Giant pelagic larvae of Phyllodocidae (Polychaeta,Annelida)

The microscopic anatomy of giant pelagic larvae of Phyllodocidae was studied using routine histological, SEM, and TEM techniques. The larvae consist of two distinct regions: a large spherical trochophore measuring up to 2 mm in diameter and a posterior, long (up to 10 mm length), narrow rudiment of the adult body with up to 120 segments. The larvae have an unusual mixture of larval and adult features, including a very complex, well-developed brain and ganglia in the ventral nerve cord, and only a single pair of protonephridia located in the hyposphere of the trochophore. A muscular pharynx is not developed. The intestinal wall, especially in the trochophore region, consists of endodermal cells containing considerable nutritive material in the form of yolk-like globular inclusions. The digestive tract of all larvae was empty. The position of the frontal sensory organ and the prototroch, the structure of the parapodia and setae, and the three pairs of tentacular cirri dictate inclusion of the larvae in the family Phyllodocidae. The relatively enormous size and unusual pattern of development of the adult body may be adaptations for a long pelagic life and rapid settlement of the species, which inhabits slopes of islands and underwater mounts located far apart. J. Morphol. 238:93–107, 1998. © 1998 Wiley-Liss, Inc.     

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.     

Syllidae (Annelida, Polychaeta) from the Caribbean coast of Venezuela

Venezuela possesses a great variety of coastal environments allowing for a high diversity of marine species. However, systematic studies on marine invertebrates are scarce, especially on polychaetes. The family Syllidae is poorly known, and only 14 genera and 42 species have been reported from this country. A total of 13 genera and 26 species the Syllidae were identified from benthic samples collected on different substrata of the northeastern coast of Venezuela. Of these, seven genera and 16 species constitute new records for Venezuela: Odontosyllis guillermoi, Syllides floridanus, Salvatoria clavata, Salvatoria limbata, Sphaerosyllis longicauda, Parapionosyllis longicirrata, Trypanosyllis parvidentata, Trypanosyllis vittigera, Opisthosyllis sp., Syllis amica, Syllis armillaris, Syllis gracilis, Syllis pseudoarmillaris, Syllis vittata, Parasphaerosyllis indica and Myrianida convoluta.     

New records to Chile of the Family Paraonidae (Annelida: Polychaeta)

The Paraonidae are a polychaete family of small body size which have not been reported for Chile until recently. Mainly due to improved sample-processing methods, research campaigns carried out in 1994 and 1996 on three areas off southern Chile have yielded numerous records. Several species proved to be new to the Chilean polychaete fauna, including species that have been known previously only from Antarctic areas. These new records and range extensions are reported in this paper. Electronic Publication     

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.     

Ultrastructure of Chrysopetalid Paleal Chaetae (Annelida,Polychaeta)

Paleal notochaetae belonging to a number of Chrysopetalum species (Chrysopetalidae) were examined by scanning and transmission electron microscopy. Paleae are composed of broad medullary channels stacked with a regular series of horizontal fibrous diaphragms. The medullary part of the palea is surrounded by irregular rows of narrow tubular channels within the chaetal cortex. The origin and function of camerate chaetae is discussed.     

Unusual granules in the ejaculatory duct of a Microphthalmus species (Polychaeta,Annelida)

Summary The paired prominent ejaculatory ducts of the hermaphroditic polychaete Microphthalmus cf. listensis are surrounded by gland cells the processes of which penetrate the ducts themselves. These cells produce, in separate regions, two different types of spherical granules. Type I is composed of an electron dense and an electron lucent part. Type II granules contain a tubular filament that forms a single or double spiral in the periphery of a more or less unstructured electron dense material. Golgi vesicles give rise to this granule type. During the passage of sperm, these granules are obviously discharged into the lumen of the duct. Here they change form and probably dissolve. Their function is as yet unknown; capacitation of sperm is assumed.     

The epidermis of Timarete filigera (Polychaeta, Cirratulidae): histochemical and ultrastructural analysis of the gland cells

The aim of this study was to verify whether different living conditions of Polychaeta are correlated with morphological and functional differences in the organization of the integument. For this purpose, we decided to study the epidermis of Timarete filigera, a non-tubicolous polychaete. With this objective in mind, we have identified the various cellular types responsible for mucous secretion in the epidermis of this species and defined the histochemical composition of the mucus produced by different types of gland cells. Three types of gland cells have been identified by histochemical and ultrastructural studies in the epidermis of this polychaete. The histochemistry was carried out using standard techniques and peroxidase-labelled lectins. In type 1 cells, the secretory granules contain neutral glycoproteins with glucosidic residues of GalNAc, Galbeta 1,3 GalNAc, glucosidic and/or mannosidic residues. In type 2 cells, the secretory granules contain acid glycoproteins mainly sulphated with glucosidic residues of GalNAc, Galbeta 1,3 GalNAc, glucosidic and/or mannosidic residues, and some terminal sialic acid. In type 3 cells, the residual granules have the same chemical composition as that of granules present in type 2 cells. The secretion of these glandular mucous cells consists of mainly sulphated acidic glycoproteins and GAG resistant to testis jaluronidase. In these cells, the residual granules have the same chemical composition as that of their secretion. The heterogeneity of mucus composition may be correlated with its different functions.     

Global diversity of polychaetes (Polychaeta; Annelida) in freshwater

A literature review of Polychaeta (Annelida) including Aphanoneura (the oligochaete-like Aeolosomatidae and Potamodrilidae), living in freshwater yielded 168 species, 70 genera and 24 families representing all of the major polychaete clades, but less than 2% of all species. The best-represented families were, in order, Nereididae, Aeolosomatidae, Sabellidae, Spionidae and Histriobdellidae. Fourteen families were represented by a single species and genus. Regions supporting the highest diversity of freshwater polychaetes were in order, Palaearctic, Neotropical, Oriental, Nearctic, Australasian, and Afrotropical. More than half of all species and genera inhabitat lakes and rivers, followed by lagoons/estuaries, which have a high proportion of euryhaline species, and inland seas. Less common, atypical polychaete habitats include subterranean waters, the hyporheic zone of rivers and plant container habitats (phytotelmata). At least three distinct ecological/historical processes appear to account for the colonisation of continental waters: invasion of a clade prior to the break-up of Gondwana, as in Aphanoneura, Namanereis, Stratiodrilus, and Caobangia; relatively recent stranding of individual species (relicts); and the temporary visitation of euryhaline species. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Ultrastructural study of the jaw structures in two species of Ampharetidae (Annelida: Polychaeta)

Two species of jaw bearing Ampharetidae (Adercodon pleijeli (Mackie 1994) and Ampharete sp. B) were investigated in order to describe the microanatomy of the mouth parts and especially jaws of these enigmatic polychaetes. The animals of both studied species have 14–18 mouth tentacles that are about 30 µm in diameter each. In both species, the ventral pharyngeal organ is well developed and situated on the ventral side of the buccal cavity. It is composed of a ventral muscle bulb and investing muscles. The bulb consists of posterior and anterior parts separated by a deep median transversal groove. In both species, the triangular teeth or denticles are arranged in a single transversal row on the surface of the posterior part of the ventral bulb just in front of its posterior edge. There are 36 denticles in Adercodon pleijeli and 50 in Ampharete sp. B. The height of the denticles (6–12 µm) is similar in both species. Each tooth is composed of two main layers. The outer one (dental) is the electron‐dense sclerotized layer that covers the tooth. The inner one consists of long microvilli with a collagen matrix between them. The thickness of the dental layer ranges from 0.95 to 0.6 µm. The jaws of the studied worms may play a certain role in scraping off microfouling. The fine structure of the jaws in Ampharetidae is very similar to that of the mandibles of Dorvilleidae, the mandibles and the maxillae of Lumbrineridae, Eunicidae and Onuphidae, and the jaws of other Aciculata. This type of jaw is characterized by unlimited growth and the absence of replacement. The occurrence of jaws in a few smaller Ampharetidae is considered as an apomorphic state.     

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.     

Ultrastructure and mineral composition of serpulid tubes (Polychaeta,Annelida)

We have studied the tube ultrastructure of 44 recent species from 36 serpulid genera. Twelve distinct ultrastructures are identified. Serpulids possess very diverse tube ultrastructures, in contrast with the traditional point of view. Most species show single‐layered tubes, but 34% of these species have between two and four ultrastructurally different layers. Tubes are mostly bimineralic, and are composed of aragonite and calcite; however, one of the polymorphs is always dominant. All the studied single‐layered tubes with a lamello fibrillar tube ultrastructure are exclusively calcitic; prismatic structures, both in regular or irregular orientation, are either calcitic or aragonitic in composition. There is no correlation between tube mineralogy, and ultrastructure, and marine, brackish, and freshwater environments. We find that 47% of the serpulid species studied possess a unique combination of tube structure characters. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 633–650.     

Further revisions of the Sabellidae subfamilies and cladistic relationships among the Fabriciinae (Annelida: Polychaeta)

Cladistic relationships of the sabellid subfamilies Fabriciinae and Sabellinae are examined in light of recent revisions of fabriciin taxa. The potential placement of Caobangia in the Sabellinae is suggested from an initial analysis of selected fabriciin species and genera. Subsequent cladistic analyses at the family level produced over 1800 cladograms, among which Caobangia is the most plesiomorphic taxon of either the Fabriciinae or Sabellinae. Successive approximations weighting reduced this ambiguity, consistently placing Caobangia in the Sabellinae. Subfamilies are emended on the basis of these results. Cladistic relationships of fabriciin taxa, exclusive of Caobangia , display topological instability among some genera and species. Genera are, however, monophyletic in all topologies. Incorporating ontogenetic data for branching ventral filamentous appendages in Augeneriella reduces ambiguity among genera and suggests alternative transformation series for ventral filamentous appendages.     

Factors influencing latitudinal pattern of biodiversity: An example using Sabellidae (Annelida, Polychaeta)

The latitudinal gradient of biodiversity hypothesis was tested utilising the geographical distribution of species taxa belonging to the family Sabellidae (Annelida: Polychaeta), one of the polychaete families studied by only a few specialists around the world. An increase in species richness towards the Tropical region was revealed, with an asymmetric distribution between the two hemispheres. Data show that most of the species were found in Northern latitudes, producing a Northern area much richer than the Southern one. The Atlantic-Mediterranean Region was particularly rich, with the number of species close to those from the Indo-Pacific Region. Mediterranean species richness is discussed in the light of the particular history of the basin, also emphasising the importance of naturalistic studies that have made the Mediterranean marine flora and fauna one of the best known in the world. The existence of extrinsic factors that obscure the real pattern of distributions is hypothesised. Differences found between the two hemispheres could be attributed primarily to the higher number of specialists working in the Boreal Domains, including the Mediterranean area. Global trends produced by some representative genera are also discussed with respect to the distribution of the taxonomic specialists.     

Morphology and ultrastructure of the nephridial system of Hypania invalida (Grube, 1860) (Annelida,Polychaeta, Ampharetidae)

The excretory organs of the freshwater polychaete Hypania invalida have been examined using scanning and transmission electron microscopy. Three pairs of macroscopically and ultrastructurally different nephridia are present in the thorax. Intersegmental septa in the thorax are absent, with the exception of a single diaphragm between second and third chaetiger. The first pair of nephridia is anterior to this septum, the second pair crosses the septum, with the nephrostomes anterior and the ducts and the nephridiopori posterior to it, and the third pair of nephridia is entirely posterior to the diaphragm. The first two pairs of nephridia have ciliated nephrostomes of moderate size and long nephridial ducts that extend the length of the thorax. In contrast, the third pair is characterized by short ducts and very prominent nephrostomes. Macroscopically, seven different sections of nephridial duct cells can be distinguished along the length of the first two pairs of nephridia, whereas, on an ultrastructural basis, only six different regions can be identified. Only two regions of different duct cells can be recognized in the third pair of nephridia. Cells of the two anterior pairs of nephridia show typical characteristics of transport epithelia and most likely function as excretory organs. In contrast, the duct cells of the third pair are not that much differentiated and might primarily be responsible for the release of sexual products, as sperm was observed passing through these ducts. Podocyte‐like cells were observed to accompany nephridial ducts. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Fine structure of the spermatozoon of <Emphasis Type="Italic">Diopatra neapolitana</Emphasis> (Polychaeta,Onuphidae)

The identification of Diopatra species lacks of clear diagnostic features of taxonomic importance and the knowledge of their reproductive characters is scant. The spermatozoa of Diopatra neapolitana were ultrastructurally investigated by electron microscopy in order to correlate the mode of reproduction with sperm cells morphology. The mature male gamete has a depressed subspherical nucleus, a cone-like acrosome, and a long flagellum. The acrosome is conical in shape and radially symmetrical, with a base diameter twice the height. Within the acrosome vesicle, the basal region includes a very electron-dense thickened ring composed of paracrystalline substances. The subacrosomal space is filled with a poorly electron-dense material, with straight filaments axially arranged to form a perforatorium. The nucleus contains the complete axial canal, holding the hind perforatorium region. The middle piece consists of five mitochondria with well-distinct membranes and tubulo-vesicular cristae. Two centrioles are located perpendicularly to each other. The proximal one lies in the central fossa and the distal one, slightly eccentric to the sperm axis, anchors to the plasma membrane by nine satellite rays of the pericentriolar complex. The axoneme has a 9+2 arrangement of microtubules. In general, the spermatozoon of D. neapolitana conforms exteriorly to the typical ect-aquasperm; the acrosome complex ultrastructure, however, shows noticeable modifications from the basic form. This finding agrees with the previously observed reproductive pattern (broadcast spawning—free-swimming larvae) of D. neapolitana belonging to Santa Gilla population, and may be helpful to solve the taxonomic problems of the D. neapolitana complex as well.     

Ultrastructure and development of the nephridia in Anaitides mucosa (Annelida,Polychaeta)

Different developmental stages (trochophores, nectochaetae, non-mature and mature adults) of Anaitides mucosa were investigated ultrastructurally. A. mucosa has protonephridia throughout its life; during maturity a ciliated funnel is attached to these organs. The protonephridial duct cells are multiciliated, while the terminal cells are monociliated. The single cilium is surrounded by 14 microvilli which extend into the duct lumen without coming into any contact with the duct cells. Corresponding ultrastructure and development indicate that larval and adult protonephridia are identical in A. mucosa. Differences between various developmental stages can be observed only in the number of cells per protonephridium. A comparison between the funnel cells, the cells of the coelothel and the duct cells reveals that the ciliated funnel is a derivative of the duct. Due to the identical nature of the larval and postlarval protonephridia, such a funnel cannot be a secondary structure. In comparison with the mesodermally derived metanephridial funnel in phoronids it seems likely that the metanephridia of annelids and phoronids evolved convergently.     

A scanning electron microscopic study of secondary lamellae and chloride cells of rainbow trout (Salmo gairdneri)

Summary Scanning electron micrographs of gill tissue from rainbow trout fixed with 50% glutaraldehyde revealed the presence of microridges on surfaces of epithelial cells of the secondary lamellae. These microridges vary in length from 1 to 7 , with a mean height of 0.75 . Calculations show that they increase the total lamellar epithelial surface area approximately 2.5 fold. Mucus secreting cells are present on the body of the filament and on secondary lamellae. Chloride cells are located primarily in the interlamellae filamental epithelium and on the basal area of lamellae. Extensions of the chloride cell epithelium are microvillous in nature and their height is only slightly greater than that of the microridges of typical lamellar epithelial cells. A reduction in number or complete absence of microvilli on chloride cells appeared to be related to degenerative changes in these cells observed in transmission electron micrographs. Non secretory interlamellae filamental epithelial cells have microridges of very attenuated lengths.This research was supported by EPA Grant R-801034, USPHS Training Grant HL-05873, the Mich. Agr. Exp. Sta., Proj. 122 (Journal Article No. 5801), and OWRR Grant A-064. Acknowledgements: The authors wish to express their gratitude to Mrs. J. Mack and Mr. Wm. McAffe for their technical assistance with the electron microscopes.     

On the phylogeny of the Phyllodocidae (Polychaeta Annelida): an alternative

The phylogeny of the Phyllodocidae (Polychaeta Annelida) is examined in an analysis which includes previously unnoticed characters of the nervous system. Concentrating on four problems of importance in understanding the phylogeny of the family, a study of the nervous system demonstrated that: 1. A median antenna, such as that present in species of Eulalia, is not homologous to the nuchal papilla of, for example, Eteone and Phyllodoce; 2. Nuchal organs, presumably found in all phyllodocids, are basically homologous, but four separate types may be recognized; 3. The anterior concentration of the first segments (particularly their tentacular cirri), which is evident in most phyllodocids, is accompanied by different alterations of the ventral nervous system. The polarities of these changes are not self-evident; 4. A study of the ventral nervous system of Eteone reveals that the first apparent segment, bearing two pairs of tentacular cirri, must be regarded as homologous to the second segment of all other phyllodocids. Furthermore, this genus descended from forms in which a segment was present anterior to the first apparent segment, the first two segments possibly bearing tightly packed tentacular cirri. In connection with other characters, this information was used to produce a cladogram, which has some surprising features. The most unexpected result is the prediction that Eteone is the sister group of a taxon containing the genera Paranaitis, Chaetoparia, and Phyllodoce. The results are discussed and compared to those of other investigators.