Surveying keratose sponges (Porifera, demospongiae, Dictyoceratida) reveals hidden diversity of host specialist barnacles (Crustacea, Cirripedia, Balanidae).

Sponges represent one of the most species-rich hosts for commensal barnacles yet host utilisation and diversity have not been thoroughly examined. This study investigated the diversity and phylogenetic relationships of sponge-inhabiting barnacles within a single, targeted host group, primarily from Western Australian waters. Specimens of the sponge order Dictyoceratida were surveyed and a total of 64 host morphospecies, representing four families, were identified as barnacle hosts during the study. Utilising molecular (COI, 12S) and morphological methods 42 molecular operational taxonomic units (MOTUs) of barnacles, representing Acasta, Archiacasta, Euacasta and Neoacasta were identified. Comparing inter- and intra-MOTU genetic distances showed a barcode gap between 2.5% and 5% for COI, but between 1% and 1.5% in the 12S dataset, thus demonstrating COI as a more reliable barcoding region. These sponge-inhabiting barnacles were demonstrated to show high levels of host specificity with the majority being found in a single sponge species (74%), a single genus (83%) or a single host family (93%). Phylogenetic relationships among the barnacles were reconstructed using mitochondrial (12S, COI) and nuclear (H3, 28S) markers. None of the barnacle genera were recovered as monophyletic. Euacasta was paraphyletic in relation to the remaining Acastinae genera, which were polyphyletic. Six well-supported clades of molecular operational taxonomic units, herein considered to represent species complexes, were recovered, but relationships between them were not well supported. These complexes showed differing patterns of host usage, though most were phylogenetically conserved with sister lineages typically occupying related hosts within the same genus or family of sponge. The results show that host specialists are predominant, and the dynamics of host usage have played a significant role in the evolutionary history of the Acastinae.

Comparative transcriptomic analysis of deep- and shallow-water barnacle species (Cirripedia, Poecilasmatidae) provides insights into deep-sea adaptation of sessile crustaceans.

BACKGROUND: Barnacles are specialized marine organisms that differ from other crustaceans in possession of a calcareous shell, which is attached to submerged surfaces. Barnacles have a wide distribution, mostly in the intertidal zone and shallow waters, but a few species inhabit the deep-sea floor. It is of interest to investigate how such sessile crustaceans became adapted to extreme deep-sea environments. We sequenced the transcriptomes of a deep-sea barnacle, Glyptelasma gigas collected at a depth of 731 m from the northern area of the Zhongjiannan Basin, and a shallow-water coordinal relative, Octolasmis warwicki. The purpose of this study was to provide genetic resources for investigating adaptation mechanisms of deep-sea barnacles. RESULTS: Totals of 62,470 and 51,585 unigenes were assembled for G. gigas and O. warwicki, respectively, and functional annotation of these unigenes was made using public databases. Comparison of the protein-coding genes between the deep- and shallow-water barnacles, and with those of four other shallow-water crustaceans, revealed 26 gene families that had experienced significant expansion in G. gigas. Functional annotation showed that these expanded genes were predominately related to DNA repair, signal transduction and carbohydrate metabolism. Base substitution analysis on the 11,611 single-copy orthologs between G. gigas and O. warwicki indicated that 25 of them were distinctly positive selected in the deep-sea barnacle, including genes related to transcription, DNA repair, ligand binding, ion channels and energy metabolism, potentially indicating their importance for survival of G. gigas in the deep-sea environment. CONCLUSIONS: The barnacle G. gigas has adopted strategies of expansion of specific gene families and of positive selection of key genes to counteract the negative effects of high hydrostatic pressure, hypoxia, low temperature and food limitation on the deep-sea floor. These expanded gene families and genes under positive selection would tend to enhance the capacities of G. gigas for signal transduction, genetic information processing and energy metabolism, and facilitate networks for perceiving and responding physiologically to the environmental conditions in deep-sea habitats. In short, our results provide genomic evidence relating to deep-sea adaptation of G. gigas, which provide a basis for further biological studies of sessile crustaceans in the deep sea.

Comparative Transcriptomic Analysis Reveals Candidate Genes and Pathways Involved in Larval Settlement of the Barnacle Megabalanus volcano.

Megabalanus barnacle is one of the model organisms for marine biofouling research. However, further elucidation of molecular mechanisms underlying larval settlement has been hindered due to the lack of genomic information thus far. In the present study, cDNA libraries were constructed for cyprids, the key stage for larval settlement, and adults of Megabalanus volcano. After high-throughput sequencing and de novo assembly, 42,620 unigenes were obtained with a N50 value of 1532 bp. These unigenes were annotated by blasting against the NCBI non-redundant (nr), Swiss-Prot, Cluster of Orthologous Groups (COG), and Kyoto Encyclopedia of Genes and Genomes (KEGG) databases. Finally, 19,522, 15,691, 14,459, and 10,914 unigenes were identified correspondingly. There were 22,158 differentially expressed genes (DEGs) identified between two stages. Compared with the cyprid stage, 8241 unigenes were down-regulated and 13,917 unigenes were up-regulated at the adult stage. The neuroactive ligand-receptor interaction pathway (ko04080) was significantly enriched by KEGG enrichment analysis of the DEGs, suggesting that it possibly involved in larval settlement. Potential functions of three conserved allatostatin neuropeptide-receptor pairs and two light-sensitive opsin proteins were further characterized, indicating that they might regulate attachment and metamorphosis at cyprid stage. These results provided a deeper insight into the molecular mechanisms underlying larval settlement of barnacles.

Darwin's "Mr. Arthrobalanus": Sexual Differentiation, Evolutionary Destiny and the Expert Eye of the Beholder.

Darwin's Cirripedia project was an exacting exercise in systematics, as well as an encrypted study of evolution in action. Darwin had a long-standing interest and expertise in marine invertebrates and their sexual arrangements. The surprising and revealing sexual differentiation he would uncover amongst barnacles represented an important step in his understanding of the origins of sexual reproduction. But it would prove difficult to reconcile these findings with his later theorizing. Moreover, the road to discovery was hardly straightforward. Darwin was both helped and hindered by the tacit expectations generated by his transformist theorizing, and had to overcome culturally-embedded assumptions about gender and reproductive roles. Significant observational backtracking was required to correct several oversights and misapprehensions, none more so than those relating to the chronically misunderstood "Mr. Arthrobalanus." With careful attention to chronology, this paper highlights some curious and overlooked aspects of Darwin's epic project.

Phylogenetic relationships of Darwin's "Mr. Arthrobalanus": The burrowing barnacles (Cirripedia: Acrothoracica).

The barnacles of the superorder Acrothoracica are small, burrowing, epibiotic, and dioecious (large female with dwarf male) crustaceans largely found in the carbonate sediments and skeletons of marine invertebrates. The acrothoracicans represent the Cirripedia with the most plesiomorphic characters and have prominently featured in phylogenetic speculations concerning these crustaceans. Traditionally, Acrothoracica was divided into two main orders, Pygophora and Apygophora. The Apygophora had uniramus cirri and no anus. The Pygophora had biramus terminal cirri and an anus and was further divided into two families, Lithoglyptidae and Cryptophialidae. Kolbasov (2009) revised the superorder Acrothoracica on the basis of morphological examinations of females, dwarf males, and cyprids and rearranged the acrothoracican species into two new orders, Lithoglyptida and Cryptophialida. The present study is the first attempt to reconstruct the phylogenetic relationships of acrothoracican barnacles by sequencing two mitochondrial (cytochrome C oxidase I and 16S ribosomal DNA) and two nuclear (18S ribosomal DNA and histone H3) markers of 8 of the 11 genera comprising 23 acrothoracican species. All monophylies of the eight acrothoracican genera sampled in this study were strongly supported. The deep interfamilial relationship constructed is consistent with the recent morphological phylogenetic relationship proposed by Kolbasov, Newman, and Høeg (Kolbasov, 2009) that Cryptophialidae (order Cryptophialida) is the sister group to all other acrothoracicans (order Lithoglyptida). According to an ancestral character state reconstruction analysis, the posterior lobes of females; armament of opercular bars, attachment stalk, lateral projections of the body, and aperture slits in dwarf males; and habitat use appear to have phylogenetic importance.

Evolutionary and biogeographical patterns of barnacles from deep-sea hydrothermal vents.

The characterization of evolutionary and biogeographical patterns is of fundamental importance to identify factors driving biodiversity. Due to their widespread but discontinuous distribution, deep-sea hydrothermal vent barnacles represent an excellent model for testing biogeographical hypotheses regarding the origin, dispersal and diversity of modern vent fauna. Here, we characterize the global genetic diversity of vent barnacles to infer their time of radiation, place of origin, mode of dispersal and diversification. Our approach was to target a suite of multiple loci in samples representing seven of the eight described genera. We also performed restriction-site associated DNA sequencing on individuals from each species. Phylogenetic inferences and topology hypothesis tests indicate that vent barnacles have colonized deep-sea hydrothermal vents at least twice in history. Consistent with preliminary estimates, we find a likely radiation of barnacles in vent ecosystems during the Cenozoic. Our analyses suggest that the western Pacific was the place of origin of the major vent barnacle lineage, followed by circumglobal colonization eastwards through the Southern Hemisphere during the Neogene. The inferred time of radiation rejects the classic hypotheses of antiquity of vent taxa. The timing and the mode of origin, radiation and dispersal are consistent with recent inferences made for other deep-sea taxa, including nonvent species, and are correlated with the occurrence of major geological events and mass extinctions. Thus, we suggest that the geological processes and dispersal mechanisms discussed here can explain the current distribution patterns of many other marine taxa and have played an important role shaping deep-sea faunal diversity. These results also constitute the critical baseline data with which to assess potential effects of anthropogenic disturbances on deep-sea ecosystems.

The origins and evolution of dwarf males and habitat use in thoracican barnacles.

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.

Molecular phylogeny of the acorn barnacle family Tetraclitidae (Cirripedia: Balanomorpha: Tetraclitoidea): validity of shell morphology and arthropodal characteristics in the systematics of Tetraclitid barnacles.

Shell structure is a crucial aspect of barnacle systematics. Within Tetraclitidae, the diametric and monometric growth patterns and number of rows of parietal tubes in the shells are key characteristics used to infer evolutionary trends. We used molecular analysis based on seven genes (mitochondrial COI, 16S and 12S rRNA, and nuclear EF1, RPII, H3, and 18S rRNA) to test two traditional phylogenetic hypothesis: (1) Tetraclitid barnacles are divided into two major lineages, which are distinguished according to monometric and diametric shell growth patterns, and (2) the evolutionary trend in shell parietal development began with a solid shell, which developed into a single tubiferous shell, which then developed into multitubiferous shells. The results indicated that Tetraclitinae and Newmanellinae are not monophyletic, but that Austrobalaninae and Tetraclitellinae are. The phylogram based on the genetic data suggested that Bathylasmatidae is nested within the Tetraclitidae, forming a sister relationship with the Austrobalaninae and Tetraclitinae/Newmanellinae clade. Within the Tetraclitinae/Newmanellinae clade, the genera Tetraclita (multitubiferous shell), Tesseropora (single tubiferous shell), and Yamaguchiella (multitubiferous shell) are polyphyletic. The results suggested that shell morphology and growth patterns do not reflect the evolutionary history of Tetraclitidae, whereas the arthropodal characteristics are informative.

A range extension of a deep-sea barnacle of the genus Aurivillialepas (Cirripedia, Scalpellomorpha), a Macaronesian and amphitropical refugial genus having Mesozoic affinities.

The scalpellomorph barnacle, Aurivillialepas calycula (Aurivillius, 1898), previously known only from Macaronesia, is reported from Banco de Galicia, off the NW corner of the Iberian Peninsula. One of the two specimens was attached to the scleractinian coral, Madrepora oculata Linnaeus, 1758. Since such pedunculate barnacles are little known, the potentially hermaphroditic specimens and its complemental male are illustrated photographically, and a key to the genus Aurivillialepas is provided. The genus, together with Scillaelepas Seguenza, 1876 and Gruvelialepas Newman, 1980, has long been considered to constitute a natural group of scalpellomorphs within the Calanticidae, and therefore the Scillaelepadinae subfam. nov. is proposed to accommodate them. Biogeographical aspects of these deep-sea barnacles support the hypothesis that not only the islands but the banks and guyots of Macaronesia constitute refugia for ancient as well as more recent forms, some of which may stem back to the late Mesozoic.

Molecular phylogeny, systematics and morphological evolution of the acorn barnacles (Thoracica: Sessilia: Balanomorpha).

The Balanomorpha are the largest group of barnacles and rank among the most diverse, commonly encountered and ecologically important marine crustaceans in the world. Paradoxically, despite their relevance and extensive study for over 150years, their evolutionary relationships are still unresolved. Classical morphological systematics was often based on non-cladistic approaches, while modern phylogenetic studies suffer from severe undersampling of taxa and characters (both molecular and morphological). Here we present a phylogenetic analysis of the familial relationships within the Balanomorpha. We estimate divergence times and examine morphological diversity based on five genes, 156 specimens, 10 fossil calibrations, and six key morphological characters. Two balanomorphan superfamilies, eight families and twelve genera were identified as polyphyletic. Chthamaloids, chionelasmatoid and pachylasmatoids split first from the pedunculated ancestors followed by a clade of tetraclitoids and coronuloids, and most of the balanoids. The Balanomorpha split from the Verrucidae (outgroup) in the Lower Cretaceous (139.6 Mya) with all the main lineages, except Pachylasmatoidea, having emerged by the Paleocene (60.9 Mya). Various degrees of convergence were observed in all the assessed morphological characters except the maxillipeds, which suggests that classical interpretations of balanomorphan morphological evolution need to be revised and reinterpreted.

Characterization of microsatellite loci and repeat density in the gooseneck barnacle, Pollicipes elegans, using next generation sequencing.

Pollicipes elegans is a commercially important and biogeographically significant rocky-shore gooseneck barnacle found along the eastern Pacific coasts of Peru, El Salvador, and Mexico. Little is known about its reproductive biology, and no genetic resources exist despite its growing importance as a fisheries species in the region. Next generation sequencing methods can provide rapid and cost-effective development of molecular markers such as microsatellites, which can be applied to studies of paternity, parentage, and population structure in this understudied species. Here, we used Roche 454 pyrosequencing to develop microsatellite markers in P. elegans and made genomic comparisons of repeat density and repeat class frequency with other arthropods and more distantly related taxa. We identified 13 809 repeats of 1-6 bp, or a density of 9744 bp of repeat per megabase queried, which was intermediate in the range of taxonomic groups compared. Comparison of repeat class frequency distributions revealed that P. elegans was most similar to Drosophila melanogaster rather than the more closely related crustacean Daphnia pulex. We successfully isolated 15 polymorphic markers with an average of 9.4 alleles per locus and average observed and expected heterozygosities of 0.501 and 0.597, respectively. Four loci were found to be out of Hardy-Weinberg equilibrium, likely due to the presence of null alleles. A preliminary population genetic analysis revealed low but significant differentiation between a Peruvian (n = 47) and Mexican (n = 48) population (F(ST) = 0.039) and markedly reduced genetic diversity in Peru. These markers should facilitate future studies of paternity, parentage, and population structure in this species.

A checklist of the barnacles (Crustacea: Cirripedia: Thoracica) of the Persian Gulf and Gulf of Oman with nine new records.

The present annotated checklist contains 43 species of thoracican barnacles known to date from the area, 33 and 26 from the Persian Gulf and the Gulf of Oman, respectively. Nine species are new records for the area including Amphibalunus subalbidus (Henry, 1973), Armatobalanus allium (Darwin, 1854), Chelonibia patula (Ranzani, 1818), Conchoderma hunteri (Owen, 1830), Lepas anserifera Linnaeus, 1767, Lithotrya valentiana Reinhardt, 1850, Megabalanus coccopoma (Darwin, 1854), Megabalanus occator (Darwin, 1854) and Platylepas hexastylos (Fabricius, 1798), of which A. subalbidus and M. coccopoma are reported as alien species from the region.

Phylogenetic position and evolutionary history of the turtle and whale barnacles (Cirripedia: Balanomorpha: Coronuloidea).

Barnacles of the superfamily Coronuloidea are obligate epibionts of various marine mammals, marine reptiles and large crustaceans. We used five molecular markers: 12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA and Histone 3 to infer phylogenetic relationships among sixteen coronuloids, representing most of the recent genera of barnacles of this superfamily. Our analyses confirm the monophyly of Coronuloidea and that this superfamily and Tetraclitoidea are sister groups. The six-plated Austrobalanus clusters with these two superfamilies. Based on BEAST and ML trees, Austrobalanus is basal and sister to the Coronuloidea, but the NJ tree places Austrobalanus within the Tetraclitoidae, and in the MP tree it is sister to both Coronuloidea and Tetraclitoidae. Hence the position of Austrobalanus remains unresolved. Within the Coronuloidea we identified four clades. Chelonibia occupies a basal position within the Coronuloidea which is in agreement with previous studies. The grouping of the other clades does not conform to previous studies. Divergence time analyses show that some of the time estimates are congruent with the fossil record while some others are older, suggesting the possibility of gaps in the fossil record.

Morphometric and molecular identification of individual barnacle cyprids from wild plankton: an approach to detecting fouling and invasive barnacle species.

The present study used DNA barcodes to identify individual cyprids to species. This enables accurate quantification of larvae of potential fouling species in the plankton. In addition, it explains the settlement patterns of barnacles and serves as an early warning system of unwanted immigrant species. Sequences from a total of 540 individual cypris larvae from Taiwanese waters formed 36 monophyletic clades (species) in a phylogenetic tree. Of these clades, 26 were identified to species, but 10 unknown monophyletic clades represented non-native species. Cyprids of the invasive barnacle, Megabalanus cocopoma, were identified. Multivariate analysis of antennular morphometric characters revealed three significant clusters in a nMDS plot, viz. a bell-shaped attachment organ (most species), a shoe-shaped attachment organ (some species), and a spear-shaped attachment organ (coral barnacles only). These differences in attachment organ structure indicate that antennular structures interact directly with the diverse substrata involved in cirripede settlement.

Trevathana noae, a new species of coral inhabiting barnacle (Cirripedia: Thoracica: Pyrgomatidae).

We describe a new species from the genus Trevathana Anderson, 1992, collected from the Cocos/Keeling Islands in the Indian Ocean. Trevathana noae sp. nov. is similar to other species of Trevathana by its external shell and opercular valve morphology. It is distinct from congeners in that the tergum has a distinct spur which lacks an internal tooth in adult specimens. A key to the known species of Trevathana is given. 

Two new species of scalpelliform barnacles (Cirripedia: Thoracica) from the Plio-Pleistocene of Cotentin, northwest France.

Two new species of pedunculate cirripede are described from the Plio-Pleistocene of Cotentin, northwest France. Scalpellum carentanensis sp. nov. is only the third species of this genus to be recorded from Pliocene strata in Europe. It is characterised by a triangular tergum with an apico-basal fold, without an outward extension, the inner surface of which has the raised part marked by wide growth lines, cut by a groove; the scutum has a ridge between the umbo and the tergo-lateral angle, lacks an apico-basal ridge, and possesses a line formed by the convergence of the growth zones; the length of the upper lateral exceeds its width by about 1.25 times, the umbo being situated at one-fifth the length of the plate from the apex. Arcoscalpelluin concavitectum sp. nov. is the second species of this genus on record from the Pliocene of Europe. It possesses a carina with a concave tectum bordered on each side by a smooth rounded rib; the tergum has an apico-basal line formed by the convergence of growth zones; and the upper lateral has the basi-scutal angle more widely truncated than the basi-tergal angle.

Vulcanolepas scotiaensis sp. nov., a new deep-sea scalpelliform barnacle (Eolepadidae: Neolepadinae) from hydrothermal vents in the Scotia Sea, Antarctica.

A new deep-sea stalked barnacle, Vulcanolepas scotiaensis sp. nov. is described from hydrothermal vents at depths of 2400-2600 metres along segments of the East Scotia Ridge and from 1400 metres in the Kemp Caldera. Both locations are areas of volcanic activity that lie on the Antarctic-South American Ocean Ridge complex near the South Sandwich Islands. This discovery confirms a wide distribution in southern seas for Vulcanolepas, complementing the previous records from deep-sea vents in the Lau Basin and Kermadec Ridge in the southwest Pacific, and the Pacific Antarctic Ridge in the southeast Pacific. V. scotiaensis sp. nov., the third described species of Vulcanolepas shows an extraordinary range in morphology, requiring a reassessment of the original diagnosis for Vulcanolepas. Although the morphological envelope of V. scotiaensis sp. nov. includes representatives with a peduncle to capitulum ratio similar to that observed in most neolepadines, the peduncle generally shows greater proportional length than in species in any neolepadine genus except Leucolepas; it is distinguished from other species of Vulcanolepas by a broader capitulum, much smaller imbricating scales on the peduncle and more ornamented capitulum plates. The morphological diversity of V. scotiaensis sp. nov. is interpreted as having arisen due to abrupt changes in water temperature.LSID: urn:lsid:zoobank.org:act:AA2AFDA5-0B08-466A-A584-D3FDBDE9DA61.

Acrothoracican barnacles (Lithoglyptida) in Taiwan, including the taxonomic status of Balanodytes taiwanus Utinomi, 1950 and cryptic diversity of Auritoglyptes bicornis (Aurivillius, 1892).

We list five acrothoracican barnacles of the order Lithoglyptida currently found in Taiwanese waters, including two new records, Trypetesa habei Utinomi, 1962 and Berndtiapurpurea Utinomi, 1950 and a new undescribed species of the genus Lithoglyptes. We also investigate the morphology and molecular genetics of Balanodytes taiwanus Utinomi, from its type locality, Taiwan. The original description of B. taiwanus in Utinomi (1950a) reported the absence of caudal appendages. Re-examination of the somatic body of one of the syntype specimens (the other syntype only having an opercular bar remaining) of B. taiwanus deposited in the Seto Marine Laboratory in Japan, revealed the presence of caudal appendages. The morphology of our specimens collected in various locations in Taiwan fits the description in Utinomi (1950a) and all have caudal appendages. The diagnosis of Balanodytes Utinomi, 1950 is revised herein. The genus Armatoglyptes Kolbasov & Newman, 2005 is a nomen nudum and ajunior synonym of Balanodytes. Our results also show that the widespread species Lithoglyptes habei Tomlinson, 1963 is a junior synonym of B. taiwanensis. The molecular sequence divergencse of the 16S RNA and COI genes were studied for samples of another monotypic genus Auritoglyptes (A. bicornis) from different regions of Taiwan. It was shown that there are at least three phylogenetic clades in Taiwan, suggesting that Auritoglyptes represents a cryptic species complex.

Genealogical approaches to the temporal origins of the Central American Gap: speciation and divergence in pacific Chthamalus (Sessilia: Chthamalidae).

A large section of the tropical Eastern Pacific coastline is nearly devoid of reef or consolidated habitat, and is known as the Central American Gap as it is associated with a biogeographic transition in fish and invertebrate species. We analyze phylogeographic data for intertidal barnacles (Chthamalus) to identify relevant temporal patterns that describe the origins of this biogeographic transition (the Mexican-Panamic Transition Zone). These contrasts of populations on either side of the transition zone include two pairs of closely related species (C. panamensis and C. hedgecocki; C. southwardorum and a Southern form of C. southwardorum), as well as gene flow data within one species (C. panamensis) that currently is found on both sides of the boundary between provinces. Using sequence data from a prior phylogenetic study, we used traditional (net nucleotide divergence) measures as well as coalescent analyses that incorporate the isolation-migration model to identify the likely time of separation between Northern and Southern taxa in two species pairs. A total of 67 individuals were sequenced at two mitochondrial (cytochrome c oxidase I, 16S) and one nuclear (elongation factor 1-alpha) gene regions. Our analyses indicate that the regional isolation of these intertidal barnacles occurred approximately 315-400kya, with subsequent expansion of C. panamensis from the Southern region into the North much more recently. There are insufficient survey data to conclusively document the absence of species from this group within the Central American Gap region near the Gulf of Tehuantepec. However, appropriate habitat is quite sparse in this region and other environmental factors, including upwelling and water temperature, are likely to be associated with isolation of many species in the Mexican and Panamic provinces sensu stricto. Some taxa may maintain gene flow across this region, but very few genetic studies have been completed on such taxa. Until further work is done, distinguishing between prior hypotheses of a faunal gap, or a faunal transition zone, is somewhat speculative. Additional taxonomic revision will be necessary in Chthamalus but is beyond the scope of this paper.

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.