particular emphasis on its appendage and chaetal arrangements. The main features of the three associations are discussed in light of the current knowledge on symbiotic polychaetes, particularly cnidarian-associated syllids.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 455–477.     

Zur Morphologie und Ökologie vonPolydora ciliata undP. ligni (Polychaeta,Spionidae)

The polychaete wormsPolydora ciliata andP. ligni were investigated with regard to the morphology of their fifth chaetigerous segment, bearing bundles of modified bristles in a special arrangement. The development of these chaetae from late larval to adult stages is described, considering the loss of provisional bristles, the variation of shape caused by the wear and the shedding of old chaetae after losing their function. In addition an epidemic shell disease of the mussel,Mytilus edulis, induced by infestation withP. ciliata is reported.P. ligni was observed to regularly inhabit the inflorescenses of the eelgrass,Zostera marina. The relationships between this plant andP. ligni are discussed.