Larvae of Barnacles (Cirripedia: Thoracica) in the White Sea Plankton

The barnacle fauna of the White Sea is briefly described. The morphology of barnacle larvae in this water body is comparatively analyzed. The characters important for the larvae identification are given particular attention. A classification key for the naupliar and cyprid larvae of White Sea barnacles is proposed.     

Biodiversity and Biogeography of Chthamalid Barnacles from the North-Eastern Pacific (Crustacea Cirripedia)

The biogeography and ecology of the species of Chthamalus present on the west coast of America are described, using data from 51 localities from Alaska to Panama, together with their zonation on the shore with respect to that of other barnacles. The species present were C. dalli, Pilsbry 1916, C. fissus, Darwin, 1854, C. anisopoma Pilsbry 1916 and four species in the C. panamensis complex. The latter are C. panamensis Pilsbry, 1916, C. hedgecocki, Pitombo & Burton, 2007, C. alani nom. nov. (formerly C. southwardorum Pitombo & Burton, 2007) and C. newmani sp. nov.). These four species were initially separated by enzyme electrophoresis. They could only be partially separated by DNA bar coding but may be separated using morphological characters.     

Scanning Electron Microscopy of Mouth Appendages in Six Species of Barnacles (Crustacea Cirripedia Thoracica)

The morphology and setation of mouth appendages (trophi) are investigated with scanning electron microscopy in the balanomorph barnacles Semibalanus balanoides, Balanus balanusand B. nubilusand in the pedunculate barnacles Pollicipes polymerus, P. cornucopiaand Lepas anatifera.It is difficult to uphold a clearcut distinction between denticles and setae and several types of setae also intergrade with each other. The trophi of Pollicipes polymerusand P. cornucopiahave the most ‘generalized’ morphology. The palps of Pollicipesand L. anatiferaare simpler than those in balanomorphs and carry a single type of seta. In L. anatiferathe mandibles and maxillules are adapted to a semi-predatory feeding habit by carrying large, pointed teeth, but this species lacks the foliate-serrate setae which populate the palps in the other species studied. Compared with Pollicipesand Lepas, the three balanomorphs have palps with a complex setation. The trophi in S. balanoidesdiffer from the two species of Balanusin numerous features of setation and denticulation, notably in having the palps populated by plumodenticulate setae homologous to purely denticulated types in Balanus.It is suggested that SEM studies of barnacle trophi will provide characters of use in estimating phylogeny.     

Cladistic analysis of the Cirripedia Thoracica

We present a cladistic analysis of the Cirripedia Thoracica using morphological characters and the Acrothoracica and Ascothoracida as outgroups. The list of characters comprised 32 shell and soft body features. The operational taxonomic units (OTUs) comprised 26 well-studied fossil and extant taxa, principally genera, since uncertainty about monophyly exists for most higher ranking taxonomic units. Parsimony analyses using PAUP 3.1.1 and Hennig86 produced 189 trees of assured minimal length. We also examined character evolution in the consensus trees using MacClade and Clados. The monophyly of the Balanomorpha and the Verrucomorpha sensu stricto is confirmed, and all trees featured a sister group relationship between the ‘living fossil Neoverruca and me Brachylepadomorpha. In the consensus trees the sequential progression of ‘pedunculate‘sister groups up to a node containing Neolepas also conforms to current views, but certain well-established taxa based solely on plesiomorphies stand out as paraphyletic, such as Pedunculata (= Lepadomorpha); Eolepadinae, Scalpellomorpha and Chthamaloidea. The 189 trees differed principally in the position of shell-less pedunculates, Neoverruca, the scalpelloid Capitulum, and the interrelationships within the Balanomorpha, although the 50% majority rule consensus tree almost fully resolved the latter. A monophyletic Sessilia comprising both Verrucomorpha and Balanomorpha appeared among the shortest trees, but not in the consensus. A tree with a monophyletic Verrucomorpha including Neoverruca had a tree length two steps longer than the consensus trees. Deletion of all extinct OTUs produced a radically different tree, which highlights the importance of fossils in estimating cirripede phylogeny. Mapping of our character set onto a manually constructed cladogram reflecting die most recent scenario of cirripede evolution resulted in a tree length five steps longer than any of our shortest trees. Our analysis reveals that several key questions in cirripede phylogeny remain unsolved, notably the position of shell-less forms and the transition from ‘pedunculate‘to ‘sessile‘barnacles. The inclusion of more fossil species at this point in our understanding of cirripede phylogeny will only result in even greater levels of uncertainty. When constructing the character list we also identified numerous uncertainties in the homology of traits commonly used in discussing cirripede evolution. Our study highlights larval ultrastructure, detailed studies of early ontogeny, and molecular data as the most promising areas for future research.     

The ectoparasitic barnacle Anelasma (Cirripedia,Thoracica, Lepadomorpha) on the shark Centroscyllium nigrum (Chondrichthyes,Squalidae) from the Pacific sub-Antarctic

We report the occurrence of the ectoparasitic lepadomorph barnacle Anelasma sp. on the deep-sea squaloid shark Centroscyllium nigrum from the Pacific sub-Antarctic off southern Chile. Anelasma has previously been documented only from the northeast Atlantic on the squaloid shark Etmopterus spinax; this new record extends the known range of Anelasma into the Pacific Ocean and into the Southern Hemisphere, and documents a new host for this parasitic barnacle.     

The functional morphology of the alimentary tract of barnacles (Cirripedia: Thoracica)

The alimentary tract of barnacles is made up of cuticle-lined foregut and hindgut with an intervening U-shaped midgut associated anteriorly with a pair of pancreatic glands and perhaps midgut caeca. Epithelial salivary glands secrete acid mucopolysaccharide, glycoprotein or both. Cells of all the midgut regions are capable of absorption which is carried out mainly by the anterior midgut and caeca. Midgut cells of Balanus balanoides (L.) show a seasonal variation in the distribution of intracellular lipid droplets. Midgut cells rest on an elastic basal lamina and secrete a peritrophic membrane which contains mucopolysaccharide and protein. Cells of the stratum perintestinale connect with the midgut epithelial cells via cell processes which probably translocate absorbed materials. Glycoprotein globules and lipid droplets accumulate in the body parenchyma of B. balanoides and are transported to the ovaries to form yolk (glycolipovitellin). The pancreatic gland cells of all barnacles are active secretory cells secreting proteinaceous material (probably digestive enzymes).     

Molecular phylogeny and character evolution of the chthamaloid barnacles (Cirripedia: Thoracica)

The Chthamaloidea (Balanomorpha) present the most plesiomorphic characters in shell plates and cirri, mouthparts, and oral cone within the acorn barnacles (Thoracica: Sessilia). Due to their importance in understanding both the origin and diversification of the Balanomorpha, the evolution of the Chthamaloidea has been debated since Darwin's seminal monographs. Theories of morphological and ontogenetic evolution suggest that the group could have evolved multiple times from pedunculated relatives and that shell plate number diminished gradually (8→6→4) from an ancestral state with eight wall plates surrounded by whorls of small imbricating plates; but this hypothesis has never been subjected to a rigorous phylogenetic test. Here we used multilocus sequence data and extensive taxon sampling to build a comprehensive phylogeny of the Chthamaloidea as a basis for understanding their morphological evolution. Our maximum likelihood and Bayesian analyses separate the Catophragmidae (eight shell plates and imbricating plates) from the Chthamalidae (8-4 shell plates and no imbricating plates), but do no support a gradual reduction in shell plates (8→6→4). This suggests that evolution at the base of the Balanomorpha involved a considerable amount of homoplasy.     

Littoral barnacles of The Snares islands,southern New Zealand (Cirripedia: Thoracica)

Abstract

Ibla idiotica, Lepas australis, Calantica villosa, Chamaesipho columna, Tetraclitella purpurascens, Notobalanus vestitus, and Notomegabalanus campbelli are reported for the first time from The Snares. The occurrence of Calantica spinosa and Epopella plicata is confirmed. Habitat information and recorded distribution in New Zealand are given for each species. The Snares barnacle fauna is restricted both in diversity and in numbers of individuals. This may be due to high wave energies, strong winds which aid desiccation, and counter-flow of currents, which might disrupt larval dispersal from the north and north-west.     

Aspects of the circulatory system of Pollicipes polymerus J. B. Sowerby (Cirripedia: Thoracica)

The circulatory system of Pollicipes polymerus exhibits a high degree of organization which precludes it from being referred to as an open system. The system is arbitrarily divided into four parts: (1) the circulation of the peduncle and mantle; (2) the distributive circulation of the body, which provides hemolymph to most of the cephalic gut, to the maxillary gland, and to the cirri; (3) the peripheral circulation which distributes blood from the cirri to the peripheral areas of the thoracic region, to most of the thoracic gut, and from the scutal sinus to the peripheral areas of the cephalic region; and (4) the collecting circulation, which conveys hemolymph mostly from the peripheral circulation of the body to the peduncle. There also may be a circulation that is comparable to the vertebrate lymphatic system. Pumping of hemolymph can be attributed to three pairs of skeletal muscles that compress the dorsolateral channels. These muscles are unique for crustacean muscles in that they do not appear to be striated. The rostral vessel appears to be a vestige of a heart in which the pump muscles have been lost. There is a similarity of the rostral vessel to the heart of Calanus finmarchicus (a copepod). This is additional evidence linking the cirripeds with the copepods within the Maxillopoda. Electron microscope observations of the walls of the midsagittal vessels indicate that there is a more or less random layering of cellular and noncellular elements within the wall. Muscle cells appear to be incorporated in the vessel wall.     

Seasonal species composition of barnacle larvae (Cirripedia: Thoracica) in Rhode Island waters, 1977-1978

Cirripede larvae can occur year-round in temperate and tropicalwaters, often in significant numbers, yet the species compositionof the ‘Balanus sp.’ component is rardy studied.Weekly plankton samples were analyzed qualitatively for larvalcirripede species and stage over a year (1977–1978) attwo Rhode Island stations. Six species of larvae were foundin Lower Narragansett Bay (30° salinity). Semibalanus balanoidesand Balanus balanus have a single winter brood. S. balanoidesis the predominant winter breeder with a minor release of naupliiin early December and major release in March followed by cypridsin mid-April.B. balanus populations release all larvae in Marchwith cyprids in mid-April. Balanus crenatus is mainly a winterbreeder, but has multiple broods; it does not breed in July-Septemberwhen the water temperature is above 18°C. Balanus venustusis the predominant summer breeder, and larvae were observedfrom May through December (water >8°C). Larvae of Chthamalusfragilis and Balanus eburneus occur in low numbers from May-October.At the Pettaquamscutt River site (12 salinity), Balanus improvisuslarvae predominate and early stage nauplii (I-II) occur in samplesyear round (0–27°C). Two peaks of later stage naupliiand cyprids occur in late spring (May) and early winter (Nov.-Jan.).Continued temperatures bdow 5°C or above 20°C appearto inhibit larval development. Comparison of results with existing literature reveals severalsignificant findings. The bimodal rdease of S. balanoides larvaeis unusual and may be in response to the phytoplankton dynamicsof the year; however, the existence of distinct races of S.balanoides may also be a factor. Larvae of B. venustus predominatein the lower bay during the summer, yet this species is unreportedin past studies. B. improvisus nauplii are more cold tolerantthan previously reported. Comparison of findings with reportedbreeding patterns in Florida indicate significant differencesin temperature responses between northern and southern populations. ¹Contribution No. 188 from EPA Environmental Research Laboratory,South Ferry Rd., Narragansett, RI 02882     

Taphonomy and systematics of a new Late Cretaceous verrucid barnacle (Cirripedia,Thoracica) from Canterbury,New Zealand

Abstract: Cirripede remains (Thoracica, Verrucomorpha), found associated with the mosasaur Prognathodon waiparaensis  Welles and Gregg, 1971 in glauconitic sands of the Late Cretaceous Conway Formation exposed along the Waipara River bank (mid‐Canterbury, New Zealand), are identified as a new species, Verruca sauria sp. nov. On the basis of taphonomy, it is deduced that these verrucids grew on a postmortem accumulation of mosasaur bones under very quiescent conditions. The current amphitropical distribution of the earliest known verrucids, i.e. V. sauria sp. nov., V. prisca  Bosquet, 1854 , V. pusilla  Bosquet, 1857 and V. tasmanica  Buckeridge, 1983 , is rationalized in the light of Tethyan palaeogeography.     

The distribution of cirripede larvae (Cirripedia: Thoracica) in the Amursky and Ussuriisky bays of the Sea of Japan

Based on data from plankton surveys that were carried out in the waters of the Amursky and Ussuriisky bays of the Sea of Japan from May to October 2007 and 2008, the species composition, time of occurrence, density, and distribution of cirripede larvae (Cirripedia: Thoracica) were studied. The larvae of five species of this group were revealed. Their average density does not exceed 400 ind./m3 and their proportion in the total meroplankton does not exceed 22%. In the summer season, the larvae of Amphibalanus improvisus prevailed and in the autumn season the nauplii of Balanus rostratus prevailed. The number of larvae of barnacles was higher in Amursky Bay, with deep-sea species nauplii being concentrated in its central part and shallow water ones in the northern and central parts, in the zones of costal gyres. In Ussuriisky Bay, the density of larvae of most species decreased seaward from the coast. The abundance of larvae of invasive species A. improvisus show that it has successfully become naturalized in Amursky Bay and is gradually replacing the native species Chthamalus dalli and Balanus crenatus in the lower littoral and upper sublittoral zones.     

On nine Tropodiaptomus-species (Copepoda,Calanoida) from equatorial East Africa

Of the nine species of Tropodiaptomus occurring in Equatorial East Africa, seven are redescribed and figured in great detail. Their relationships are discussed. A primary homonym is eliminated, and types are indicated wherever possible.     

Boschmaella japonica,New Species,a Parasite on the Barnacles Chthamalus challengeri and Balanus amphitrite amphitrite from Japan (Crustacea: Cirripedia: Rhizocephala: Chthamalophilidae)

Abstract A new rhizocephalan parasite which infests the barnacles Chthamalus challengeri and Balanus amphitrite amphitrite is described. It is known only from the type locality, Aburatsubo, Kanagawa Prefecture, Japan. Its position in the family Chthamalophilidae is assured by the cyprids lacking a thorax, by the position of the mesenteric canal opposite to the stalk, and by its host being a balanomorph barnacle. Among the three previously described species of the Chthamalophilidae, the new species most closely resembles Boschmaella balani, but it differs in details of the internal root system and in the externa being found both on the host body and on the inner mantle. About two-thirds of the infested hosts carry more than one externa.     

The barnacles (Cirripedia: Thoracica) collected by the Viking expedition in chemobiotic benthic communities of the Norwegian and Barents Seas

The barnacles (Cirripedia Thoracica) collected by the Viking expedition in chemobiotic benthic communities of Arctic Ocean-Nyegga area in Norwegian Sea and Håkon Mosby Mud Volcano area in the Barents Sea are investigated. The species (Weltnerium nymphocola in Håkon Mosby Mud Volcano area and Hamatoscalpellum hamatum in Nyegga area) that were found are common inhabitants of the Barents and Norwegian sea benthic communities. The absence of obligatory barnacle fauna that was revealed in investigated chemobiotic benthic communities of the Arctic Ocean shows they are similar to corresponding communities of the Atlantic Ocean, while the specific obligatory barnacle fauna is noted for chemobiotic benthic communities of the Pacific and Indian oceans.