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Most individuals of the rhizocephalan Peltogasterella gracilis (Boschma, 1927) produce only male (larger) or female (smaller) offspring, but some mixed-sex broods appear in autumn. Such variability may be an adaptation to the seasonally changing availability of hosts (for female larvae) and conspecific females (for male larvae), but details of the seasonal variation in the sex ratio and its ultimate causes have not been fully studied in P. gracilis. We studied the life cycle, host availability, and seasonal sex-ratio variation in P. gracilis by samplings from spring to autumn in 2017–2020 in Otaru, and additionally from winter to spring of 2007 in Oshoro Bay, Hokkaido, northern Japan. We measured its prevalence on the hermit crab Pagurus lanuginosus De Haan, 1849, the number of males per externa, the proportion of externae brooding different embryonic stages, and embryo size that are linked to sex. The prevalence was about 20% in most of the sampling period. The number of males per externa was 0.01–0.74 in June 2018–2020, but increased in later months to nearly 1.0. Eyed embryos and nauplii before hatching were common from June or July to September, but they were rare in other seasons. Female embryos were observed in all sampling periods, but male embryos were not found in July or August, by which time most females already have males implanted within their receptacles. It is suggested that P. gracilis releases both female and male larvae according to the availability of unparasitized hosts and of virgin externae, respectively.

From a number of partial moult series obtained by laboratory culture of plankton-caught specimens, we described all naupliar instars of the facetotectan species, Hansenocaris itoi. It reveals seven naupliar instars, instead of the five that were previously supposed for the Facetotecta. This number of naupliar instars is unique not only for Facetotecta, but also for Thecostraca and Hexanauplia as well. We studied the external morphology of each naupliar instar in detail with light microscopy and SEM. Nauplius 1 is entirely non-feeding and differs from subsequent instars in having a smooth, unsculptured cuticle and undeveloped armament of the labrum and the limbs. The subsequent naupliar development is characterized by an increase in size and in the number of cuticular plates and by the appearance of different armament of the limbs. We compare nauplius 1 across all Thecostraca and discuss its nature in relation to the larval development of Crustacea. We also discuss the presence of seven naupliar instars in Facetotecta and the concept of Hexanauplia.

Molecular and morphological methods are used to describe Lernaeodiscus rybakovi, a new rhizocephalan species parasitizing the porcellanid crab Pachycheles stevensii Stimpson, 1858, collected in Russian waters of the Sea of Japan. Molecular analysis of three species, including the new one, confirms the monophyly of the genus Lernaeodicus Muller, 1862 and its recent transfer to the family Peltogastridae. The main morphological features of the new species are also common with characters of the other species of the genus Lernaeodiscus Muller, 1862. Externa of L. rybakovi differs from the well studied Lernaeodiscus porcellanae from the E. Pacific by molecular markers, color and the absence of pronounced marginal lobes. Retinacula on the internal cuticle of L. rybakovi are found in the genus Lernaeodiscus for the first time. Some of the female hosts with adult externae were unusual in also carrying their own eggs on the pleopods, a rare situation among rhizocephalans. The complete larval development in the genus Lernaeodiscus is described here for the first time and includes five naupliar and one cypris instar. The main morphological features of Lernaeodiscus nauplii (the presence of flotation collar, morphology of the frontolateral horns and furcal spines, and the arrangement of dorsal setae on the shield head) are common with those of other known peltogastrid larvae, but also resemble nauplii of Peltogasterella in the presence of naupliar eyes and thin structure of the flotation collar. The sizes of male and female cyprids of L. rybakovi overlap slightly. In summer months, larval sex ratio is male-biased. We briefly review important larval characters in the Rhizocephala.

Although parasitic castration is widespread among rhizocephalan barnacles, Boschmaella japonica Deichmann & Høeg, 1990 does not completely sterilise the host barnacle Chthamalus challengeri Hoek, 1883. As little information is available on the relationships with the host in “barnacle-infesting parasitic barnacles” (family Chthamalophilidae), we studied the life cycles of both B. japonica and C. challengeri and the effects of the parasite on the host reproduction. Specimens of C. challengeri were collected from an upper intertidal shore at Shirahama, Wakayama, western Japan from April 2017 to September 2018 at 1–3 mo intervals. We recorded the body size, number of eggs, egg volume, and the presence of the parasite for each host. Moreover, settlement and growth of C. challengeri were followed in two fixed quadrats. Chthamalus challengeri brooded from February to June. The prevalence of B. japonica was high (often exceeded 10%) from April to July, and was rarely observed from September to next spring. The life cycle of the parasite matched well with that of the host. The parasite reduced the host’s brooding rate and brood size, to the extent that no hosts brooded in 2018.

How larvae of whale and dolphin epibionts settle on their fast-swimming and migrating hosts is a puzzling question in zoology. We successfully reared the larvae of the whale and dolphin barnacle Xenobalanus globicipitis to the cyprid stage. We studied the larval developmental ecology and antennular morphology in an attempt to assess whether an epibiotic lifestyle on this extreme substratum entails any unique larval specializations. Morphological parameters were compared with five other barnacle species that also inhabit extreme substrata. We found no larval specializations to a lifestyle associated with marine mammals. The external morphology of the antennules in Xenobalanus cyprids is morphologically similar to species from strikingly different substrata. We found variation only in the structures that are in physical contact with the substratum, i.e., the third segments carrying the villi-covered attachment disc. The third segments of the Xenobalanus cyprid antennules are not spear-shaped as in the stony coral barnacles, which are here used to penetrate the live tissue of their hosts. The presence of a cyprid cement gland implies that Xenobalanus uses cement protein when attaching to its cetacean host. Naupliar instars developed outside of the mantle cavity, indicating dispersal is planktonic. Our results militate against the idea that the cyprids settle during ocean migrations of their hosts. We suggest cyprids settle during coastal aggregations of the cetacean hosts. We conclude that the ecological success of barnacles has ultimately depended on a larva that with little structural alteration possesses the ability to settle on an amazingly wide array of substrata, including cetaceans.

The invasion of novel habitats is recognized as a major promotor of adaptive trait evolution in animals. We tested whether similar ecological niches entail independent and adaptive evolution of key phenotypic structures related to larval host invasion in distantly related taxa. We use disparately related clades of coral barnacles as our model system (Acrothoracica: Berndtia and Thoracica: Pyrgomatidae). We analyze the larval antennular phenotypes and functional morphologies facilitating host invasion. Extensive video recordings show that coral host invasion is carried out exclusively by cypris larvae with spear-shaped antennules. These first exercise a series of complex probing behaviors followed by repeated antennular penetration of the soft host tissues, which subsequently facilitates permanent invasion. Phylogenetic mapping of larval form and function related to niche invasion in 99 species of barnacles (Thecostraca) compellingly shows that the spear-phenotype is uniquely associated with corals and penetrative behaviors. These features evolved independently in the two coral barnacle clades and from ancestors with fundamentally different antennular phenotypes. The larval host invasion system in coral barnacles likely evolved adaptively across millions of years for overcoming challenges associated with invading and entering demanding coral hosts. This article is protected by copyright. All rights reserved.

Symbiosis is increasingly recognized as being an important component in marine systems, and many such relationships are initiated when free-swimming larvae of one partner settle and become sedentary on a host partner. Therefore, several crucial questions emerge such as the larva's mechanism of locating a host, selection of substratum and finally settlement on the surface of its future partner. Here, we investigated these mechanisms by studying how larvae of the fire coral-associated barnacle

Movement is a fundamental characteristic of life, yet some invertebrate taxa, such as barnacles, permanently affix to a substratum as adults. Adult barnacles became ‘sessile’ over 500 Ma; however, we confirm that the epizoic sea turtle barnacle, Chelonibia testudinaria , has evolved the capacity for self-directed locomotion as adults. We also assess how these movements are affected by water currents and the distance between conspecifics. Finally, we microscopically examine the barnacle cement. Chelonibia testudinaria moved distances up to 78.6 mm yr −1 on loggerhead and green sea turtle hosts. Movements on live hosts and on acrylic panels occasionally involved abrupt course alterations of up to 90°. Our findings showed that barnacles tended to move directly against water flow and independent of nearby conspecifics. This suggests that these movements are not passively driven by external forces and instead are behaviourally directed. In addition, it indicates that these movements function primarily to facilitate feeding, not reproduction. While the mechanism enabling movement remained elusive, we observed that trails of cement bore signs of multi-layered, episodic secretion. We speculate that proximal causes of movement involve one or a combination of rapid shell growth, cement secretion coordinated with basal membrane lifting, and directed contraction of basal perimeter muscles.

Molecular and morphological methods are used to describe the rhizocephalan Lernaeodiscus kasyanovi sp. nov. from Russian waters of the Sea of Japan, the second of two Lernaeodiscus species parasitizing the porcellanid crab Pachycheles stevensii Stimpson, 1858. Lernaeodiscus kasyanovi sp. nov. differs from the other species found on this host, Lernaeodiscus rybakovi Korn et al., 2020a, by molecular markers, by a smaller size, a lighter color, by the shape of receptacles, by the presence of marginal lobes in mature specimens, and by the retinacula type on the internal cuticle. Lernaeodiscus kasyanovi sp. nov. is the sister taxon to Lernaeodiscus porcellanae in the monophyletic genus Lernaeodiscus. Lernaeodiscus kasyanovi is rather rare compared to L. rybakovi, their prevalence on the porcellanid crab P. stevensii does not exceed 2%. It is the second example of two rhizocephalan species infesting the same brachyuran species in Peter the Great Bay.

Sponges are common in coral reefs and provide secondary habitats and shelter to a very diverse associated biota. To examine the symbiotic relationships between crustacean associates and their sponge hosts, the most important step is to collect live crustaceans and sponges for subsequent taxonomic identification as well as for larval rearing and experiments on larval biology. Using sponge-inhabiting barnacles as a model, we describe a set of collection procedures, identification methods, and laboratory-rearing systems for maintaining living barnacles and their host sponges. These methods also permit observing the behavior of the barnacle symbionts, including feeding, mating, as well as larval development and settlement, information that can be applied to the study of host-specificity, larval biology, and host selection.

We present a comprehensive revision and synthesis of the higher-level classification of the barnacles (Crustacea: Thecostraca) to the genus level and including both extant and fossils forms. We provide estimates of the number of species in each group. Our classification scheme has been updated based on insights from recent phylogenetic studies and attempts to adjust the higher-level classifications to represent evolutionary lineages better, while documenting the evolutionary diversity of the barnacles. Except where specifically noted, recognized taxa down to family are argued to be monophyletic from molecular analysis and/or morphological data. Our resulting classification divides the Thecostraca into the subclasses Facetotecta, Ascothoracida and Cirripedia. The whole class now contains 14 orders, 65 families and 367 genera. We estimate that barnacles consist of 2116 species. The taxonomy is accompanied by a discussion of major morphological events in barnacle evolution and justifications for the various rearrangements we propose.

The regression and salinization of the Aral Sea, largely caused by water diversion for irrigation, is among the most severe ecological disasters of the 20th century, and has had severe health and economic consequences for the local population. Introductions of alien species to enhance commercial fisheries before the regression had already impacted the ecology of this system. Crustaceans made up about one-quarter of the original metazoan species and constituted the principal food for native and introduced fish. From 1960 on, crustaceans were recorded at numerous fixed sampling stations, including thanatocoenoses (dead animals from sediment cores). We use this previously unpublished information to document changes in species abundance and discuss their causes in the context of species interactions and changes to physical and chemical parameters. Competition from alien crustaceans led to declines in or even extinction of some native species, but eventually severe salinization became the main detriment, and resulted in the complete collapse of commercial fisheries. This seriously hurt a critical trade, which provided the principal protein source for the local population. We document how comparatively modest conservation efforts enabled the northern Small Aral Sea to partially recover and commercial fishing to resume.

Symbiotic relations and range of host usage are prominent in coral reefs and crucial to the stability of such systems. In order to explain how symbiotic relations are established and evolve, we used sponge-associated barnacles to ask three questions. (1) Does larval settlement on sponge hosts require novel adaptations facilitating symbiosis? (2) How do larvae settle and start life on their hosts? (3) How has this remarkable symbiotic lifestyle involving many barnacle species evolved? We found that the larvae (cyprids) of sponge-associated barnacles show a remarkably high level of interspecific variation compared with other barnacles. We document that variation in larval attachment devices are specifically related to properties of the surface on which they attach and metamorphose. Mapping of the larval and sponge surface features onto a molecular-based phylogeny showed that sponge symbiosis evolved separately at least three times within barnacles, with the same adaptive features being found in all larvae irrespective of phylogenetic relatedness. Furthermore, the metamorphosis of two species proceeded very differently, with one species remaining superficially on the host and developing a set of white calcareous structures, the other embedding itself into the live host tissue almost immediately after settlement. We argue that such a high degree of evolutionary flexibility of barnacle larvae played an important role in the successful evolution of complex symbiotic relationships in both coral reefs and other marine systems.

Rhizocephalans are abundant members of marine ecosystems and are important regulators of crustacean host populations. Morphological and ecological variation makes them an attractive system for evolutionary studies of advanced parasitism. Such studies have been impeded by a largely formalistic taxonomy, because rhizocephalan morphology offers no characters for a robust phylogenetic analysis. We use DNA sequence data to estimate a new phylogeny for 43 species and use this to develop a revised taxonomy for all Rhizocephala. Our taxonomy accepts 13 new or redefined monophyletic families. The traditional subdivision into the suborders Kentrogonida and Akentrogonida is abandoned, because both are polyphyletic. The three ‘classical’ kentrogonid families are also polyphyletic, including the species-rich Sacculinidae, which is split into a redefined and a new family. Most species of large families remain to be studied based on molecular evidence and are therefore still assigned to their current genus and family by default. We caution against undue generalizations from studies on model species until a more stable species-level taxonomy is also available, which requires more extensive genus- and species-level sampling with molecular tools. We briefly discuss the most promising future studies that will be facilitated by this new phylogeny-based taxonomy.

The morphology of Tantulocarida, a group of minutely sized ectoparasitic Crustacea, is described here using for the first time transmission electron microscopy (TEM) and confocal laser scanning microscopy (CLSM). This enabled a detailed analysis of their internal anatomy to a level of detail not possible with previous light microscopic investigations. We studied the infective stage, the tantulus larva, attached on the crustacean host in two species, Arcticotantulus pertzovi and Microdajus tchesunovi, and put special emphasis on cuticular structures, muscles, adhesive glands used in attachment, sensory and nervous systems, and on the organs used in obtaining nutrients from the host. This allowed description of structures that have remained enigmatic or unknown until now. A doubly folded cuticular attachment disc is located at the anterior-ventral part of the cephalon and used for gluing the larva to the host surface with a cement substance released under the disc. Four cuticular canals run from a cement gland, located ventrally in the cephalon, and enter an unpaired, cuticular proboscis. The proboscis can be protruded outside through a separate opening above the mouth and is used for releasing cement. An unpaired and anteriorly completely solid cuticular stylet is located centrally in the cephalon and is used only for making a 1-μm-diameter hole in the host cuticle, through which passes a rootlet system used for obtaining nutrients. The rootlet system is a direct extension of the anterior gut of the tantulus, but inside the host, it consists of cuticle only. Several muscular systems seem to degenerate very quickly after the tantulus has settled on the host. The central nervous system is reduced to an absolute minimum in terms of both cell size and number. We discuss the morphology of the tantulus in light of the Tantulocarida being a sister group to the Thecostraca or even nested within that taxon.

Mating behaviour between a dwarf male and its hermaphrodite partner was observed for the first time in cirripedes using the androdioecious barnacle Scalpellum scalpellum. Mating between hermaphrodites was also observed. The dwarf males are located on the rim of the mantle cavity of the hermaphrodite partner. When mating, the male extends the penis, which is four times longer than its body. The penis first assumes a straight stance where it is waved around in a searching mode. Upon touching the cirri of the hermaphrodite, the penis and the cirri engage in prolonged contact during which hermaphrodite feeding is suspended. Thereafter the penis assumes a U-bend to reach into the brood chamber, where after the mantle valves are closed tightly around the penis. The nearly transparent penis is a tube of very thin cuticle, equipped with pairs of side branches but not containing any visible tissue. The penis enables the minute male, situated outside the brood chamber, to securely deposit sperm into its partner. Adjacently situated hermaphrodites interact socially in between feeding sessions by reorienting themselves on the peduncle to touch each other with their cirri. This can be followed by precopulatory behaviour, where one or both individuals extend the penis to touch their partner, leading again to actual copulation where the penis of one individual is inserted into the other. We discuss the results in the context of the diverse reproductive strategies found in cirripede barnacles.

Barnacle cypris antennules are important for substratum attachment during settlement and on through metamorphosis from the larval stage to sessile adult. Studies on the morphology of cirripede cyprids are mostly qualitative, based on descriptions from images obtained using a scanning electron microscope (SEM). To our knowledge, our study is the first to use scanning electron microscopy to quantify overall structural diversity in cypris antennules by measuring 26 morphological parameters, including the structure of sensory organs. We analyzed cyprids from seven species of balanomorphan barnacles inhabiting rocky shore communities; for comparison, we also included a sponge-inhabiting balanomorphan and a verrucomorphan species. Multivariate analysis of the structural parameters resulted in two distinct clusters of species. From nonmetric multidimensional scaling plots, the sponge-inhabiting Balanus spongicola and Verruca stroemia formed one cluster, while the other balanomorphan species, all from hard bottoms, grouped together in the other cluster. The shape of the attachment disk on segment 3 is the key parameter responsible for the separation into two clusters. The present results show that species from a coastal hard-bottom habitat may share a nearly identical antennular structure that is distinct from barnacles from other habitats, and this finding supports the fact that such species also have rather similar reactions to substratum cues during settlement. Any differences that may be found in settlement biology among such species must therefore be due either to differences in the properties of their adhesive mechanisms or to the way that sensory stimuli are detected by virtually identical setae and processed into settlement behavior by the cyprid.

Barnacles and their allies (Thecostraca) are a biologically diverse, monophyletic crustacean group, which includes both intensely studied taxa, such as the acorn and stalked barnacles, as well as cryptic taxa, for example, Facetotecta. Recent efforts have clarified phylogenetic relationships in many different parts of the barnacle tree, but the outcomes of these phylogenetic studies have not yet been combined into a single hypothesis for all barnacles. In the present study, we applied a new “synthesis” tree approach to estimate the first working Barnacle Tree of Life. Using this approach, we integrated phylogenetic hypotheses from 27 studies, which did not necessarily include the same taxa or used the same characters, with hierarchical taxonomic information for all recognized species. This first synthesis tree contains 2,070 barnacle species and subspecies, including 239 barnacle species with phylogenetic information and 198 undescribed or unidentified species. The tree had 442 bifurcating nodes, indicating that 79.3% of all nodes are still unresolved. We found that the acorn and stalked barnacles, the Thoracica, and the parasitic Rhizocephala have the largest amount of published phylogenetic information. About half of the thecostracan families for which phylogenetic information was available were polyphyletic. We queried publicly available geographic occurrence databases for the group, gaining a sense of geographic gaps and hotspots in our phylogenetic knowledge. Phylogenetic information is especially lacking for deep sea and Arctic taxa, but even coastal species are not fully incorporated into phylogenetic studies.

We use Rhizocephala to illustrate the problems inherent in estimating the phylogeny of parasitic crustaceans. The adult rhizocephalan parasite has such a reduced morphology that little else than the presence of a moulted cuticle relegates them to Arthropoda. Therefore, until the advent of molecular phylogenetic analysis relegation of the Rhizocephala to taxon relied exclusively on larval characters. The reduced adult morphology also entailed that very few characters were available for intrinsic rhizocephalan systematics and virtually none that could be compared with any outgroup. Thus, rhizocephalan taxonomy relied only on analysis within the group and, with few exceptions, was not based on any phylogenetic principles. The advent of DNA methods in phylogeny confirmed, with high confidence, rhizocephalans as cirripedes and nested them within the taxon as the sister group to Thoracica. This result was supported by SEM studies of the cypris larvae. Additional molecular phylogenetic studies yielded detailed insight into rhizocephalan relationships, culminating in the phylogeny presented here, which includes species from all but one family. Contrary to traditional hypotheses, the new phylogeny shows that infection of the host using a kentrogon stage represents the plesiomorphic condition, while rhizocephalans (Akentrogonida) without this stage are advanced. In this analysis, both Kentrogonida and the species-rich Sacculinidae are paraphyletic. In addition, recent hypotheses on family-level relationships in Akentrogonida, based on larval structure and the sexual system, were largely confirmed. This shows that when accurately analysed in a strict homology regime, morphological characters can be powerful partners to molecular data in elucidating rhizocephalan phylogeny.

This study used morphometric analyses to compare the structure of the third antennular segment, also called the attachment organ, in cyprid larvae from cirripede species representing a diverse set of taxonomic groups. The aim was to investigate the degree of morphological variation in view of the diversity of habitats, settlement substrata, and modes of life found in the Cirripedia. In all cyprids the third segment features a flat surface (the attachment disc) covered with small cuticular villi thought to function in adhesion. The parameters analyzed were the angle of this disc relative to the long axis of the antennule, its shape (outline), the density of cuticular villi, and the type of cuticular structure encircling the disc. The 10 species studied came from most major groups of cirripedes, and comprised shallow-water forms inhabiting hard bottoms (Capitulum mitella, Pollicipes pollicipes, Semibalanus balanoides, Austrominius modestus, Megabalanus rosa), sublittoral forms (Verruca stroemia, Scalpellum scalpellum), epibiotic forms settling on live, soft tissues (Balanus spongicola, Savignium crenatum), and a parasite (Peltogaster paguri). Significant structural variation was found among the species, but due to limited taxon sampling it was unclear whether the differences relate to ecological factors or phylogenetic affiliation. The disc perimeter is guarded by either a series of long and thin cuticular fringes overreaching the rim of the disc (= a velum) or a few low, but very broad cuticular flaps (= a skirt). The presence of a velum (in all rocky-shore species) or a skirt (all other species) around the attachment disc was the only parameter that was clearly correlated with habitat. The shape of the third antennular segment varied from a symmetrical bell shape with a distally facing attachment disc having a circular disc outline, to segments that were elongated in side view, with a very tilted ventral disc surface having an elliptical disc outline. The bell shape may be most common in forms from rocky shores, but in our test of morphometric parameters only Scalpellum scalpellum (sublittoral), Savignium crenatum (epibiotic in corals), and Peltogaster paguri (parasitic) had shapes that differed significantly from the other species. The density of villi on the attachment disc varied significantly, but also showed no clear-cut correlation with substratum or habitat. Attachment organ structure is clearly the most variable feature in cirripede cyprids. To evaluate the degree to which attachment organ structure is correlated with habitat, settlement substratum, and mode of life, future studies should employ a more refined statistical analysis on an enlarged dataset, with much increased taxon sampling and a more multifaceted definition of ecological variables.

Barnacle cypris larvae show high morphological variation in the organs used in search of and attaching to a substratum. This variation may represent adaptation to the habitat of the species. Here, we studied SEM level morphologies of cypris antennular sensory and attachment organs in a deep-sea vent endemic species (Neoverruca sp.) and a vent/seep inhabiting species (Ashinkailepas seepiophila). We compare them with three species from other environments. The antennular morphologies of Neoverruca sp. and A. seepiophila were similar, which is consistent with recent molecular studies showing a close relationship of the two species. The setation pattern of the antennules was very conservative among species from various environments. In contrast, striking differences were observed in the structure of the attachment organ (the third antennular segment). Neoverruca sp. and A. seepiophila had no velum or a skirt surrounding the attachment disc on the third segment, while other cirripede cyprids almost always have either of these structures. In addition, both cyprids of A. seepiophila and Neoverruca sp. had the attachment disc angled toward the substratum, whereas it faces distally in cyprids from hard bottom inhabiting barnacles. We suggest that both velum/skirt and the angle of the attachment disc play an important role, when the antennules are contacting the substratum during surface exploration. Differences in attachment organ structures may be highly adaptive, enabling cirripede species to enter new habitats during evolution.

New taxa of thoracican cirripedes are recorded from the Eocene Mo Clay of northwest Jutland, Denmark, namely Stipilepas molerensis Carriol n. gen., n. sp., a scalpelliform gooseneck barnacle, and Plesiobrachylepas jutlandica Carriol n. gen., n. sp., a brachylepadomorph sessile form. This material is of importance in that the former not only represents the first extinct example of the tribe Ashinkailepadini, it also is the second fossil neolepadine, while the latter adds to the diversity of the family Brachylepadidae, which previously embraced only three genera. The geological context of these taxa is outlined and their phylogenetic interest discussed.

The ecologically important shore crab Carcinus maenas is commonly infected in its native range by the rhizocephalan Sacculina carcini. However, several aspects of this host-parasite interaction are poorly understood. Here, we analyse data from approximately 60000 Danish crabs to unravel factors governing infection patterns in time and space, and according to host sex and size. Female crabs were more frequently infected (12.6%) than males (7.9%). Sites with high salinity supported the highest infection prevalence. Infection prevalence peaked in summer (10 to 15%) and winter (20 to 35%) due in part to emergence of virginal externae in summer (main outbreak) and autumn (minor outbreak) preceded by peaks in crabs with lost externa (scars). Younger externae and scars dominated among males, whereas adult externae were most frequent among females. Infection prevalence increased with size in females but decreased in males, and modified (feminized) males showed lower scar frequency than unmodified ones. Modified males occurred frequently among the smaller size classes, whereas unmodified males dominated the larger size classes. Externa size was positively related to host size in both genders (same linear relationship). Molecular analyses suggested that hosts below 16 mm in carapace width do not become infected. Dissections of infected hosts revealed marked reduction of ovaries, whereas testes were unaffected by sacculinization. Our study demonstrates great spatio-temporal variation in infection prevalence mainly related to the parasite's life history. S. carcini appears capable of infecting all host sizes except the smallest. Owing to incomplete feminization of males, infections are rapidly lost from the larger and highly profitable male hosts.

Facetotecta, or crustacean "y-larvae," occur in all the world's oceans although the adult forms remain completely unknown. At the metamorphic molt from the last naupliar instar to the terminal cypris larval stage a free carapace, six pairs of natatory thoracopods, and a segmented thorax and abdomen all develop anew. Unlike in earlier molts, the cephalic shield and the so-called "faciotruncal integument" usually remain together at this last naupliar molt, and the posterior "trunk" portion of the exuviae, while hollow, is not empty. In mounted preparations examined by phase contrast or differential interference contrast (DIC) microscopy, a ghost-like image of part of the cypris thorax, particularly the thoracopods and even their setae, is commonly visible inside the naupliar exuviae, and may be universally present in the Facetotecta. To investigate this "ghost," we used DIC and digital photographic stacking, and also scanning electron microscopy, on slide or stub-mounted final naupliar exuviae of an assortment of undescribed species of Facetotecta that had been reared from planktonic lecithotropic nauplii to the cypris stage at Sesoko Island, Okinawa, Japan, and at Keelung and Green Island, Taiwan. These techniques showed that the "ghost" is a delicate, three-dimensional, fibrous structure, essentially a sling-like mold or matrix with struts attached to the outer cuticle and pairs of deep pockets that previously held the thoracopods of the developing cypris y. Whether it is endoskeletal in nature, the (partial) exuvia of an additional instar, remnants of apoptosis, or something else is currently unknown. Nothing similar has been reported in other thecostracans, or in other crustaceans that undergo a similarly abrupt metamorphosis at the last naupliar molt.

How androdioecy (coexistence of hermaphrodites and males) is maintained is still poorly understood. Therefore, sex determination was studied in the androdioecious barnacle Scalpellum scalpellum L. First, 247 cypris larvae from seven broods were investigated for sexual dimorphism in larval morphology and found to be all identical. Second, experiments with cyprids showed that males and hermaphrodites differ distinctly in morphology as soon as 4–5 days after settlement. Third, 14 252 cyprids were allowed to settle on the bottom of their culture cages, and all surviving larvae developed into hermaphrodites and none into dwarf males. Fourth, larvae settled in hermaphrodite receptacles (i.e. future males) were removed at increasing intervals after settlement to study if the male and hermaphrodite sexual expressions are fixed or plastic. All larvae became dwarf males if allowed to stay there for more than 8 h after settlement. But if removed within 3 h after settlement, half of them developed into hermaphrodites. We conclude that an environmental sex determination mechanism operates in S. scalpellum. Together with a 1:1 hermaphrodite/male ratio observed in previously reported experiments offering a free choice of settlement, we suggest that all larvae are potential hermaphrodites, but only 50% can settle in hermaphrodite receptacles and yield males.

Barnacles are exceptional in having various sexual systems (androdioecy, hermaphroditism, dioecy) and with a high morphological diversity of males, though these are always minute (dwarf) compared to their female or hermaphrodite partners. For the first time, we use a multiple DNA marker-based phylogeny to elucidate the ancestral states and evolution of (1) dwarf males, (2) their morphology when present, (3) their attachment site on the partner, and (4) habitat use in thoracican barnacles. Our taxon sampling was especially rich in rare deep-sea Scalpelliformes and comprised species with diverse sexual systems and dwarf male morphologies. Within the thoracican barnacles dwarf male evolution is subject to extensive convergence, but always correlated to similar ecological conditions. Males evolved convergently at least four times from purely hermaphroditic ancestors, in each case correlated with the invasion into habitats with low mating group sizes. The independent evolution of dwarf males in these lineages dovetails with the males having different morphologies and occurring in several different locations on their sexual partner.

Rhizocephala, a group of parasitic castrators of other crustaceans, shows remarkable morphological adaptations to their lifestyle. The adult female parasite consists of a body that can be differentiated into two distinct regions: a sac-like structure containing the reproductive organs (the externa), and a trophic, root like system situated inside the hosts body (the interna). Parasitism results in the castration of their hosts, achieved by absorbing the entire reproductive energy of the host. Thus, the ratio of the host and parasite sizes is crucial for the understanding of the parasite’s energetic cost. Using advanced imaging methods (micro-CT in conjunction with 3D modeling), we measured the volume of parasitic structures (externa, interna, egg mass, egg number, visceral mass) and the volume of the entire host. Our results show positive correlations between the volume of (1) entire rhizocephalan (externa + interna) and host body, (2) rhizocephalan externa and host body, (3) rhizocephalan visceral mass and rhizocephalan body, (4) egg mass and rhizocephalan externa, (5) rhizocephalan egg mass and their egg number. Comparing the rhizocephalan Sylon hippolytes, a parasite of caridean shrimps, and representatives of Peltogaster, parasites of hermit crabs, we could match their different traits on a reconstructed relationship. With this study we add new and significant information to our global understanding of the evolution of parasitic castrators, of interactions between a parasitic castrator and its host and of different parasitic strategies within parasitic castrators exemplified by rhizocephalans. publishedVersion

The larval phase of metazoans can be interpreted as a discrete post-embryonic period. Larvae have been usually considered to be small, yet some metazoans possess unusually large larvae, or giant larvae. Here, we report a possible case of such a giant larva from the Upper Jurassic Solnhofen Lithographic limestones (150 million years old, southern Germany), most likely representing an immature cirripede crustacean (barnacles and their relatives). The single specimen was documented with up-to-date imaging methods (macro-photography, stereo-photography, fluorescence photography, composite imaging) and compared with modern cirripede larvae. The identification is based on two conspicuous spine-like extensions in the anterior region of the specimen strongly resembling the so-called fronto-lateral horns, structures exclusively known from cirripede nauplius larvae. Notably, at 5 mm in length the specimen is unusually large for a cirripede nauplius. We therefore consider it to be a giant larva and discuss possible ecological and physiological mechanisms leading to the appearance of giant larvae in other lineages. Further findings of fossil larvae and especially nauplii might give new insights into larval evolution and plankton composition in the past.

Summary In his monograph on Cirripedia from 1851 [1], Darwin pointed to a highly unusual, plateless, and most likely parasitic barnacle of uncertain phylogenetic affinity. Darwin’s barnacle was Anelasma squalicola, found on deep-water sharks of the family Etmopteridae, or lantern sharks [2]. The barnacle is uncommon and is therefore rarely studied. Recent observations by us have shown that they occur at an unusually high prevalence on the velvet belly lantern shark, Etmopterus spinax, in restricted fjord areas of western Norway. A phylogenetic analysis based on ribosomal DNA data (16S, 18S, and 28S) from 99 selected barnacle species, including all available pedunculate barnacle sequences from GenBank, shows that A. squalicola is most closely related (sister taxon) to the pedunculate barnacle Capitulum mitella. Both C. mitella and species of Pollicipes, situated one node higher in the tree, are conventional suspension feeders from the rocky intertidal. Our phylogenetic analysis now makes it possible to establish morphological homologies between A. squalicola and its sister taxon and provides the evolutionary framework to explain the unprecedented transition from a filter-feeding barnacle to a parasitic mode of life.

Parasites generally influence the feeding behavior of their host and may therefore indirectly impact ecosystem structure and functioning if the host plays an ecological key role. The ecologically important shore crab ( Carcinus maenas ) is commonly infected by the rhizocephalan parasite Sacculina carcini that aside from inflicting behavioral change, castration and ceased molting, also feminizes its male host morphologically. The latter results in reduced cheliped size, and, together with the other parasite-induced effects, this may potentially impact host feeding behavior. In two separate laboratory experiments, we offered infected and uninfected adult male crabs respectively ad libitum small, easy-to-handle blue mussels ( Mytilus edulis ) (10–15 mm in shell-length), and a limited, size-structured prey population (15–45 mm in shell-length; seven size-classes, ten mussels per class) during 10–15 days. Corrected for carapace width, the per capita consumption rate of the infected and uninfected crabs was similar in either experiment, both regarding number of mussels and amount of tissue dry-weight consumed. Also, the median mussel size preyed upon when exposed to the size-structure prey population was unaffected by infection. However, infected crabs preyed less frequently (26%) on intermediate mussel sizes (25–30 mm) than uninfected crabs. For both infected and uninfected crabs the median prey size increased linearly with maximum claw height. Host dry weight was significantly reduced by infection, assumed to be the result of the morphological feminization (reduced appendage size) rather than reflecting poorer general condition of infected individuals. Infected crabs were nonetheless subjected to a higher mortality rate than uninfected crabs during the experimental period. We conclude that Sacculina infection has a very limited effect on its host crabs' feeding biology and that any measurable ecosystem ramifications of the host–parasite association must originate from other processes; for instance reduced mean size (infection inhibits molting) and density (infection increases mortality) of the crab population where parasitism is high.

Deep-sea hydrothermal vents and cold seeps are scattered and limited habitats, and species that are confined to these benthic habitats often have a planktonic larval stage in their life history to maintain dispersal and gene flow among discrete populations. We have reared the larvae of Ashinkailepas seepiophila , inhabiting inactive vents and active cold seeps, through their entire larval development under atmospheric pressure. Using two temperatures (4 °C, 10 °C) we examined whether larval development resembles other vent species in being affected by thermal stimulation. Under laboratory conditions, A. seepiophila completed larval development in 65 days at 4 °C. This is considerably shorter than the 100 days previously reported to be required at the same temperature by Neoverruca sp., a species confined to inhabiting active vents. Also in contrast to Neoverruca sp., A. seepiophila did not drastically shorten the developmental period from nauplius VI to cyprid when exposed to temperature of 10 °C. We conclude that there is no need for late naupliar stages of A. seepiophila to develop into cyprids more quickly at elevated temperatures because this species does not inhabit active vents. The difference in response to water temperature during larval development may contribute to the separated distribution of the two species at active vents ( Neoverruca sp.) and inactive vents and cold seep ( A. seepiophila ) environments.

Cypris settlement and metamorphosis into dwarf males were studied in the androdioecious barnacle Scalpellum scalpellum using field collected samples from the North Sea, and experiments with laboratory reared larvae, observed with video. In the field sample, dwarf males were always situated on the rim of the mantle aperture and almost invariably confined to two areas (receptacles) located along the scutal plates near their contact to the terga. In the laboratory experiments, cyprids settling on the mantle rim always developed into males. Those settling elsewhere, whether on the external surfaces of the adults, or on their hydroid substratum, always developed into hermaphrodites. The numbers settling as males did not differ significantly from those settling as hermaphrodites, suggesting that genetic sex determination may operate in S. scalpellum . The N. Sea sample comprised 52 adult hermaphrodites. Of these 15 (29%) lacked males altogether, while 37 (71%) carried males with an average of 4.7 per hermaphrodite. On the hermaphrodite, a thin lamella along the mantle rim protects the settling and metamorphosing male from accidental damage or dislocation by the beating cirri. The cyprid gains additional protection by starting almost immediately after settlement to penetrate into the receptacle tissue. After 12–24 h the developing dwarf male can be almost completely buried. It appears that the cyprid does not use any preformed burrow, but actively descends into the hermaphrodite integument.

The thecostracans include the Facetotecta, Ascothoracida, and Cirripedia and show great diversity in both morphology and biology. This makes them ideal models for studying evolutionary adaptations of the larval and adult body-plan, lifestyle, and reproduction. Surprisingly, despite all the work published since Darwin's seminal monographs, few studies have tested evolutionary hypotheses about Thecostraca within a phylogenetic context. In this review, we combine a Bayesian phylogenetic method and multilocus sequence data to reconstruct the evolutionary history of 12 key thecostracan phenotypic traits associated with their lifecycle, larval biology, reproduction, and adult morphology. Our analyses show that thecostracan biological diversity resulted both from unique innovations and from events of convergence. This provides an opportunity to reevaluate previous classifications of the Thecostraca and the theories relating to the origin and diversification of this taxon.

How and why diverse sexual systems evolve are fascinating evolutionary questions, but few empirical studies have dealt with these questions in animals. Pedunculate (gooseneck) barnacles show such diversity, including simultaneous hermaphroditism, coexistence of dwarf males and hermaphrodites (androdioecy), and coexistence of dwarf males and females (dioecy). Here, we report the first phylogenetically controlled test of the hypothesis that the ultimate cause of the diverse sexual systems and presence of dwarf males in this group is limited mating opportunities for non-dwarf individuals, owing to mating in small groups. Within the pedunculate barnacle phylogeny, dwarf males and females have evolved repeatedly. Females are more likely to evolve in androdioecious than hermaphroditic populations, suggesting that evolution of dwarf males has preceded that of females in pedunculates. Both dwarf males and females are associated with a higher proportion of solitary individuals in the population, corroborating the hypothesis that limited mating opportunities have favoured evolution of these diverse sexual systems, which have puzzled biologists since Darwin.

article i nfo We used video, light and scanning electron microscopy to study the cypris larvae in species of coral inhabiting barnacles (Pyrgomatidae) in search of adaptations to settlement on their highly unusual substratum. Species studied were Savignium crenatum, Trevathana jensi, Trevathana margaretae, Trevathana mizrachae and Trevathana sarae. In all five species the third antennular segment was shaped like a spearhead with only an extremely narrow attachment disc. This morphology represents the most extreme antennular specialization known from cirripedes, and it is not even matched by those parasitic barnacles that use the antennule for active penetration of cuticles. Compared to the third segment, the first, second and fourth segments exhibit no obvious specializations, and the armament of sensory setae is also as very comparable to that seen in balanomorphan cirripedes. The coral barnacles also have the nauplius eye, compound eyes, frontal filaments, lattice organs and cement glands known from other barnacles. Only T. sarae differed by having two unusually shaped setae terminally on the fourth segment. Video observations showed that the coral barnacle cyprids display the exploratory walking behaviour known from cyprids settling on hard bottom substrata. We interpret the pointed structure of the third antennular segment as an adaptation to penetrate through live coral tissue. Projecting this character onto a recent phylogenetic tree of balanomorphan species indicates that it is an apomorphy for a large clade of coral inhabiting barnacles. © 2010 Elsevier B.V. All rights reserved.

Androdioecy (co-existence of hermaphrodites and dwarf males) is a fascinating yet poorly understood phenomenon. The pedunculated barnacle Scalpellum scalpellum is an emerging model species for the system. In S. scalpellum, dwarf males and hermaphrodites are very different in adult morphology (e.g., in feeding structures and reproductive organs), but they share the same larval development with nauplii followed by cypris larvae. Recently, it was found that S. scalpellum cypris larvae display both genetic and environmental sex determination, but no detailed morphological study has yet investigated how the settled cypris larvae differ subsequent to settlement. This study investigates the morphological aspects of the onset of sex determination in the cyprids of S. scalpellum by examining their metamorphosis into either dwarf males or hermaphrodites under laboratory conditions. This study emphasizes morphological differences, such as size and shape of primordial shell plates, development of a flexible peduncle and of thoracopods. It was shown that the cypris larvae start to differ already one day after settlement on either a hydroid (leading to hermaphrodites) or an adult hermaphrodite (leading to dwarf males). Dwarf males gradually developed an ovoid body shape and two pairs of circular scutal and tergal primordia. Such cyprids developed neither a carina nor any peduncle or cirri for feeding. The study concludes that the dwarf males of S. scalpellum are not just hermaphrodites arrested early in development. This entails that dwarf males constitute their own separate developmental pathways and points to S. scalpellum dwarf males being more specialized than previously stated. Finally, the study compares differences in dwarf male morphology between S. scalpellum with two other androdioecious species with less specialized dwarf males and use this to discuss evolutionary implications for the adaptive evolution of dwarf males across the Cirripedia.

Cirripedes are one of the major groups of fouling organism in the marine environment. The cyprid can, before a permanent attachment, actively explore and walk on the substratum using its antennules in a bipedal fashion without leaving the surface. Studying the structure of the cyprid antennule is therefore important for understanding the events that culminate in biofouling by barnacles. There are at present no complete, standardised accounts of the structure of the cyprid antennules in thoracican barnacles, and moreover, the existing accounts vary in their use of terminology. This article describes the cyprid antennule of the barnacle Megabalanus rosa. This barnacle species is common in E Asia, and the cyprids have previously been used in several biofouling studies. All externally visible setae on the antennules have been mapped; these comprise both chemosensors with a terminal pore, a putative aesthetasc-like seta and mechano-sensory setae. More setae were found on the attachment disc than in previous scanning electron microscope-based studies, but not all structures that can be seen with transmission electron microscopy were visible. The disc itself seems to have a variable surface area, which could assist in exploring rough surfaces. The various lengths of the antennular setae, coupled with the disposition of the segments, enable the cyprid to cover a wide swath of substratum during exploratory walking. A new terminology is proposed for cyprid antennular setae, which will form a basis for future comparative and functional studies of cirripede settlement.

We used scanning electron microscopy (SEM) to describe cypris morphology in species of the barnacles Ibla and Paralepas, both of which are pivotal in understanding cirripede evolution. In Ibla, we also studied late naupliar stages with video and SEM. Special emphasis was put on the lattice organs, the antennules and the thorax and telson. In Paralepas we had settled specimens only and could therefore only investigate the carapace with the lattice organs. Cyprids of Ibla quadrivalvis and Paralepas dannevigi have five sets of lattice organs, grouped as two anterior and three posterior pairs. The organs are of the pore-field type and the terminal pore is situated anteriorly in the first pair, just as in the Rhizocephala and the Thoracica. In Ibla the armament of antennular sensilla resembles that found in the Thoracica but differs from the Rhizocephala. The absence of setules on the A and B setae sited terminally on the fourth antennular segment is a similarity with the Acrothoracica. The attachment disc is angled rather than facing distally and is encircled by a low cuticular velum. The thoracopods have two-segmented endopods and exopods as in the Thoracica, but the number, shape, and position of thoracopodal setae differ somewhat from other species of that superorder. Both Ibla and Paralepas cyprids have a deeply cleaved telson, but no independent abdominal part. In cypris morphology, Ibla and Paralepas show several synapomorphies with the clade comprising Rhizocephala and Thoracica and there are no specific apomorphies with either the Acrothoracica, the Rhizocephala or any particular subgroup within the Thoracica. This is in agreement with recent molecular evidence that Ibla (Ibliformes) is the sister taxon to all other Thoracica and the ibliforms therefore become the outgroup of choice for studying character evolution within the superorder. Paralepas, and other pedunculated barnacles without shell plates, are apparently not primitive but are secondarily evolved and nested within the Thoracica. J. Morphol., 2009. © 2008 Wiley-Liss, Inc.

Thoracic cirripedes are sessile crustaceans that use six pairs of thoracic appendages (the cirri) to catch and handle food. We used scanning electron microscopy to examine the cirri, which include one to three pairs of maxillipeds in six species of thoracican barnacles, in search of correlations between cirral setation and feeding mode. The species studied comprise both pedunculate and sessile forms and represent a wide range of marine habitats as well as morphologies, viz., Ibla cumingi, Octolasmis warwickii, Capitulum mitella, Pollicipes polymerus, Tetraclita japonica japonica and Megabalanus volcano. Of the pedunculates, I. cumingi has the least complex setation pattern consisting of only serrulate types. This is consistent with its very simplified feeding mode and an apparent inability to discriminate between food items. Octolasmis warwickii is slightly more modified, while both P. polymerus and C. mitella have a more diversified setation. The balanomorphan species exhibit by far the most complex cirral setation. This is consistent with the several types of suspension feeding seen in these species, their ability to identify and sort captured food items and even to perform microfiltration in the mantle cavity using the setae on their three pairs of maxillipeds. Our results indicate that in thoracican barnacles, adaptations in feeding behaviour are associated with changes in the setation pattern of the cirri. In addition, the setal types and their distribution on the cirri are potential new characters in future morphology-based analyses of the phylogeny of the Cirripedia Thoracica.

We describe the external morphology of the two cypridiform larval instars (first and second ascothoracid-larvae, or “a-cyprids”) of the ascothoracidan genus Dendrogaster . Ascothoracid-larvae of five species were studied with light and scanning electron microscopy, including both ascothoracid-larval instars in Dendrogaster orientalis Wagin. The first and second instars of the ascothoracid-larvae differ in almost all external features. The carapace of instar 1 has a smooth surface and lacks pores, setae, and lattice organs, while instar 2 has all these structures. The antennules of the first instar have only a rudimentary armament, the labrum does not encircle the maxillae, thoracopods 2–3 are not armed with a plumose coxal seta, and the abdomen is four-segmented (versus five-segmented in instar 2). Thus, the first ascothoracid-larva of Dendrogaster represents a transitional, generally brooded stage between the naupliar stages and the dispersive and fully functional second ascothoracid-larva that accomplishes settlement. The presence of two instars of ascothoracid-larvae (a-cyprids) in members of the order Dendrogastrida differs from the single cypridiform instar found in the Cirripedia (cyprid) and Facetotecta (y-cyprid), and we discuss the evolutionary significance of these ontogenies. We found lattice organs in both the second ascothoracid-larvae and in adult males of Dendrogaster . We could not observe both ascothoracid-larvae and males in any single species, but our data suggests that the lattice organs change significantly at the molt between these two instars. The lattice organs of second ascothoracid-larvae have no distinct keel and are situated in wide, shallow pits, whereas they have the ground pattern “crest-in-a-trough” morphology in adult males of two additional species examined for comparison. The positions of the terminal pores of lattice organs 1 and 2 also seem to change during maturation. These findings show that comparative data on lattice organ morphology for phylogenetic purposes must derive from strictly homologous instars, viz., the second ascothoracid-larva (a-cyprid) of the Ascothoracida, the y-cyprid of the Facetotecta, and the cyprid of Cirripedia. The ascothoracid-larvae of Dendrogaster and those of the family Ascothoracidae have four pairs of lattice organs, which suggests that this genus and family form a monophylum, to the exclusion of Ulophysema , which then brings into question the monophyly of the Dendrogastridae. Ulophysema is currently placed in the Dendrogastridae, but its second ascothoracid-larva has lattice organs of different and more plesiomorphic number and morphology. We briefly review lattice organ morphology across the Thecostraca. These organs are normally considered structures of the cypridiform larva and their presence in adult (males) Ascothoracida is unique in the Thecostraca. The continued morphological modification of these sensory structures in males compared to ascothoracid-larvae may suggest that they originated in adult thecostracans, but have come to be functional in the cypridiform larvae as well.