Tantulocarida versus Thecostraca: inside or outside? First attempts to resolve phylogenetic position of Tantulocarida using gene sequences

Complete 18S rDNA sequences of two species of the Tantulocarida Arcticotantulus pertzovi (Basipodellidae) and Microdajus tchesunovi (Microdajidae) were obtained and used for estimating the relationship of the class with other Crustacea. This constitutes the first use of tantulocaridan gene sequences, and we conclude that the Tantulocarida are very close relatives of the class Thecostraca, which comprise cirripedes, ascothoracidans and the enigmatic facetotectans. With much lower confidence, the Tantulocarida are also indicated as nested within the Thecostraca, being sister group to the Cirripedia. We therefore discuss morphological similarities and differences between tantulocaridans and the thecostracans in search of potential synapomorphies, including a possible relation to the parasitic barnacles (Rhizocephala). We conclude that the cement gland of the tantulus larva and the cirripede cyprid might be homologous structures, but that similarities in host infection and root systems between the Tantulocarida and the Rhizocephala are, on present evidence, likely to be homoplasies evolved by convergent evolution into advanced parasitism. The precise position of the Tantulocarida in relation to or within the Thecostraca must be pursued by a more extensive database of genetic markers.     

Serratotantulus chertoprudae gen. et sp. n. (Crustacea, Tantulocarida, Basipodellidae): A new tantulocaridan from the abyssal depths of the Indian Ocean

A single tantulus larva was found at the abyssal depth of the Indian Ocean attached to a harpacticoid host of the family Cletodidae. It represents a new genus and species of Tantulocarida, family Basipodellidae. Its ultrastructure was studied with SEM. This genus can be easily distinguished from the other genera of Basipodellidae by the pore pattern, bilobed oral disk with strong longitudinal ridges and the posterior projection of the cephalic shield. A morphological analysis of two related families Basipodellidae and Deothertridae shows that they represent polyphyletic taxa and need further revision.     

Microdajus tchesunovi sp. n. (Tantulocarida, Microdajidae) - A new crustacean parasite of from the White Sea

A new species of Tantulocarida, Microdajus tchesunovi sp. n., was found on tanaid host, Typhlotanais sp. of the family Nototanaidae, collected from silty sediment in the Kandalaksha Bay in the White Sea. Several tantulus larvae, developing males at different stages of metamorphosis, and early stages of parthenogenetic females were found attached to different sites of their hosts. Ultrastructure of the new species was studied with SEM. Microdajus tchesunovi sp. n. can be easily distinguished from other species of the genus Microdajus Greve by the presence of a pair of longitudinal dorsal lamellae at the anterior end of the cephalon. It is also characterized by the presence of an endopod seta on the sixth thoracopod and the absence of thoracopodal endites. A morphological comparison of species of the family Microdajidae is presented in tabular form.     

A new species ofMicrodajus (Crustacea: Tantulocarida) parasitic on a tanaid in the northeastern Atlantic,with observations onM. langi Greve

The tantulus larva ofMicrodajus langi is redescribed using Scanning Electron Microscopy. Thoracopods 2–5 have poorly developed but fully armed endites. Thoracopods 1–5 are biramous, with each ramus represented by a minute segment. A new species,M. pectinatus, is described from a tanaid host collected in the Rockall Trough, off the west coast of Scotland. A system of homologies for the portopodal segments of the thoracopods of the tantulus larva is proposed. Some aspects of the musculature of the adult male are described for the first time.     

Cuticular morphogenesis during continuous growth of the final instar larva of a moth

The larvae of the tobacco hornworm, Manduca sexta, grow continuously. During the feeding period of the fifth larval instar their weight increases ten-fold (ca. 1·2–12 g) accompanied by a four-fold expansion of the surface area of the abdominal cuticle. We have found that this cuticle contains structures which facilitate its expansion. Folds in the epicuticle (papillae) flatten as the cuticle expands. The endocuticle, in contrast, does not unfold but rather is plastically deformed. This plastic deformation is assisted by vertical structures in the cuticle (cuticular columns) which are more easily deformed than the surrounding lamellate cuticle. The head capsule cuticle, which does not expand as the larva grows, lacks papillae and cuticular columns. Thus, these are specialized structures that are reserved for cuticle that must expand as the larva grows.     

Fine structure and probable function of ring organs in the mite Histiostoma feroniarum (Acari: Actinotrichida: Acaridida: Histiostomatidae)

Histiostoma feroniarum, like other histiostomatid mites, possesses peculiar ring organs that are visible under the light microscope as ventrally located, characteristic rings of sclerotized cuticle. The ring organ is composed of three elements: a disc of modified cuticle, ring organ cells located underneath the disc, and an "empty" chamber frequently visible between the cuticular disc and the cells. The cuticle of the disc is not perforated and differs from the surrounding unmodified cuticle as revealed by special staining developed for light microscopy and by electron microscopy. The ring organ cells show a polarity, with a practically smooth apical surface and an extremely folded basal membrane. The basal invaginations reach the apical cell portion, where they form tubular canaliculi distributed beneath the apical cell membrane. The cytoplasm contains many mitochondria, which are usually in contact with the cell membrane invaginations. Structurally, the ring organ cells closely resemble the transport cells described in osmoregulatory organs both in water-inhabiting and terrestrial arthropods. Thus, our results support earlier suggestions of an osmoregulatory function performed by sclerotized rings (=ring organs), as an adaptation to aqueous environments. A possible homology with similar organs of other mites is discussed.     

A redescription of Cumoniscus Bonnier, 1903 (Crustacea: Tantulocarida) from a cumacean host in the Bay of Biscay

Cumoniscus kruppi Bonnier, the only tantulocarid known from a cumacean host, is redescribed from a tantulus larva collected from the Bay of Biscay. The host is a male leuconid cumacean. This genus is placed in the family Deoterthridae, and can be distinguished from other genera by the pore pattern on the cephalic shield, the reduced setae on the sixth legs and the presence of tiny curved spinules on the endopod of thoracopods 2 to 5.     

Exoskeletal cuticle differentiation during intramarsupial development of Porcellio scaber (Crustacea: Isopoda)

Exoskeletal crustacean cuticle is a calcified apical extracellular matrix of epidermal cells, illustrating the chitin-based organic scaffold for biomineralization. Studies of cuticle formation during molting reveal significant dynamics and complexity of the assembly processes, while cuticle formation during embryogenesis is poorly investigated. This study reveals in the terrestrial isopod Porcellio scaber, the ultrastructural organization of the differentiating precuticular matrices and exoskeletal cuticles during embryonic and larval intramarsupial development. The composition of the epidermal matrices was obtained by WGA lectin labelling and EDXS analysis. At least two precuticular matrices, consisting of loosely arranged material with overlying electron dense lamina, are secreted by the epidermis in the mid-stage embryo. The prehatching embryo is the earliest developmental stage with a cuticular matrix consisting of an epicuticle and a procuticle, displaying WGA binding and forming cuticular scales. In newly hatched marsupial larva manca, a new cuticle is formed and calcium sequestration in the cuticle is evident. Progression of larval development leads to the cuticle thickening, structural differentiation of cuticular layers and prominent cuticle calcification. Morphological characteristics of exoskeleton renewal in marsupial manca are described. Elaborated cuticle in marsupial larvae indicates the importance of the exoskeleton in protection and support of the larval body in the marsupium and during the release of larvae in the external environment.     

The attachment complex of brachiolaria larvae of the sea star <Emphasis Type="Italic">Asterias rubens</Emphasis> (Echinodermata): an ultrastructural and immunocytochemical study

The attachment complex of brachiolaria larvae of the asteroid Asterias rubens comprises three brachiolar arms and an adhesive disc located on the preoral lobe. The former are used in temporary attachment and sensory testing of the substratum, whereas the latter is used for permanent fixation to the substratum at the onset of metamorphosis. Brachiolar arms are hollow structures consisting of an extensible stem tipped by a crown of dome-like ciliated papillae. The papilla epidermis is composed of secretory cells (type A, B and C cells), non-secretory ciliated cells, neurosecretory-like cells and support cells. Type A and B secretory cells fill a large part of the papilla epidermis and are always closely associated. They presumably form a duo-gland adhesive system in which type A and B cells are respectively adhesive and de-adhesive in function. The adhesive disc is an epidermal structure mainly composed of secretory cells and support cells. Secretory cells produce the cement, which anchor the metamorphic larva to the substratum until the podia are developed. The relatedness between the composition of the adhesive material in the brachiolaria attachment complex and in the podia of adults was investigated by immunocytochemistry using antibodies raised against podial adhesive secretions of A. rubens. Type A secretory cells were the only immunolabelled cells indicating that their temporary adhesive shares common epitopes with the one of podia. The attachment pattern displayed by the individuals of A. rubens during the perimetamorphic period—temporary, permanent, temporary—is unique among marine non-vertebrate Metazoa.     

Studies on the host parasite relationship between the parasitic prosobranch Thyca crystallina and the asteroid starfish Linckia laevigata

A study of the Blue starfish Linckia laevigata in the Banda Islands, Moluccas, Indonesia, revealed 62% infestation by the molluscan prosobranch parasite Thyca crystallina. Infestation rate varies directly with the degree of water movement. A marked difference in settlement pattern according to parasite size is observed. The smallest limpets tend to settle on the upper surface of the distal parts of the arms and are randomly oriented. The largest parasites are found on the oral arm surface on the right hand side of the ambulacral groove and facing the starfish mouth. All of the attached parasites were female. Most of the large females had a dwarf male, one tenth the length, attached, under the mantle, at the anterior end. Amongst the small limpets, less than half had made a proboscis hole in the host integument and none had made an attachment disc lesion; of the large parasites 90% had the proboscis embedded in the host and 80% left an attachment disc lesion on the host. Small parasites can move very slowly over the host surface. Larger limpets are probably permanently attached. A detached parasite of any size is probably unable to reattach. There is evidence of an extended parasite breeding period and repetitive settlement on the hosts, which could cause the observed overdispersion without invoking a mature female parasite pheromone.     

Cuticular cavities: storage chambers for cyanoglucoside-containing defensive secretions in larvae of a Zygaenid moth

As a defensive reaction against predators the larvae of Zygaena trifolii Esper. 1783, release highly viscous fluid droplets out of cuticular cavities. The fluid appears on the cuticular surface upon contraction of the irritated segments, with no specialized muscles being involved. Two morphologically different types of cavities have been found: the larger ones are located beneath pigment spots, the smaller ones occupy the remaining surface except in the ventral region. Both types have complicated cuticular opening structures. The defensive fluid contains the cyanoglucosides linamarin and lotaustralin, the amino acid beta-cyano-L-alanine, proteins and water. Although a considerable amount of fluid (3-6 microliter per sixth-instar larva) is stored in the cuticle, fine structural examinations of the epidermis do not show any specific cells or cell areas with morphological adaptations for secretion. Further, there do not exist any major cytological differences between the cells below the cavities and in the ventral region, where those cavities are absent.     

Disentangling the assembly mechanisms of ant cuticular bacterial communities of two Amazonian ant species sharing a common arboreal nest

Bacteria living on the cuticle of ants are generally studied for their protective role against pathogens, especially in the clade of fungus‐growing ants. However, little is known regarding the diversity of cuticular bacteria in other ant host species, as well as the mechanisms leading to the composition of these communities. Here, we used 16S rRNA gene amplicon sequencing to study the influence of host species, species interactions and the pool of bacteria from the environment on the assembly of cuticular bacterial communities on two phylogenetically distant Amazonian ant species that frequently nest together inside the roots system of epiphytic plants, Camponotus femoratus and Crematogaster levior. Our results show that (a) the vast majority of the bacterial community on the cuticle is shared with the nest, suggesting that most bacteria on the cuticle are acquired through environmental acquisition, (b) 5.2% and 2.0% of operational taxonomic units (OTUs) are respectively specific to Ca. femoratus and Cr. levior, probably representing their respective core cuticular bacterial community, and (c) 3.6% of OTUs are shared between the two ant species. Additionally, mass spectrometry metabolomics analysis of metabolites on the cuticle of ants, which excludes the detection of cuticular hydrocarbons produced by the host, were conducted to evaluate correlations among bacterial OTUs and m/z ion mass. Although some positive and negative correlations are found, the cuticular chemical composition was weakly species‐specific, suggesting that cuticular bacterial communities are prominently environmentally acquired. Overall, our results suggest the environment is the dominant source of bacteria found on the cuticle of ants.     

Learning and discrimination of individual cuticular hydrocarbons by honeybees (Apis mellifera)

In social insect colonies, recognition of nestmates, kinship, caste and reproductive status is crucial both for individuals and for the colony. The recognition cues used are thought to be chemical, with the hydrocarbons found on the cuticle of insects often cited as being particularly important. However, in honeybees (Apis mellifera) the role of cuticular hydrocarbons in nestmate recognition is controversial. Here we use the proboscis extension response (PER) conditioning paradigm to determine how well honeybees learn long-chain linear alkanes and (Z)-alkenes present on the cuticle of worker bees, and also how well they can discriminate between them. We found large differences both in learning and discrimination abilities with the different cuticular hydrocarbons. Thus, the tested hydrocarbons could be classified into those which the bees learnt and discriminated well (mostly alkenes) and those which they did not (alkanes and some alkenes). These well-learnt alkenes may constitute important compounds used as cues in the social recognition processes.     

Factors influencing the developmental capacity of Galleria larval epidermis

The nature of the cuticle secreted by integument from a day-1 penultimate instar larval Galleria when cultured in vivo in the abdomen of a last instar larva varied with the age of the host. When placed in a day-5 last instar larva, the implanted integument secreted a pupal cuticle at the time the host metamorphosed and became a pupa. However, when placed in a day-7 last instar larva the implant, from the same stage donor, secreted a larval cuticle at the time the host pupated. Experimental studies involving implantation of the integument for a 24 hr period, into various developmental stages of normal and ligated last instar larvae, pupae, and pharate adults, prior to placing it in a day-7 last instar larva suggest that a non-hormonal factor present in day-4 and -5 last instar larvae is important to initiate pupal syntheses.     

The effect of host species on growth and variability of Echinorhynchus salmonis Müller, 1784 (Acanthocephala: Echinorhynchidae), with special reference to the status of the genus

Summary The sex and age (as measured by length) of Echinorhynchus salmonis Müller, 1784 and the host species of this acanthocephalan considerably affected worm body form and size, as well as size of proboscis, proboscis hooks, proboscis receptacle, lemnisci, testes and cement glands. Linear regression analysis indicated that curves describing the growth pattern of these characters by worm length were significantly different as a function of host species. The larger worms recovered from bloater (Coregonus hoyi: Salmonidae) almost invariably showed higher regression coefficient compared to those from smelt (Osmerus mordax: Osmeridae) in all characters. Taxonomic implications of these findings are discussed. Abnormalities in body wall, proboscis hook orientation, lemnisci and cement gland ducts as well as variations in proboscis hook and cement gland numbers are reported, some for the first time. Findings from studies of cement gland pattern invalidate Petrochenko's (1956) splitting up of the genus into three: Echinorhynchus, Metechinorhynchus and Pseudoechinorhynchus. It is proposed that the designation of the last two genera as junior synynoms of the first be accepted. ac]19791205     

Larval morphology of the Nemertean Carcinonemertes epialti (Nemertea: Hoplonemertea)

The morphology of the newly hatched larva of Carcinonemertes epialti Coe has been examined by light and electron microscopy. The newly hatched larva is covered with cilia and measures about 110 μm in length. Four types of epidermal cells are recognizable: (1) Multiciliated cells, (2) vacuolated cells, (3) mucous cells, and (4) “knob cells”. The knob cells protrude from the posterior end of the larva and contain granules and bundles of microfilaments. The gut is incomplete and is located ventral to the bipartite proboscis. A bilobed brain and two subepidermal ocelli are found in the anterior end of the larva. The anterior and posterior cirri are composed of long, tightly appressed cilia that arise from an invagination of the epidermis at each end of the larva. The anterior cirrus is surrounded by two types of glandular cells. It is proposed that the knob cells have a role in larval attachment, combining the functions of the adhesive cells and anchor cells described in the duo-gland system of turbellarians. The cirri are believed to be larval sensory structures that function in substrate selection. Histological and ultrastructural observations suggest that the larvae of Carcinonemertes are relatively long lived and develop into juveniles without a drastic metamorphosis.     

Comparative Proteomics Studies of Insect Cuticle by Tandem Mass Spectrometry: Application of a Novel Proteomics Approach to the Pea Aphid Cuticular Proteins

The cuticle is a biological composite material consisting principally of N‐acetylglucosamine polymer embedded in cuticular proteins (CPs). CPs have been studied and characterized by mass spectrometry in several cuticular structures and in many arthropods. Such analyses were carried out by protein extraction using SDS followed by electrophoresis, allowing detection and identification of numerous CPs. To build a repertoire of cuticular structures from Bombyx mori, Apis mellifera and Anopheles gambiae the use of SDS and electrophoresis was avoided. Using the combination of hexafluoroisopropanol and of a surfactant compatible with MS, a high number of CPs was identified in An. gambiae wings, legs and antennae, and in the thoracic integument cuticle of Ap. mellifera pupae. The exoskeleton analysis of B. mori larvae allowed to identify 85 CPs from a single larva. Finally, the novel proteomics approach was tested on cuticles left behind after the molt from the fourth instar of Acyrthosiphon pisum. Analysis of these cast cuticles allowed to identify 100 Ac. pisum CPs as authentic cuticle constituents. These correspond to 68% of the total putative CPs previously annotated for this pea aphid. While this paper analyzes only the recovered cuticular proteins, peptides from many other proteins were also detected.     

Investing in attachment: evolution of anchoring structures in acanthocephalan parasites

Secure attachment to host tissues is essential for survival and reproduction in parasitic organisms. The production of elaborate attachment structures must be costly, however, and investments in attachment should be approximately proportional to the likelihood that a parasite will be dislodged. In the present study, relative investments in attachment as a function of body size and the type of host used were examined across 138 species of acanthocephalans. These worms live anchored to the intestinal wall of a vertebrate host by inserting their hooked proboscis into host tissues. Taking proboscis volume into account, there is a negative interspecific relationship between the number of hooks borne on the proboscis and their mean length, reflecting a trade‐off between hook number and hook length. This supports the assumption that hooks are costly to produce, because any given species cannot simultaneously maximize both the relative number and relative length of the hooks it produces. There is a positive relationship between total worm size and total hook length, but it is weak, with a slope indicating that, as total body volume increases, total hook length also increases but at a slower rate. Indeed, relative investments in attachment, measured as hook length per unit body volume, decrease as worm size increases. Independently of total body size, investments in hook production are higher in species exploiting endothermic hosts, especially birds, than in those living in ectothermic hosts. Given the greater amounts of food passing through the gut of endotherms, and the richer and denser communities of intestinal parasites that they harbour, they are likely to select for greater investments in attachment. These results support the prediction that investments in attachment are influenced by the probability of being dislodged, and allow comparisons with other groups of intestinal parasites such as cestodes or trematodes. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 637–645.     

<Emphasis Type="Italic">Prodajus curviabdominalis</Emphasis> n. sp. (Isopoda: Epicaridea: Dajidae), an ectoparasite of mysids,with notes on morphological changes,behaviour and life-cycle

Adults and larvae of a new ectoparasitic isopod, Prodajus curviabdominalis n. sp., are described from the mysid Siriella okadai Ii collected from the Seto Inland Sea, western Japan. The adult female is found within the host marsupium with the cephalon directed posteriorly, whereas the dwarf adult male attaches to the ventral surface of the female pleon. The cryptoniscid larva usually attaches to the second or third abdominal somite of the host, using an oral sucker. Mature adults of the new species are distinguished from all other congeners by: pleon of ovigerous female strongly curved dorsally, with large swellings on ventral side; pereon of ovigerous female narrow; exopods on male uropods present; male pleon short and thick. This is the third record of a member of the Dajidae from Japan. The behaviour of the cryptoniscid larvae of the new species on the host mysid was also observed using a video camera. Larvae moved from the first attachment site, usually the second or third abdominal somite of the host mysid, into the marsupium. When host oostegites were not fully developed, larvae entered beneath the host carapace until her marsupium was fully formed. The host infected by a female P. curviabdominalis moved the oostegites rhythmically, an action which may aid the respiration of the parasite.