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A new species of chionelasmatid sessile vent barnacle, Eochionelasmus coreana sp. nov., is described and illustrated on the basis of specimens collected from the Solitaire hydrothermal vent field in the Central Indian Ridge of the Indian Ocean. This new species is morphologically very similar to E. ohtai, the type species of the genus Eochionelasmus. However, it differs from E. ohtai in its distribution, the status of the notch on the maxillule, and the positions of rl1 and cl1 on whorls of the imbricating plates. In addition, a molecular phylogenetic tree indicated that the chionelasmatid Eochionelasmus was closely related to the waikalasmatid Waikalasma with high supporting values rather than the other chionelasmatid Chionelasmus. The new species is not only the first record of a sessile vent barnacle from outside of the Pacific Ocean, but is also the first sessile barnacle from the Indian Ocean.

Octomerisis a chthamalid intertidal barnacle with eight shell plates. There are currently two species of such barnacles:O. brunneaDarwin, 1854 (type locality in the Philippines), common in the Indo-Pacific region, andO. angulosaSowerby, 1825, only recorded in South Africa.Octomeris intermediaNilsson-Cantell, 1921, identified from the Mergui Archipelago in Myanmar, was considered to be conspecific withO. brunneaby Hiro (1939) based on samples collected in Taiwan. The morphological differences in shell and opercular plates betweenO. brunneaandO. intermediaare believed to be intra-specific variations due to different degrees of shell erosion. In the present study, the genetic and morphological differentiations ofOctomerisin the Indo-Pacific region were examined. This study found two molecular clades (with inter-specific differences) based on the divergence in the COI genes, and the species also have distinct geographical distributions. TheOctomeris brunneaclade covers samples collected from the Philippines and Taiwan waters and the other clade, which we argue isO. intermedia, is distributed in Phuket and Krabi, Thailand and Langkawi, Malaysia. Phuket and Krabi are located approximately 300 km south of the Mergui Archipelago, the type locality ofO. intermedia. The morphology of samples collected from Thailand fits the type description ofO. intermediain Nilsson-Cantell (1921). Our study concludes thatO. intermediais a valid species based on morphological and molecular evidence.

In the present study, the Indo-Pacific coral associated barnacle Cantellius euspinulosum (Broch, 1931) was found to have cryptic species in Korea, Taiwan and other regions based on molecular studies. However, the original specimens of C. euspinulosum from Broch have not been previously described or illustrated, making it difficult to assign which cryptic species to the original C. euspinulosum. The original specimen of C. euspinulosum was examined and illustrated here, and the species identity of C. cf. euspinulosum collected from Jejudo Island in the present study and other cryptic species (based on literature illustrations) in the Indo-Pacific were evaluated.C. euspinulosum from Singapore, Java, Mergui Archipelago in Andaman Sea and Nha Trang represented the C. euspinulosum identified by Broch (1931). It is a generalist on Acropora, Favia, Favites, Leptoria, Montipora, Pachyseris and Pocillipora corals and distributed in the Indo-Pacific region. Morphological examination and DNA sequencing (COI, 12S DNA sequences) in the present study showed that C. cf. euspinulosum from Jejudo Island, Korea represents a distinct species, herein named C. alveoporae sp. nov. Cantellius alveroporae sp. nov. is a specialist species that only grows on Alveopora and also present in Palau, and Ogasawara Island in Japan. Cantellius cf. euspinuloum in Taiwan, the Moscos Island, and Australia belong to several other distinct species awaiting further morphological and molecular studies. At least five cryptic species of C. euspinulosum were identified in the present study, including both specialist and generalists.

A new species of coral associated barnacle (Balanomorpha: Pyrgomatidae) sharing morphological features of Darwiniella (Anderson, 1992) and Galkinius Perreault, 2014 is described. It has a fused shell and opercular plates, characteristic of Darwiniella. However, the morphology of the tergum and somatic body are closer to Galkinius. Sequence divergence of mitochondrial DNA 12S rDNA and COI reveals this new species clusters with the Galkinius clade. Therefore this new form is assigned to the genus Galkinius, as G. maculosus sp. n. Concomitantly the diagnosis of Galkinius is emended to include species with fused or four- plated shells and fused opercular plates. The new species is distinct from all Galkinius species in having a fused shell. It inhabits the corals Lobophyllia spp. and is distributed from the Dongsha Atoll in the South China Sea, Orchid Island of Taiwan in the Pacific Ocean, to Madang in Papua New Guinea waters.

The morphology and ultrastructure of the shells of two balanid species have been examined, paying special attention to the three types of boundaries between plates: (i) radii-parietes, (ii) alae-sheaths, and (iii) parietes-basal plate. At the carinal surfaces of the radii and at the rostral surfaces of the alae, there are series of crenulations with dendritic edges. The crenulations of the radius margins interlock with less prominent features of the opposing paries margins, whereas the surfaces of the longitudinal abutments opposing the ala margins are particularly smooth. The primary septa of the parietes also develop dendritic edges, which abut the internal surfaces of the primary tubes of the base plates. In all cases, there are chitino-proteinaceous organic membranes between the abutting structures. Our observations indicate that the very edges of the crenulations and the primary septa are permanently in contact with the organic membranes. We conclude that, when a new growth increment is going to be produced, the edges of both the crenulations and the primary septa pull the viscoelastic organic membranes locally, with the consequent formation of viscous fingers. For the abutting edges to grow, calcium carbonate must diffuse across the organic membranes, but it is not clear how growth of the organic membranes themselves is accomplished, in the absence of any cellular tissue., This research was funded by projects CGL2017-85118-P (A.G.C., C.S. and C.G.) and CGL2015-64683-P (A.B.R.N.) of the Spanish Ministerio de Economía, Industria y Competitividad, the Unidad Científica de Excelencia UCE-PP2016-05 of the University of Granada (A.G.C. and A.B.R.N.) and the Research Group RNM363 of the Junta de Andalucía (A.G.C.). N.A.L. acknowledges support from CONICYT-Chile through grant nos. FONDECYT 1140938, PCI REDES 170106 and PIA ANILLOS ACT172037, for international collaborative research with A.G.C. and A.B.R.N.

An extraordinarily diverse assemblage of cirripedes is described from a shallow-water deposit of late Oligocene age from Cosy Dell farm, near Waimumu, Southland, New Zealand. It is unusual not only because it represents a rarely preserved intertidal to shallow subtidal fauna, but also because it contains at least nine species, five of which are new to science. The deposit contains the earliest known representatives of the genera Chamaesipho and Notobalanus and these are associated with abundant Verruca, an association that no longer occurs in present-day New Zealand waters. Although the remains are intimately associated with each other, the deposit is interpreted as a condensed heterogeneous taphocoenosis, with intertidal zone taxa (Verruca, Chamaesipho, Tetraclitella, Austrobalanus, Protelminius) mixing with upper subtidal species (Eolasma, Notobalanus, Tasmanobalanus) and pelagic cirripedes (Lepas).urn:lsid:zoobank.org:pub:74EECF88-AF4A-4EA7-BF7B-7487D0E35D5Durn:lsid:zoobank.org:act:8D81F5AD-3861-48A5-A...

The family Catophragmidae is one of the lower balanomorphs from traditional and recent multiple mitochondrial and nuclear markers molecular analysis. Here, we characterized the first mitogenome of ...

In an ecologically oriented paper, Maruzzo et al. (2012) illustrate six developmental phases from the freshly settled cyprid larvae to the sessile juvenile in the barnacle, Amphibalanus amphitrite (Darwin, 1854), and in an evolutionarily oriented paper Hoeg et al. (2012) do likewise for Megabalanus rosa (Pilsbry, 1916). Both video-based studies illustrate 5 unadorned increasingly sac-like phases leading up to the sessile juvenile (Phase 6). In both species, Phase 5 is not only illustrated as sac-like, it resides outside the protective canopy of the cyprid shell, and this is reminiscent of the situation described by Bernard and Lane (1962, Fig. 24A). However, it has been long known that not only is the metamorphosing balanoid larva within the cyprid carapace divided into a capitulum and peduncle, but the anlagen of plates can seen with light microscopy through the largely transparent cyprid carapace (Runnstrom, 1925; Walley, 1969; Gusenbauer, 2003), and primordial plates appear in SEMs in Phase 5, after the carapace has been shed (Glenner and Hoeg, 1993; Gusenbauer, 2003). Maruzzo et al. and Hoeg et al. neither recognize such plates nor do they recognize the peduncle in any of the phases. Yet it is largely the peduncle that is involved in the struggle to shed the cyprid carapace in the transition to semi-sessile Phase 5, which they describe and illustrate as a “bag-like body shape.” Maruzzo et al. claim this phase persists for as much as 24 hours before becoming fully sessile (Phase 6), and they call for studies to assess its contribution to differential mortality. But Phase 5 has generally been observed to persist for less than three hours. Thus, the significantly longer duration being reported is either an artifact resulting from in vitro conditions and manipulations and/or failure to determine when the wall plates they illustrate in Phase 6 began to form. On the other hand, SEMs of Glenner and Hoeg, and more recently Gusenbauer, show that semi-sessile Phase 5 is by and large comparable to that illustrated by Runnstrom (1925) and, therefore, an unadorned sac-like form surely does not exist. Thus, Phase 5 of Maruzzo et al. and Hoeg et al., like that of Bernard and Lane (Fig. 14A), is most certainly an artifact.

The origin of the mainly sea-turtle fouling balanomorph family Chelonibiidae is still poorly documented. Aside from an Eocene erratic specimen assigned to an extinct subfamily, the extant subfamily Chelonibiinae did not appear in the fossil record before the Late Miocene. Protochelonibiinae Harzhauser & Newman subfam. nov. is here introduced as an extinct sister-group of Chelonibiinae. The subfamily is known so far only from the proto-Mediterranean and the Paratethys seas and ranged from Early Miocene to Late Pliocene. Members of the subfamily are characterized by large walls with tripartite rostra which display distinct sutures on the external surface. The tripartite rostrum, however, has evolved independently several times in the evolution of the balanomorphs and cannot be treated as synapomorphy. The subfamily comprises one new genus and two species. Protochelonibia Harzhauser & Newman gen. nov. is the type genus of Protochelonibiinae and Protochelonibia submersa Harzhauser & Newman sp. nov. is introdu...

We report new distribution and host records of the turtle barnacle Stephanolepas muricata Fischer. Ours is the first study to document this species' occurrence in Mexico and in the Atlantic Ocean (North Carolina, Georgia and Florida: USA). We also present the first record of S. muricata from olive ridley sea turtles, Lepidochelys olivacea (Eschscholtz). We discuss the physical and ecological characteristics associated with the specimens collected and present evidence suggesting the likelihood of chemical mediation between embedding coronuloid barnacles like S. muricata and host turtles. We provide the first data on the diet and fecundity of S. muricata.

Excavations at North Storr's Lake (SS-4) on San Salvador, Bahamas have yielded turtle barnacle wall plates (shell compartments) in association with skeletal remains of sea turtle and charcoal dated to ca. A.D. 932–1552. This is one of the few cases in which direct and indirect radiometric dates have been reported for archaeological sea turtle barnacles. The turtle barnacle has been identified as Chelonibia testudinaria (Linnaeus, 1758), a widespread, commensal species found on all genera of Cheloniidae in the world's oceans. Chelonibia testudinaria is the most often reported sea turtle barnacle and is commonly found on Caretta caretta (loggerhead) and Chelonia mydas (green turtle) dating as far back as the Miocene epoch. Wall plates of Chelonibia testudinaria occur in stratigraphic association with both loggerhead and green sea turtle remains at North Storr's Lake, appearing in the same archaeological strata as the sea turtle, therefore the association is clear. Sea turtle species were identified...

Two new species of the Pyrgomatid barnacle Trevathana are described: Trevathana synthesysae nov. sp., extracted from Plesiastrea versipora from the Indian Ocean Islands Reunion and Mauritius, and Trevathana isfae nov. sp. from a colony of Favia stelligera from French Polynesia, which, until recently, was terra incognita with regard to coral-inhabiting barnacles. The two new species are distinctive by their relatively broad scutum as compared to Trevathana dentatum, their prominent adductor ridge extending beyond the basal margin of the scutum, and their quadrangular tergum.

In situ fossil barnacle populations (Cirripedia, Balanomorpha) are described from the Miocene Natori Group of the Moniwa-Goishi area, northeast Japan. The Natori Group exhibits a transgressive sequence from subaerial lavas and volcaniclastics (Takadate Formation) to shoreface-shelf deposits (Moniwa and Hatatate formations) in ascending order. In the Takadate Formation, volcaniclastics locally intercalate bay deposits in which the fossil barnacle Balanus bisulcatus occurs on gravel surfaces in situ. The Takadate Formation is unconformably overlain by the basal conglomerate of the Moniwa Formation, which is interpreted as a ravinement deposit formed in an open coast. An extinct barnacle species, Concavus sendaicus, is found as gregarious clusters and solitary individuals on boulder surfaces in the basal conglomerate in situ. These features suggest that B. bisulcatus inhabited in bay environments at the early transgressive stage. On the other hand, C. sendaicus was distributed over open-coast enviro...

The thoracican cirripede, Xenobalanus globicipitis (Steenstrup, 1851), of the family Coronulidae (Balanomorpha), is known to occur on a variety of species of cetaceans throughout the world (see Rajaguru & Shantha, 1992 for a review; Van Waerebeek et al., 1993; Jefferson et al., 1995; Orams & Schuetze, 1998; Rittmaster et al., 1999; De Beneditto & Ramos, 2000; Aznar et al., 2005). Occurrence of X. globicipitis (hereafter referred to as Xenobalanus) has been reported from wild, captive, live-stranded, and/or dead individuals, and most describe the barnacles as commonly occurring on dorsal fins, pectoral flippers, and flukes. Though Xenobalanus may easily be confused with stalked barnacles belonging to the suborder Lepadomorpha because of their marked similarity in appearance, they belong to the suborder Balanomorpha (cf. Darwin, 1854; Ruppert et al., 2004). While Xenobalanus is often referred to as a parasitic organism, the relationship between the barnacle and its host is actually commensal, or phoretic, as the barnacle only attaches to the host for transportation purposes. While attached, the barnacle feeds via suspension techniques commonly utilized by other sessile barnacles (Ruppert et al., 2004). Xenobalanus is specialized in its ability to attach to cetaceans, yet the mechanism that drives its occurrence remains unclear. Though the larval stages of this species have not been described, similar barnacles undergo self-fertilization and release their larvae in the water column. These larvae develop and settle once an appropriate host is found, and life cycles of 5-6 months have been reported (Van Waerebeek et al., 1993; Fertl, 2002). A number of factors have been suggested to affect its settlement, including host swimming speed, the age or health of host individuals, upwelling events, and/or water temperature and

The presence of Solidobalanus auricoma (Cirripedia; Balanoidea) in the Red Sea at a depth of 112 m is reported, and its morphology is described. S. auricoma is a relatively deep water barnacle known from the Persian Gulf through Malaysia, southeastern Australia, and northeastern New Zealand at depths of 27 to 320 m. This is its first record from the Red Sea. The present finding of S. auricoma in the Gulf of Elat (Aqaba) extends the boundaries of its geographic distribution farther west and north.

This paper records a new sessile cirripede from a nearshore, shallow-water facies at Předboj near Prague, Czech Republic. The new taxon, Archaeochionelasmus nekvasilovae gen. et sp. nov., is the earliest known neobalanoform barnacle, and while it may have been an obligate commensal of a cephalopod or marine reptile, that it was a shore barnacle cannot be ruled out. (Zoobank registration: urn:lsid:zoobank.org:pub:177CC951-CEC1-425C-B693-46ADB83BD5D9)

This study examined the phylogeography of the barnacle Fistulobalanus albicostatus, which inhabits mangroves and estuarine shores in the West Pacific. Differentiation in the mitochondrial cytochrome c oxidase subunit I (COI) and 12S ribosomal RNA (12S) genes of 401 specimens of F. albicostatus was examined in samples from 16 locations in the West Pacific, ranging from Honshu to Southern China. Our results revealed that F. albicostatus comprises two major clades exhibiting a COI divergence ranging from 1.25% to 2.8%. Clade A demonstrated the widest distribution, ranging from Japan to China, and was divided into three subclades occurring in the South China Sea (A1), Okinawa (A2), and Honshu, Korea and Qingdao (A3). Clade B was determined to be endemic to Okinawa; i.e. two endemic lineages occur in this island. Thus, F. albicostatus resembles several inter-tidal species in having clades that are endemic to Okinawan waters. Nevertheless, in contrast to the rocky inter-tidal barnacles Tetraclita spp. and Chthamalus malayensis, F. albicostatus was not found to be separated into continental and oceanic populations, but instead is divided into northern and southern clades, probably because of the Yangtze River discharge, which limits gene flow between the northern and southern populations.

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 new mangrove barnacle, Balanus thailandicus sp. nov., is described from the Puyu Canal, Tamalung Bay, Satun, southwest Thailand. The new species has the common characteristics of the Balanus amphitrite complex and is more or less sympatric with B. reticulatus, B. amphitrite, B. patelliformis, and B. variegatus. Balanus thailandicus is most similar to B. reticulatus, but it is clearly distinguishable from the last three mentioned species of the complex in having the parietes folded or ribbed in adults but smooth in juveniles, parietal tubes solid in the upper half and interrupted by transverse septa in the lower part, the large number of articles in the anterior ramus of cirrus I, and erect hooks below the posterodistal angles of the articles of cirri III and IV.

A new deep sea balanomorph, Tetrachaelasma tasmanicum sp. nov., is described. This new record is based on numerous disarticulated plates, from dredge samples around deep‐sea seamounts on the South Tasman Rise. This is the deepest known occurrence of the Balanomorpha, and the first record of Tetrachaelasma from the Pacific realm. An overview of the biogeography, including aspects of the palaeontology of deep sea balano‐morphs is provided. A map showing station localities and a list of deep sea cirripedes recovered from the South and East Tasman Rise during the 1995 Rig Seismic cruise, are included.

Previous studies on the ultrastructure of barnacle cypris larvae suggest that cypris morphology, especially the antennules which play a key role in selecting the final substratum for settlement, may show variations between species existing in different habitats. In the present study, the cypris morphology of the barnacle Fistulobalanus (Balanus) albicostatus, which predominantly lives on trunks of mangrove trees was investigated using scanning electron microscopy and compared with that of species from other habitats that have already been described. The antennular segments II and III of F. albicostatus each bear one post-axial seta. The third segment consists of an attachment disc surrounded by a thin cuticular velum and the fourth segment exhibits four sub-terminal setae and five terminal setae. The morphology of the cypris antennules of F. albicostatus is similar to other balanomorph barnacles which inhabit hard shores, revealing that the antennular morphology does not have diagnostic variations between species from the different intertidal habitats. The morphology of the caudal rami (used in the evaluation of the substratum microtopography), however, appears to vary among barnacles living on different substrata.

Metamorphosis and early juvenile development was followed in the laboratory in Balanus amphitrite and B. improvisus (family Balanidae) and in Semibalanus balanoides and Elminius modestus (family Archaeobalanidae) from cyprid settlement until 24 h after ecdysis. Stages of development were studied in vivo and with scanning electron microscopy. Events in metamorphosis and early juvenile development are very similar in all four species and can be interpreted in terms of a highly modified moult. Notably, there was no indication whatsoever of the “amorphous decorticated settler” previously described from metamorphosis of B. amphitrite. The shape of a juvenile barnacle with cirri and incipient shell plates is apparent immediately after the shedding of the cypris carapace, and rudimentary peduncle can be distinguished below the developing wall plates. A basal row of cuticular hairs encircles the peduncle in all species except S. balanoides. These hairs seem to serve a restraining function during early development, when the juvenile barnacle is only attached by the initially secreted cyprid cement. Similarly situated hairs are also present in very young juveniles of the lepadomorph Scalpellum scalpellum. In Semibalanus balanoides nothing indicates that the rostrum originates from the fusion of two latera as previously claimed, since this plate is single as soon as it can be distinguished.

The Titan Acorn barnacle, Megabalanus coccopoma, a native of the tropical eastern Pacific, has become established in the western Atlantic (Brazil and the northern Gulf of Mexico to the Carolinas), northwestern Europe and the western Indian Ocean (Mauritius), and therefore its dispersal capabilities are well known. This study reports its introduction to Japan and confirms its occurrence in Australia. In an attempt to determine the source of this introduction, phylogeographic techniques, involving cytochrome c oxidase I sequences of various widely separate populations of M. rosa and M. volcano, were utilized. No significant genetic differentiation or haplotype patterns between widely separated populations of each of the three species were found. Lack of such differentiation indicates recent geographical isolation and thus negates a null hypothesis predicting that the occurrence of one of more of these species in Australia was natural.

The validity of the turtle barnacle genusChelolepashas been challenged on the grounds that the publication in which the genus was formally described does not meet the requirements of a published work under theInternational Code of Zoological Nomenclature. The present note informs thatChelolepasis a valid name that satisfies the provisions of the Code and thatTubicinella cheloniaeis a synonym ofChelolepas cheloniae.

The Thoracica includes the ordinary barnacles found along the sea shore and is the most diverse and well-studied superorder of Cirripedia. However, although the literature abounds with scenarios explaining the evolution of these barna- cles, very few studies have attempted to test these hypotheses in a phylogenetic context. The few attempts at phylogenetic analyses have suffered from a lack of phylogenetic signal and small numbers of taxa. We collected DNA sequences from the nuclear 18S, 28S, and histone H3 genes and the mitochondrial 12S and 16S genes (4,871 bp total) and data for 37 adult and 53 larval morphological characters from 43 taxa representing all the extant thoracican suborders (except the monospecific Brachylepadomorpha). Four Rhizocephala (highly modified parasitic barnacles) taxa and a Rhizocephala + Acrothoracica (burrowing barnacles) hypothetical ancestor were used as the outgroup for the molecular and morphological analyses, respectively. We analyzed these data separately and combined using maximum likelihood (ML) under "hill-climbing" and genetic algorithm heuristic searches, maximum parsimony procedures, and Bayesian inference coupled with Markov chain Monte Carlo techniques under mixed and homogeneous models of nucleotide substitution. The resulting phylogenetic trees answered key questions in barnacle evolution. The four-plated Iblomorpha were shown as the most primitive thoracican, and the plateless Heteralepadomorpha were placed as the sister group of the Lepadomorpha. These relationships suggest for the first time in an invertebrate that exoskeleton biomineralization may have evolved from phosphatic to calcitic. Ses- silia (nonpedunculate) barnacles were depicted as monophyletic and appear to have evolved from a stalked (pedunculate) multiplated (5+) scalpelloidlike ancestor rather than a five-plated lepadomorphan ancestor. The Balanomorpha (symmetric sessile barnacles) appear to have the following relationship: (Chthamaloidea(Coronuloidea(Tetraclitoidea, Balanoidea))). Thoracican divergence times were estimated under ML-based local clock, Bayesian, and penalized likelihood approaches using an 18S data set and three calibration points: Heteralepadomorpha = 530 million years ago (MYA), Scalpellomorpha = 340 MYA, and Verrucomorpha = 120 MYA. Estimated dates varied considerably within and between approaches depending on the calibration point. Highly parameterized local clock models that assume independent rates (r ≥ 15) for confamilial or congeneric species generated the most congruent estimates among calibrations and agreed more closely with the barnacle fossil record. Reasonable estimates were also obtained under the Bayesian procedure of Kishino et al. (2001, Mol. Biol. Evol. 18:352-361) but using multiple calibrations. Most of the dates estimated under the Bayesian procedure of Aris-Brosou and Yang (2002, Syst. Biol. 51:703-714) and the penalized likelihood method using single and/or multiple calibrations were inconsistent among calibrations and did not fit the fossil record. (Calibration point; Cirripedia; divergence times; DNA sequences; fossils; genetic algorithms; mixed models; phylogeny.) The barnacles (Crustacea: Cirripedia: Thoracica) are among the most commonly encountered marine crus- taceans in the world. They deviate from other crus- taceans in having irreversibly sessile (nonmobile) adults, a habit that has been accompanied by dramatic changes in both morphology and biology. These animals have dis- pensed with the normal crustacean scheme of growing by molting the entire exoskeleton and are instead clad in a protective armor of mineralized shell plates. They are thoracopodal filter feeders, and the majority of them are hermaphrodites (Anderson, 1994). These drastic adap- tations make barnacles excellent model organisms for evolutionary studies (Schram and Hoeg, 1995). Barnacles have attracted the attention of many promi- nent specialists, most notably Charles Darwin, whose seminal monographs (1851-1855), contain a prodigious body of information on barnacle biology, paleontol- ogy, and systematics. However, for a group that has been the focus of intense study for almost two cen- turies, there is still a pervasive lack of phylogenetic information (Schram and Hoeg, 1995). Multiple hy- potheses have been proposed concerning barnacle evo

The systematics of pyrgomatids, stemming from the early 1800’s, has traditionally been based on the number of plates making up the wall (six, four or one) and specializations in the opercular plates. A recent study ofthe related bryozobiines focused attention on detailed structural modifications of the basis, which we now find also applies to some highly derived pyrgomatids and an archaeobalanine. Reexamination of the Indonesian coral barnacle Pyrgoma kuri Hoek, 1913 has revealed previously unknown morphological features, including separable opercular plates, a true tergal spur, and a basis lined with ladder to arch-like calcareous structures covering “atrial passageways”. Thus, the present study expands our knowledge of such specializations and our understanding of the evolution and relationships of the derived pyrgomatids as well as the archaeobalanines and bryozobiines. The complex modifications of the basis found in these three groups evidently function as an avenuefor chemical mediationof growth between the barnacle and its host. Although monophyly of the archaeobalanines and bryozobiines based on such structures is possible, there is no obvious connection between them and the few derived pyrgomatids in which these features occur. There is apparently a propensity toward such modifications in archaeobalanids resulting in parallel evolution in association with distinctly different hosts (e.g., sponges, corals and bryozoans).

The coronuloid barnacle family-group names Cylindrolepadinae, Stomatolepadinae, Chelolepadinae, Cryptolepadinae and Tubicinellinae of Ross & Frick, 2007 are considered nomen nudum according to Article 8.6 of the International Code of Zoological Nomenclature, but appear in several subsequent published works and internet taxonomic databases. It is the purpose of this communication to rectify this situation. These five subfamilial names are proposed and defined herein anew, as Cylindrolepadinae subfam. nov., Stomatolepadinae subfam. nov., Chelolepadinae subfam. nov., Cryptolepadinae subfam. nov. and Tubicinellinae subfam. nov. The remaining valid family-group names within the Coronuloidea are also listed and defined herein.

A new species of coral-inhabiting barnacle, Pyrgopsella youngi, is described. It was found in a colony of the coral Symphyllia radians Milne Edwards & Haime, 1849 from Sulawesi, Indonesia. The barnacles were suspended in the coral tissue and were easily detached. A unique feature of Pyrgopsella is its membranous basis; in P. youngi the calcareous basis is reduced to a vestige by which the barnacle is attached to the coral. Pyrgopsella has been regarded as a genus of sponge-inhabiting barnacle, thus unique in the otherwise coral-associated family Pyrgomatidae, but our findings confirm that this genus too comprises coral-inhabiting barnacles. We propose a new genus, Pyrgospongia, to accommodate the sponge-inhabiting barnacle originally described as Pyrgopsella stellula Rosell, 1973. The relationships of both Pyrgopsella and Pyrgospongia within the Pyrogomatidae are discussed.

Bryozobia synaptos n. gen., n. sp., from the Grand Recif of Madagascar and the Mascarene Plateau north of Mauritius, is the only cirriped known that has evolved an intimate association with cheilostome bryozoans. This small barnacle settles on a bryozoan colony and in time overgrows a number of zooids, modifying its shell to accommodate them by developing numerous passages (atria) through its basis and wall by a process involving substrate replication. Calcareous, tubular passages formed in the basis of the barnacle grow radially, keeping pace with marginal growth of the barnacle's wall. Eventually, the openings to the zooecial tubes at the wall margin are encircled by the barnacle, thereby forming the first whorl of atria through which the bryozoan deploys its lophophores. During subsequent growth the process is repeated

The presence of embryos in the mantle cavity of A. imperator, T. purpurascens and T. rosea was monitored over two years. Tesseropora rosea showed a clearly defined breeding season in summer and early autumn. Seasonal breeding in A. imperator and T. purpurascens was less well defined, a small number of gravid individuals being present during most of the year. Breeding increased during the late autumn and early winter in A. imperator and during winter, spring and early summer in T. purpurascens. The larval stages of A. imperator, T. purpurascens and T. rosea are described from larvae reared in the laboratory. Separation of the larval stages between these species depends on fine morphological detail. Distinctive larval features permit the separation of coronuloid larvae from those of the other balanomorph superfamilies, but coronuloid nauplii resemble balanoid nauplii more than chthamaloid nauplii.

Epibionts on a nesting female loggerhead sea turtle Caretta caretta (Linnaeus, 1758; 92.1 cm straight carapace length and 70.7 cm straight carapace length) were examined on the island of Yakushima, Kagoshima, Japan (30.4156°N, 130.4394°S) on 16 June 2009 (Fig. 1a). Specimens were photographed, counted and identified at the lowest taxonomic level, and four species of barnacles were found: Chelonibia testudinaria (Linnaeus, 1758), Platylepas hexastylos (Fabricius, 1798), Stephanolepas muricata Fischer, 1886, and Fistulobalanus albicostatus (Pilsbry, 1916). The turtle barnacles C. testudinaria (n = 6), P. hexastylos (n = 42) and S. muricata (n = 1) are common on Japanese turtles (Hayashi 2012), while this is the first record of F. albicostatus (n = 4) on sea turtles. The acorn barnacle F. albicostatus is a common species in estuaries and mangroves and as a biofouling species on ships, and occurs in the waters of Japan, China, Taiwan and Korea (Newman and Ross 1976; Chan 2007). The barnacle has also been recorded in California waters as an invasive species (Bonnot 1935; Henry and McLaughlin 1975). Japanese loggerhead turtles migrate over the Pacific Ocean as developmental migration in their ontogenetic stages (Bowen et al. 1995); sea turtle migration allows F. albicostatus and possibly other organisms biofouling sea turtles to invade new habitats, as seen in a case of rapa whelk Rapana venosa (Valenciennes, 1846) recorded in Harding et al. (2011). In one example for F. albicostatus, a new geographic expansion into Norway by biofouling plastic tags on migrating birds was reported (Tottrup et al. 2010). Further studies are needed to examine the potential role of migratory animals such as sea turtles and birds as dispersal vectors of these hitchhikers.