Ultrastructure of the Genital Organs in the Interstitial Syllid Petitia amphophthalma (Annelida,Polychaeta)

Abstract The gonochoristic syllid Petitia amphophthalma is one of the truly interstitial polychaetes. P. amphophthalma does not show any epitokous modifications at maturity such as those that usually occur in syllids. The reproductive structures are unique: the male genital organs consist of a seminal vesicle in chaetigers 6–10, subdivided into a dorsal part tightly filled with spermatozoa and a ventral part with contents in different stages of spermatogenesis, one pair of sperm ducts and conspicuous gland cells situated in chaetigers 10 and 11. Their glandular secretions are discharged into the sperm duct together with those of other types of gland cells that form the duct. The oocytes develop freely within the body cavity of the females. Each of the fertile segments possesses a paired oviduct ending in a large ciliated funnel. Sperm ducts and oviducts are probably modifications of excretory organs; nephridia are absent in segments where gonoducts occur. A direct sperm transfer by lytic opening of the integument of the female and internal fertilization are inferred. Copyright © 1996 Published by Elsevier Science Ltd on behalf of the Royal Swedish Academy of Sciences     

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.     

Anatomy and ultrastructure of ventral pharyngeal organs and their phylogenetic importance in Polychaeta (Annelida)

Summary A comparative anatomical and ultrastructural study of ventral pharyngeal organs (pharyngeal bulbs) was carried out in two species of the Dinophilidae: Dinophilus gyrociliatus and Trilobodrilus axi. Special attention was paid to the fine structure of the stomodeal epithelium, cuticle, glands, muscles, and myoepithelial junctions. The differences between the species are very slight. The pharyngeal organ of the Dinophilidae is characterized by the following features: solid muscle bulbus made up of muscle cells only, bulbus muscle cells with two myofilament systems crossing at an angle of about 90°, gap junctions between these muscle cells, bulbus projects into a pharyngeal sac and bears rostrally a specific epithelium and cuticle, no bulbus glands, no investing (= sagittal) muscles, specific cuticle ultrastructure, cilia of ascending oesophagus with asymmetric tips, specific structure and position of salivary gland openings. The phylogenetic importance of these structures is discussed. Some of these characters are clearly autapomorphic features of the Dinophilidae and no common derived structures to other families with a ventral pharyngeal organ are present. Therefore, it is most likely that the dinophilid pharyngeal organ evolved independently. These findings do not agree with the hypothesis of the unity of the archiannelid families (Polygordiidae, Protodrilidae, Saccocirridae, Nerillidae, Dinophilidae, and Diurodrilidae) established on the basis of an assumed structural similarity of their ventral pharyngeal organs.Abbreviations bb basal body - bep bulbus epithelium - bl basal lamina - bm bulbus muscle - c cilium - cc coelenchyme cell - cm circular muscle - cr caudal rootlet - cu cuticle - dblm dorsal bulbus longitudinal muscle - dlm dorsal longitudinal muscle - dsn dorsal stomatogastric nerve - dy dyad - el electron-dense layer - fl fibrous layer - fi filaments - g Golgi apparatus - gl gland cell - hv homogeneous vesicle - l lipid droplet - la external lamina - lal lamellar layer - ll lower lip - lm longitudinal muscle - ly lysosome - m mitochondrion - mo mouth opening - mt microtubule - mv microvillus - mvp microvillar process - n nucleus - nu nucleolus - oes oesophagus - pcom preoral commissure - phf pharyngeal fold - phl pharyngeal lumen - phs pharyngeal sac - pms peripheral myofilament system - r rootletlike structure - rer rough endoplasmic reticulum - rr rostral rootlet - s sarcoplasmic reticulum - sc salivary canal - scom suboesophageal commissure - sd septate desmosome - ser smooth endoplasmic reticulum - sg secretory granule - sgl salivary gland - sn stomatogastric nerve - st stomach - step stomodeal epithelium - tep transitional epithelium - tf tonofilaments - va vacuole - vlm ventral longitudinal muscle - vsn ventral stomatogastric nerve - z z-element - za zonula adherens     

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.     

Ultrastructure of Presumed Ocelli in Parenterodrilus taenioides (Polychaeta,Protodrilidae) and their Phylogenetic Significance

Abstract Three different types of presumed unpigmented ocelli have been found in the anterior end of Parenterodrilus taenioides, a small gutless interstitial polychaete. The type-1 ocelli are located in the palps and four ocelli have been found along the length of each palp. The type-2 and type-3 ocelli lie close together in the head segment and are located in posterior ganglionic expansions of the brain. There is one pair of the minute type-2 ocelli but at least two pairs of the type-3 organs, which are the largest ocelli. In each ocellus the sensory cells are of the ciliary type and possess two cilia whose plasma membranes branch into numerous microvilli. With the exception of the type-1 ocelli they consist of a sensory cell and a supportive cell. In each ocellus the supportive cell forms a thin cup-shaped envelope around the densely packed ciliary branches. The type-1 ocelli consist of a single cell forming an intracellular vacuole (phaosome) which contains less densely packed microvillus-like structures. In particular, the structure of these ocelli is compared with that in other polychaetes, with special emphasis on the remaining genera of the Protodrilida.     

A new genus and species of Dorvilleidae (Annelida, Polychaeta) from Bermuda, with a phylogenetic analysis of Dorvilleidae, Iphitimidae and Dinophilidae

Neotenotrocha sterreri gen. et sp. n. is described from Bermuda. It is a gonochoristic. interstitial polychaete exhibiting sexual dimorphism. The maximum length of males is only 140 μm. while females may be up to 255 μm long. They are thus the smallest polychactes known to contain a mature ovary. The new species is referred to the polychaete family Dorvilleidae, primarily due to the presence of a etenognath jaw apparatus. In almost all external characters the new form is strongly reduced. lacking parapodia, setae, antennae, and palps, whereas larval characters such as trochae and a neurotroch are retained in adults. In the light of earlier literature on phylogeny within Dorvilleidae and the likelihood that Iphitimidae and Dinophilidae are closely affiliated with the latter family, the unique combination of characters in Neotenotrocha is of special interest. A phylogenetic analysis of all apparently valid genera in the three families leads to the hypothesis that Dinophilidae and Iphitimidae represent monophyletic and paraphyletic subgroups. respectively, within Dorvilleidae. Accordingly. the definition of Dorvilleidae is emended and a key to the genera provided. The species Ougia macilenta (Oug, 1978) is referred to Parougia Wolf. 1986 and the generic status of Meiodorvillea apalpata Jumars. 1974 and Protodorvillea gaspeensis Pettibone. 1961 is discussed. Some remarks on the applicability of concepts such as neoteny and progenesis in the characterisation of interstitial dorvilleids are included. On the basis of the cladogram mid these considerations. it is hypothesized that this group may have originated in the single evolutionary event.     

Ultrastructure and functional morphology of the female reproductive organs inProtodrilus (Polychaeta,Annelida)

The morphology and function of the female reproductive organs in 6Protodrilus species are investigated by light- and transmission electron microscopy. Possible ways in which spermatozoa may enter the female coelom after leaving the spermatophore are discussed for species with and without special female reception organs. Only femaleP. rubropharyngeus andP. flavocapitatus have “dorsal organs” for spermatophore reception. The structure and function of these organs are described, as well as those of the oviduct found in 3 of the species investigated. The possible phylogenetic origin of gonoducts and different modes of oviposition within the genus are discussed. Finally, the high taxonomic significance of female traits such as dorsal organs, oviducts, cocoon glands and lateral ciliary rows in this genus is stressed.     

Evaluation of the pair-culture effect inOphryotrocha puerilis (Polychaeta: Dorvilleidae). I. Pair-culture effect and sex ratio

Pairs and larger groups of femaleOphryotrocha puerilis puerilis were formed from formerly isolated specimens. Neither the diameter of the oocytes present in the coelomic fluid nor the number of setigerous segments (ss) of the partners of a newly formed pair allow us to predict which one of the two animals will exhibit sex reversal. Amputation of the palps showed that these ventrolateral appendages of the prostomium are not responsible for the transmission of the mutual influence which is exerted during the pair-culture effect. Isolated females do not produce egg masses but keep their oocytes in the body cavity until they are eventually resorbed. The shedding of oocytes in one of the females of a newly formed pair was formerly considered to be the first step in the pair-culture effect. The present results demonstrate that egg laying in these cases is unspecific and due rather to the end of isolation than to specific interactions with the partner. In groups consisting of up to 50 animals the sex ratio oscillates around 1:1. The presence of primary males does not influence the sex ratio of adult specimens, although these males are capable of fertilizing egg masses produced by adult females. As a result, a high percentage of both young and old males are found in densely populated bowls. The significance of the pair-culture effect in natural populations is discussed in the light of these findings.     

Phylogenetic relationships and evolution of Orbiniidae (Annelida, Polychaeta) based on molecular data

The phylogenetic relationships of orbiniid taxa were reconstructed based on sequence data of the mitochondrial 16S rRNA and nuclear 18S rRNA genes. Both genes were analysed separately and in combination using maximum likelihood, Bayesian inference and maximum parsimony. Regardless of the method used, a clade consisting of the investigated Orbiniidae, Methanoaricia dendrobranchiata and Questa was strongly supported by the 18S dataset. The analysis of the combined dataset suggests inclusion of M. dendrobranchiata within the Orbiniidae with close relationships to species of Orbinia and Phylo, rather than as a sister taxon to all other orbiniids. Evidence is given for the paraphyletic status of Leitoscoloplos , Naineris , Orbinia , Phylo and Scoloplos , which represent the most species-rich genera of the Orbiniidae. It is thus reasoned that the morphological characters presently used for genus diagnosis are not informative for cladistic analysis. No support is found for the hypothesis that taxa of the Protoariciinae represent juveniles of Orbiniinae. Instead, in the case of Protoaricia oerstedi , strong support for a progenetic origin is found.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 59−73.     

Structure of the nephridium in Ophryotrocha puerilis (Polychaeta,Dorvilleidae)

Summary The nephridia of Ophryotrocha puerilis are segmental organs. The nephrostome opens at the posterior margin of a setigerous segment into the coelomic cavity of this segment. The nephridial canal is made up of about 15 cells. These cells form an S-shaped tubule which extends into the following segment. The lumen of the nephridial canal ranges from 2 to 7 m in diameter. The nephropore opens laterally on the ventral surface of the body wall.In cross sections, one, two, or three cells are seen forming the canal. The inner surfaces of the canal cells are of different appearances along the canal. Since no regular pattern of cell distribution was found along the canals of different nephridia it is assumed that changes in cell structure along the canal are due to functional states or properties rather than to anatomically fixed regional differences. The canal cells either show smooth contours or they form brush borders of microvilli or sponge-like inner surfaces with a system of vacuolar canals running through the cytoplasm. Most of the canal cells are filled with various kinds of vesicles. Usually two or three cells contain larger vesicles up to 2.5 m in diameter with more or less electron-dense contents. Some of these vesicles resemble lysosomes. There are at least three bundles of cilia in each canal. In young specimens the number of cilia in one bundle is smaller (10–15) than in adult specimens (60–70). The nephridia do not show sex specific differences. The female nephridia do not function as genital ducts. As judged from the sizes of sperm and nephridia it appears to be possible that sperm are shed via male nephridia.     

Evaluation of the pair-culture effect inOphyryotrocha puerilis (Polychaeta: Dorvilleidae). II. Conditions for the moult of the upper jaw

The conditions for moult of the upper jaw ofOphryotrocha puerilis were determined in isolated individuals and in groups of various sizes. The frequency of formation of the complicated upper jaw in both isolated individuals and in groups varies to a considerable extent. Although formation of the upper jaw and sex reversal normally are associated processes, the relationship of these two processes is not very well understood. Histological investigations at the light microscopic level demonstrate that the dental apparatus is an elaboration of the ectodermal stomodaeum which is considered to be a highly specialized part of the cuticle in the pharyngeal region of the gut.     

The development of the serpulid Pomatoleios kraussii (Annelida, Polychaeta)

The development of Pomatoleios kraussii is extremely similar to that of Pomatoceros triqueter , and is completed under laboratory conditions in 1–3 weeks at 25–27°C. Mature larvae settle and metamorphose preferentially on the shells of adult conspecifics.     

Exchanges of sodium and chloride at low salinities by Nereis diversicolor (Annelida, Polychaeta).

1. Experiments to compare the exchange (total influx) of sodium and chloride in the polychaete Nereis diversicolor in steady-state adaptation to very low salinities are reported. 2. The Na-uptake mechanism shows a high affinity for sodium, reaching half the maximal uptake rate at an external Na-concentration of 8-10 mM/liter (ca. 2% SW), and becomes "saturated" or reaches a plateau of uptake at concentrations of 40-50 mM/liter (ca. 10% SW) up to ca. 350 mM/liter (75% SW), above which Na-exchange is proportional to the external concentration. 3. The Cl-uptake curve differs from the Na-uptake curve in showing a relative depression at very low salinities before reaching "saturation" at Cl-concentrations of 50-60 mM/liter (ca. 10% SW). Cl-uptake becomes proportional to external concentration in salinities of 50% SW or greater, suggestive of passive diffusion in the ionic and osmotic conforming range. 4. It is shown that the permeability of the body wall, both to Na and to Cl, is reduced at very low salinities, thus destroying one of the assumptions upon which a previously-presented balance-sheet for chloride exchanges in N. diversicolor was based (Smith, 1970a). 5. Attempts to demonstrate an activation of the Na-uptake mechanism at very low salinities were inconclusive; reduction of body-wall permeability to sodium masks any possible activation. 6. It is suggested that the inside-negative body-wall potential is related to the depression of the Cl-uptake curve in salinities below 10% SW.     

Morphology and Life History of Ophryotrocha maculata sp. n. (Polychaeta, Dorvilleidae)

The type locality of the new species is a muddy bottom at a depth of 25 meters in the Gullmarsfjord on the Swedish west coast. On that locality the fauna is recovering from previous sulphite pulp mill pollution. The shape and morphology of O. maculata conform with most other members of the species. It is distinguished by segmental transverse rows of dark spots. It is a contemporary hermaphrodite with anterior male and posterior female segments. Different from other known species the germ cells are released directly into the water. As a consequence the brood is not protected by the parents.     

Feeding behavior of the cirratulid Cirriformia filigera (Delle Chiaje, 1825) (Annelida: Polychaeta).

Observations of the feeding behavior of Cirriformia filigera (Delle Chiaje, 1825) (Annelida: Polychaeta) from the intertidal zone of S?o Francisco and Engenho D'água beaches (S?o Sebasti?o, State of S?o Paulo) were made in the laboratory. This species, like other cirratulids, is a deposit feeder, feeding mainly on sediment surface with the aid of its grooved and ciliated palps, which are used to capture food particles. The worm lies just beneath the substrate surface in a J-shaped tube. When feeding, it extends up to 4 palps over the sediment surface, capturing food particles which pass down the groove of each palp directly to the mouth. Only fine sand grains are ingested. The worm frequently extends 4 branchial filaments into the overlying water for aeration. When it moves with the prostomium sideways, it collects and transports sand grains that pass backwards along its ventral region until reaching the middle part of its body. Next, the parapodia and palps move the sand grains to the dorsal posterior end of the animal, covering this area with sand. Some sand grains are also ingested as the worm moves.