A Study of the Morphology and Biology of Thompsonia littoralis (Crustacea: Cirripedia: Rhizocephala)

Interna development within the intertidal tropical crab, Leptodius exaratus, requires up to 4.5 months; the roots become distributed from the CNS along the major nerves and, when the crab moults, results in emergence of numerous externae located symmetrically on the abdomen and bases of the thoracic appendages. As many externae drop off, their number is continuously reduced; the survivors reach sexual maturity and ovulate after 2.5 months and complete embryonic development after 2 more months. At the next moult a new and more numerous generation of externae appears, and this is repeated until the crab dies. Presence of externae significantly prolongs the host's normal intermoult period and minimizes its growth. The parasite has little effect on the host's sexual characters although females are normally sterilized. The cuticle of the externa is moulted once, leaving a distinct annulus around the stalk. Spermatogonia are presumably injected by male cyprids into the female externa and develop into sperm within one or more vesicular spermatogenic bodies in its mantle. The eggs are laid through a single oviduct into the mantle cavity where they become fertilized and develop into cyprids that are liberated at the disintegration of the externa.     

Thompsonia dofleini,a colonial akentrogonid rhizocephalan with dimorphic,ova- or sperm-producing,externae (Crustacea,Cirripedia)

Summary Specimens of the blue swimming crab,Portunus pelagicus, are often infected with many thousands of externae ofThompsonia dofleini, all of which are connected through a common root system within the host crab. The species is unique in that the production of sperm cells takes place within the visceral mass of a small minority of the population of the externae. Spermatogonia are probably introduced by male cyprids into these externae when they are young, and they multiply and develop at the expense of the oocytes which rapidly disintegrate and ultimately disappear. It is assumed that the sperm cells are transferred to the ovary of the ordinary, egg-producing externae through the root system. Shortly after the eggs have been fertilized within the ovary they are transferred to the mantle cavity where they develop into cyprid larvae. The larvae become liberated when the externae drop off and the mantle wall disintegrates.Abbreviations used in the figures a annulus - an antennule - bm basal membrane around ovary - ce compound eye - cg cerebral ganglion - cr connecting root - cy cyprid larva(e) - c1 cuticle 1 - c2 cuticle 2 - do degenerating oocytes - e eggs in early division - ec embryonic cells - em embryos - en endocuticle of host - ep epidermis of host - ex exocuticle of host - hc hemocoelic cavity - m mantle - mc mantle cavity - mcy male cyprid - o ovary - oo oocyte - r rupture in wall of visceral mass - rs root system - sc scar - se spawned eggs - so spent ovary - sp spermatogonia - ss sperm and spermatids - st stalk - tc thickened cuticle of mantle cavity - th thorax - vm visceral mass - y yolk granules     

Anatomy of virgin and mature externae of Loxothylacus texanus,parasitic on the dark blue crab Callinectes rathbunae (Crustacea: Cirripedia: Rhizocephala: Sacculinidae)

Rhizocephalan parasites are dioecious organisms, in that one or several dwarf males are implanted into the external part of the female parasite soon after it emerges from the interior of the host animal. The structure of the female externa and its resident males is crucial for understanding both the reproductive biology and the taxonomy of these specialized parasites. We use scanning electron microscopy and histological methods to study the anatomy of juvenile and the mature externae of the rhizocephalan barnacle Loxothylacus texanus parasitizing the blue crab Callinectes rathbunae. We put emphasis on the implantation of males and the histology of the female reproductive organs. In the virgin externae, male cyprids attach around a cuticular hood covering the mantle aperture, which is partially blocked by a plug of cuticle so only trichogon larvae, not cyprids, can access the mantle cavity. This resembles the situation known from Sacculina carcini. The mature externa is characterized by a visceral mass that contains the ovary, paired colleteric glands, a single male receptacle, but paired receptacle ducts. The proximal attachment of the visceral mass is located at some distance from the basal stalk, as is characteristic for the genus Loxothylacus. The internal anatomy of the mature externa of L. texanus is in most features similar to that seen in other species of the Sacculinidae, which comprises the majority of rhizocephalan species. However, the single receptacle creates a situation where the two implanted males cannot be kept separate as in most other rhizocephalans, but pass through spermatogenesis in a common chamber. This may have unknown effects on the reproductive biology such as male–male competition. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Reproductive Stage of the Life Cycle in the Rhizocephalan Barnacle Polyascus polygenea (Crustacea: Cirripedia)

The specific features of the reproductive stage of the life cycle have been studied in the rhizocephalan barnacle Polyascus polygenea, a parasite of the coastal crab Hemigrapsus sanguineus. It is shown that a single crab can bear 1 to 8 externae of P. polygenea. The fecundity of the parasite depends on the size of the externae and their number on the host and may reach as much as 50000 eggs for one externa. In Peter the Great Bay, Sea of Japan, this species repeatedly reproduces during the entire spring–autumn period; externae with developing embryos in the mantle cavity occur from May to September, and planktonic larvae occur from June to October. One externa produces during the season of reproduction no less than three generations of larvae. Thus, the reproductive strategy in P. polygenea comprises a three-stage cascade of reproduction: asexual reproduction via budding of the interna; the development of several generations of one or several externae; and several reproduction cycles of each externa. This allows the parasite to produce a very great number of larvae and ensures the parasitization of a significant proportion of the host crab population. The structure of the ovaries and oogenesis in rhizocephalans and free-living cirripede barnacles have many common features, which provides evidence for integration of these two groups within one monophyletic taxon.     

Comparative morphology and phylogeny of the family Thompsoniidae (Cirripedia, Rhizocephala, Akentrogonida), with descriptions of three new genera and seven new species

Akentrogonid rhizocephalans morphologically resembling the genus Thompsonia are revised as a result of examination of new material. The species concerned are all obligatorily colonial and have ovoid or cylindrically shaped externae with a terminal stalk and a much reduced anatomy. A numerical cladistic analysis of all Rhizocephala Akentrogonida using the Hennig 86 program leads to a redefinition of the Thompsoniidae HOeg and Rybakov, 1992. Autapomorphies for the Thompsoniidae are primarily the morphology of the attachment to the host and the total absence of a mesentery. The cladistic analysis refutes that the Thompsoniidae should have a plesiomorphic morphology and branch off very low on the rhizocephalan phylogeny. The family now comprises four genera: Pottsia gen. n. (monotypic), Diplothylacus gen. n. with two species, Thompsonia Kossmann with five species. A revived and redefined Thylacoplethus Coutière includes eight species. The genera are distinguished by the location of the spermatogenic tissue, the site where the eggs are fertilized, the presence or absence of a mantle pore and the way it is formed, the number or absence of oviducts, and the number of cuticular annuli on the stalk. All 16 species, of which six are new to science, are described when necessary, and, if possible, illustrated. A phylogeny for the redefined family is proposed. Thylacoplethus is morphologically closest to the hypothetically ancestral thompsoniid and is likely paraphyletic. The new genus Polysaccus with two species, one of them new to science, and the monotypic genus Pirusaccus Lützen resemble thompsoniids in externa morphology and in being obligatorily colonial.     

The anatomy and development of the rhizocephalan barnacle Clistosaccus paguri Lilljeborg and relation to its host pagurus bernhardus (L.)

The anatomy of all developmental stages of Clistosaccus paguri Lilljeborg including internal primordia only 200 μm across has been studied by conventional and ultrastructural techniques. The earliest primordia lack any roots and organs. A receptacle and ovary arise as separate structures. The colleteric gland develops by infolding of the mantle cavity epithelium. The primordium becomes external during a host intermoult. The externa is apparently female. It is proposed that male cyprids, through the integument, have implanted cells found in the mantle of the externa. These male cells travel through the mantle into the single receptacle, where they proceed with spermatogenesis. Externae lacking male cells will not grow. The larvae are released as cyprids, in which no preformed kentrogon is present. About half of the infected hosts carry more than one externa. Multiple externae are separate and may represent different invading female cyprids. C. paguri infects all sizes of its host, and castrates it. The effect of the parasite on the host is discussed. The life span of the parasite seems to be one year and the parasite population shows an annual cycle.     

Distribution,prevalence and mean numbers of larvalAnisakis simplex (Nematoda: Ascaridoidea) in Atlantic salmon,Salmo salar L. and their use as biological indicators of host stocks

Synopsis An estimate of the arithmetic mean number of larvalAnisakis simplex per salmon,Salmo salar, in the north Atlantic was obtained from data collected off west Greenland where salmon from both sides of the Atlantic intermingle. An average of 5.34 larvae was found in the body cavity or associated viscera of 771 age 1 salmon representing some 60.8% of the total number of larvae present. LarvalA. simplex were found at all sampling stations throughout the north Atlantic. In most samples prevalence varied only slightly from area to area.The mean numbers of larvae in the body cavity and on associated organs of salmon in samples from North America were significantly different between samples. Comparisons of the mean numbers of larvae with several known variables in the data were, therefore, considered: host's sex, host's sea-age, geographic locality, year and season of capture.There was evidence that the mean number of larvae per host was greater in salmon from Europe than in those from North America, but the data could not be used to providereliable estimates of the mean numbers of larvae from each area because of heterogeneity within areas. Thus the data from west Greenland and from continental home waters could net be used in calculations to determine the percentage composition of North American and European salmon in the Greenland fishery although they supported earlier findings of mixing of salmon stocks off west Greenland.The use of parasitological data in the identification of different Atlantic salmon stocks is discussed.     

Postembryonic development of pycnogonids: A deeper look inside

Sea spiders form a small, enigmatic group of recent chelicerates, with an unusual bodyplan, oligosegmented larvae and a postembryonic development that is punctuated by many moults. To date, only a few papers examined the anatomical and ultrastructural modifications of the larvae and various instars. Here we traced both internal and external events of the whole postembryonic development in Nymphon brevirostre HODGE 1863 using histology, SEM, TEM and confocal microscopy. During postembryonic development, larvae of this species undergo massive reorganization: spinning apparatus and chelar glands disappear; larval legs redifferentiate; three new segments and the abdomen are formed with their corresponding internal organs and appendages; circulatory and reproductive systems develop anew and the digestive and the nervous systems change dramatically. The body cavity remains schizocoelic throughout development, and no traces of even transitory coeloms were found in any instar. In Nymphon brevirostre, just like in Artemia salina LINNAEUS 1758 the heart arises through differentiation of the already existing schizocoel, and thus the circulatory systems of arthropods and annelids are not homologous. We found that classical chelicerate tagmata, prosoma and opisthosoma, are inapplicable to adult pycnogonids, with the most striking difference being the fate and structure of the seventh appendage-bearing segment.     

Regional differences in the apical F-actin cytoskeleton of silk gland cells of Calpodes ethlius (Stöll) (Lepidoptera : Hesperiidae)

Silk glands of the larvae of Calpodes ethlius, Stöll (Lepidoptera : Hesperiidae) were labelled with rhodaminyl-phalloin, which is specific for f-actin. Most f-actin lies below the apical cell surface in bundles oriented in rings around the lumen of the gland. By the fifth larval stadium, the density of bundles differs in the 5 regions of the gland, the greatest density occurring in the narrower anterior regions. The greater bundle density may be correlated with the increased hoop stress where the gland has the smallest diameter. F-actin from the bundles redistributes during larval moults to coat vacuoles in the cytoplasm, except in the 2 most anterior regions where it keeps its arrangement from stage to stage, even through the moult.     

The monophyletic origin of a remarkable sexual system in akentrogonid rhizocephalan parasites: A molecular and larval structural study

We use sequences from the nuclear ribosomal genes, 18S and 28S to analyze the phylogeny of the Rhizocephala Akentrogonida including two species, Clistosaccus paguri and Chthamalophilus delagei, that are critical for understanding rhizocephalan evolution but have not previously been part of a molecularly based study. In addition we use light and scanning electron microscopy to compare the cypris larvae of C. paguri, Sylon hippolytes and two species of the family Thompsoniidae, since this larval stage offers a suite of characters for analyzing the evolution of these otherwise highly reduced parasites. The Rhizocephala Akentrogonida form a monophyletic group nested within a paraphyletic “Kentrogonida”. C. paguri and S. hippolytes are sistergroups confirming the monophyly of the Clistosaccidae that was originally based on similarities in the cypris larvae. We find numerous LM and SEM level similarities between the two species, many of which appear to be correlated with their specialized sexual system, where male cyprids use an antennule to implant cells into the virgin female parasite. Some of these traits are also found in cyprids of the thompsoniid species. We conclude that the special cypris morphology and the implantation of males by antennular penetration was present in the stem species to the Thompsoniidae and the Clistosaccidae and emphasize the power of larval characters in rhizocephalan systematics. C. delagei is a sister group to Boschmaella balani and the two are nested deep within the Akentrogonida. This confirms the monophyly of the Chthamalophilidae and falsifies the theory that C. delagei should represent the most primitive extant rhizocephalan. Instead, chthamalophilid rhizocephalans represent some of the most highly advanced members of the parasitic barnacles.     

Xenochironomus canterburyensis (Diptera: Chironomidae), a commensal of Hyridella menziesi (Lamellibranchia) in Lake Taupo; features of pre-adult life history

Abstract

The larvae and pupae of the univoltine chironomid Xenochironomus canterburyensis (Freeman) are inquiline commensals of the freshwater mussel Hyridella menziesi (Gray). The 1st- and 2nd-instar larvae enter the mantle/valve cavity of the mussel in midsummer, and by early winter migrate as 3rd-instar larvae to the posterior end of the valve to lodge near its margin beside the inhalant siphon. During the spring, growth of the periostracum of the valve margin between the larva and the mantle of the mussel leaves the 4th-instar larva outside the mantle/valve cavity, where it pupates before leaving the mussel for the lake surface and adult emergence.     

Siphonariid development: Quintessential euthyneuran larva with a mantle fold innovation (Gastropoda; Panpulmonata)

Recent phylogenetic revisions of euthyneuran gastropods (“opisthobranchs” and “pulmonates”) suggest that clades with a planktotrophic larva, the ancestral life history for euthyneurans, are more widely distributed along the trunk of the euthyneuran tree than previously realized. There is some indication that the planktotrophic larva of euthyneurans has distinctive features, but information to date has come mainly from traditional “opisthobranch” groups. Much less is known about planktotrophic “pulmonate” larvae. If planktotrophic larvae of “pulmonates” share unique traits with those of “opisthobranchs,” then a distinctive euthyneuran larval-type has been the developmental starting template for a spectacular amount of evolved morphological and ecological disparity among adult euthyneurans. We studied development of a siphonariid by preparing sections of larval and postmetamorphic stages for histological and ultrastructural analysis, together with 3D reconstructions and data from immunolabeling of the larval apical sensory organ. We also sought a developmental explanation for the unusual arrangement of shell-attached, dorso-ventral muscles relative to the mantle cavity of adult siphonariids. Adult siphonariids (“false limpets”) have a patelliform shell but their C-shaped shell muscle partially embraces a central mantle cavity, which is different from the arrangement of these components in patellogastropods (“true limpets”). It is not obvious how shell muscles extending into the foot become placed anterior to the mantle cavity during siphonariid development from a veliger larva. We found that planktotrophic larvae of Siphonaria denticulata are extremely similar to previously described, planktotrophic “opisthobranch” larvae. To emphasize this point, we update a list of distinctive characteristics of planktotrophic euthyneuran larvae, which can anchor future studies on the impressive evolvability of this larval-type. We also describe how premetamorphic and postmetamorphic morphogenesis of larval mantle fold tissue creates the unusual arrangement of shell-muscles and mantle cavity in siphonariids. This result adds to the known postmetamorphic evolutionary innovations involving mantle fold tissue among euthyneurans.     

Larvae of Chironomids (Insecta,Diptera) Encountered in the Mantle Cavity of Zebra Mussels,Dreissena polymorpha (Bivalvia,Dreissenidae)

The paper includes data on species composition of chironomid larvae which were encountered in the mantle cavity of zebra mussels (Dreissena polymorpha) within 7 waterbodies in the Republic of Belarus. All were found to be free‐living species commonly present in periphyton and/or benthos. A long‐term study of the seasonal dynamics of these larvae in Dreissena did not reveal any typical pattern. Our data suppose that chironomids do not have an obligate association with zebra mussels and possibly enter their mantle cavity inadvertently. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Identification of the hatching factor of the barnacle as the novel eicosanoid 10,11,12-trihydroxy-5,8,14,17-eicosatetraenoic acid

Barnacle egg hatching fator which is released into the mantle cavity where the egg masses are brooded and stimulates embryonic musculature resulting in hatching and liberation of the larvae into the sea has been isolated in a purified form from a common barnacle . Derivatised hatching factor has been analysed by GC-MS and identified as 10,11,12-trihydroxy-5,8,14,12-eicosatetraenoic acid, a novel eicosanoid probably formed from endogenous eicosapentaenoic acid (20:5w3). Hatching factor activity is the first specific physiological function to be established for this type of compound.     

The effects of cholesterol on the development of Heliothis zea

Heliothis zea was reared on an artificial diet, which lacked supplementation with plant materials, in order to determine the effects of cholesterol on the development of this insect. A number of parameters of larval development were found to be dependent upon the concentration of dietary sterol including: the number of moults which the larvae completed within a particular time interval, the ability of the larvae to pupate and the survival of the larvae. The number of moults which a larva completed prior to pupation, though, was independent of the concentration of sterol.     

Daily changes in the sensitivity of wax-moth larvae, Galleria mellonella, to cooling stress

ABSTRACT. Sensitivity to cooling stress in the last instar larvae of Galleria mellonella was measured as (a) the number of extra larval moults, (b) the number of larvae retaining the ability to secrete silk, and (c) the number showing arrested development. With respect to (a) and (b) there were considerable differences in sensitivity across the day. A relationship was observed between the number of additional larval moults induced by chilling and the ability of prepupal larvae to spin silk: the periods during the 24 h when the most larvae passed through additional larval moults were periods characterized by the smallest number of larvae capable of spinning, and vice versa. These daily changes were apparently partly independent of developmental age. Daily variations in sensitivity also occurred when larvae of the same age were cooled at different times of day. It is suggested that these rhythms in cold-sensitivity are related to a cold-sensitive rhythm in juvenile hormone secretion, or hormone sensitivity in the tissues.     

Size and settling behaviour in male and female cypris larvae of the parasitic barnacle Sacculina carcini Thompson (Crustacea: Cirripedia: Rhizocephala)

Twenty-one broods from different externae of Sacculina carcini Thompson were cultured to the cypris stage. The size of the cyprids was measured and the larvae subjected to settling upon unparasitized crabs (Garcinus maenas (L.)) and small juvenile externae. The cyprids occur in two sizes that may appear singly or together in the same brood. Small cyprids are of the female sex that settle upon crabs and are infective, while large cyprids are of the male sex and only settle upon juvenile externae. These results are in agreement with other well-studied rhizocephalans.     

Towards a mechanistic understanding of insect life history evolution: oxygen‐dependent induction of moulting explains moulting sizes

Moults characterise insect growth trajectories, typically following a consistent pattern known as Dyar's rule; proportional size increments remain constant across inter‐instar moults. Empirical work suggests that oxygen limitation triggers moulting. The insect respiratory system, and its oxygen supply capacity, grows primarily at moults. It is hypothesized that the oxygen demand increases with increasing body mass, eventually meeting the oxygen supply capacity at an instar‐specific critical mass where moulting is triggered. Deriving from this hypothesis, we develop a novel mathematical model for moulting and growth in insect larvae. Our mechanistic model has great success in predicting moulting sizes in four butterfly species, indirectly supporting a size‐dependent mechanism underlying moulting. The results demonstrate that an oxygen‐dependent induction of moulting mechanism would be sufficient to explain moulting sizes in the study species. Model predictions deviated slightly from Dyar's rule, the deviations being typically negligible within the present data range. The developmental decisions (e.g. moulting) made by growing larvae significantly affect age and size at maturity, which has important life history implications. The successful modelling of moulting presented here provides a novel framework for the development of realistic insect growth models, which are required for a better understanding of life history evolution.