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-Schematic plan views and transverse sections of grades of skeletal organization from the most primitive sessile barnacles (A, Brachylepadomorpha) through the asymmetrical sessile barnacles (B-H, Verrucomorpha). Of the eight verrucid genera currently recognized (Young 1998) only the four most representative ones (Altiverruca, Newmaniverruca, Metaverruca and Verruca s.s.) are depicted here. Right sides are to the viewers right and rostral ends are downward in plan views or coming out of the page in transverse sections. The transverse sections are slightly rostral of the rostro-carinal gap or suture. While there are numerous whorls of imbricating plates standing in tiers four plates high in brachylepadomorphs, and on the movable side of neoverrucids, only those of the R-C gap are labeled. Abbreviations: C, R, carina and rostrum; cl 1-4 , l 1-4 , rl 1-4 , carinolateral, lateral and rostrolateral tiers of imbricating plates respectively; cl, rl, the pair of imbricating plates in proverrucids; FS, FT, fixed scutum and tergum; hl, hinge line; L, median latus (completely lost in proverrucids and verrucids); m, myophore; MS, MT, movable scutum and tergum; sam, scutal adductor muscle; S, T, normal terga and scuta in brachylepadomorphs; dashed horizontal lines: membranous basis (transverse sections F-H after Young 1998).
Contexts in source publication
Context 1
... the imbricating plates between this primary wall and the substratum form a substantially greater proportion of the body chamber than in Neoverruca. The principal three tiers of lateral imbricating plates, standing four plates high (rl 1-4 , l 1-4 and cl [1][2][3][4] ), are fully rep- resented in these two apparently adult Imbri- caverruca (Figs 1A; 2A; 4C), as they are in brachylepadomorphs (Newman 1987: fig. 4A) and in the juveniles of Neoverruca (Newman & Hessler 1989;Newman 1989). ...
Context 2
... form a substantially greater proportion of the body chamber than in Neoverruca. The principal three tiers of lateral imbricating plates, standing four plates high (rl 1-4 , l 1-4 and cl [1][2][3][4] ), are fully rep- resented in these two apparently adult Imbri- caverruca (Figs 1A; 2A; 4C), as they are in brachylepadomorphs (Newman 1987: fig. 4A) and in the juveniles of Neoverruca (Newman & Hessler 1989;Newman ...
Context 3
... 1). Thus the question arises as to whether it represents a grade between Neoverruca and the proverrucids. The evidence from both juvenile and adult morphology sug- gests not. In short, Imbricaverruca has capitalized on the neoverrucid plan rather than making the reductions and acquiring proportions in the direction of those found in modern forms (Fig. 4). Thus the Neoverruca and Imbricaverruca plans are markedly divergent, the former's being better suited to the derivation of the proverru- cid/verrucid plan (Table 3). In Table 3, character 1 concerns the existence of fully pedunculate juveniles during the ontogeny of Neoverruca missing in the ontogeny of verru- cids, possibly in ...
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Citations
... In comparison with their deep-sea or shallow-water relatives, chemosynthetic barnacles have a relatively larger number of setae on the intermediate segments of the posterior three cirri (i.e., generally more than six pairs vs. less than six pairs), and a higher ratio of setal length to the width of the segment where the setae are rooted (generally 2.8-14.1 vs. mostly 2.0-3.5) (primary data summarized from Yamaguchi and Newman, 1990;Buckeridge, 1998;Newman, 2000;Southward & Jones, 2003;Buckeridge et al., 2013;Watanabe et al., 2021). We speculate that these features represent morphological adaptations of chemosynthetic barnacles, as the dense and long setae would enhance the volume of water currents flowing into the mantle cavity when their cirri are beating for respiration and feeding. ...
Thoracican barnacles represent a unique group that has evolved in parallel identical somatotype s (sessile, stalked and asymmetric) in both normal and chemosynthetic environments. Hydrothermal vents and methane seeps are typical extreme deep-sea chemosynthetic habitats for marine macrobenthos. Characterizing the evolutionary history and adaptive strategy of barnacles is fundamentally important for understanding their origin, speciation, and diversification. Herein, we performed a series of phylogenetic analyses focusing on the mitochondrial genomes of the main extant barnacle lineages. Phylogenetic inferences and topology tests contradict the view of the sister relationship between verrucomorphs and balanomorphs, instead revealing that pollicipedids, calanticids and balanomorphs share common ancestor. Selective pressure analyses indicate that the two barnacle lineages of chemosynthetic ecosystems exhibit similar patterns in their evolution of adaptive characters, but have diverse and specific positive substitution sites of mitogenomes. Divergence times suggest that chemosynthetic barnacles originated in the Cenozoic, coinciding with the origins of other metazoan animals in chemosynthetic habitats as well as the Paleogene mass extinction and oceanic anoxic events. It is reasonable to suppose that ecological niche vacancy, sitotaxis, gene specificity in adaptive stress responses, and the subdivision of the ecological niche contributed to the origin and diversification of barnacles in chemosynthetic ecosystems.
... Indeed, asymmetry side-by-side of the opercular plates is not unknown in barnacles. Specifically, members of the order Verrucomorpha, which inhabit deep-sea hydrothermal vents among other habitats, exhibit an asymmetric form of the scutum, with one side movable and the other fixed (Newman, 2000). However, apart from taxa presenting readily apparent disparity, geometric asymmetry between the paired sides of the opercular plate has never been documented in acorn barnacles, with the exception of a previous study in which linear measurements of asymmetry were reported (Barnes & Healy, 1971;Chen et al., 2014). ...
The ivory barnacle Amphibalanus eburneus is a marine crustacean, which presents near-cosmopolitan distribution due to extensive introduction and exhibits a wide spectrum of phenotypic variation. To elucidate geographical differentiation among populations through invasion, we investigated variation in genetic structure, shell morphology, and recruitment pattern for A. eburneus, from the southern Korean Peninsula where it has been established since the late 1980s. We selected samples from four populations in corresponding ecologically-relevant regions representing all surrounding South Korean waters. From these we amplified the mitochondrial genetic marker cytochrome oxidase subunit I (COI) from 57 individuals and performed a populational genetic analyses with 11 additional GenBank sequences to evaluate population structure. To examine morphological variation, we applied two-dimensional landmark-based geometric morphometrics to the scutum and tergum for 148 and 151 individuals, respectively. Furthermore, we estimated the density of year-old individuals in the field to compare recruitment responses among localities. We detected 33 haplotypes among the four locations belonging to three distinct clades based on moderate intraspecific pairwise genetic distance (≥3.5%). The haplotypes in these clades were not locality-specific in their distribution. In contrast, we did detect interpopulation variation in opercular shape and morphospace structure, and one population could be separated from the rest based on its distinct tergum morphotype alone. This morphologically distinct population was also differentiated by displaying the lowest mean recruitment density. Our results indicate that although there is no relationship between molecular variation in the COI gene and geographic regions in South Korea, association with locality for operculum morphology, and recruitment response suggest ecological adaptation by this barnacle in a new habitat.
... Advances in deep-sea scientific sampling gear and increased collections of deep-sea hydrothermal vent barnacles worldwide have led to increased biodiversity and biogeography examinations, and also improved our understanding of barnacle phylogenetics and relationships within Thoracicalcarea (Newman, 2000;Watanabe et al., 2006;Pérez-Losada et al., 2008;Herrera et al., 2015;Chan, 2018;Yu et al., 2020;Dreyer et al., 2021). Phylogenetic studies with partial gene sequences have suggested that the ancestors of extant vent barnacles settled into hydrothermal vent fields over at least two independent events (Herrera et al., 2015). ...
... Although the vent sessile neoverrucid is sister to the clade containing neobrachylepadid and neolepadid barnacles, their phylogenetic relationships could not be confirmed with high support (Herrera et al., 2015). There are currently two genera in Neoverrucidae: Imbricaverruca Newman, 2000and Neoverruca Newman, 1989. Imbricaverruca is a monotypic genus that contains I. yamaguchii and found from hydrothermal vents of the Lau Basin in the Southwest Pacific. ...
... Neoverruca is composed of two species (N. brachylepadoformis and N. intermedia), which inhabit deep-sea hydrothermal vents in the Northwest Pacific Ocean Newman, 2000;Watanabe et al., 2006;Sha and Ren, 2015). The shells of Imbricaverruca and Neoverruca are different. ...
Imbricaverruca and Neoverruca are two genera of hydrothermal vent asymmetrical barnacles in Neoverrucidae, but found in vents of the Southwest Pacific and Northwest Pacific Oceans, respectively. Imbricaverruca has a flattened operculum and the shell base with multiple whorls of imbricating plates, while Neoverruca has an inclined operculum and the shell base with fewer developed imbricating plates. It has been hypothesized that Imbricaverruca has apomorphic shell characters in Neoverrucidae. Although the monophyletic relationship of the vent barnacle members in the superfamily Neolepadoidea were confirmed based on molecular phylogeny, the relationships between Neobrachylepadidae and Neoverrucidae, and between Neoverruca and Imbricaverruca have not been determined because there are no molecular data on Imbricaverruca. In this study, we sequenced three nuclear (18S rDNA, 28S rDNA, histone 3) and one mitochondrial (CO1) genes of I. yamaguchii from the Southwest Pacific. Our phylogenetic results showed that Neobrahchylepadidae is the sister taxon to Neoverrucidae (Imbricaverruca + Neoverruca), and Imbricaverruca and Neoverruca are monophyletic sister taxa each other, which not supporting the previous hypothesis that Neoverruca is sister to the clade containing Neobrahchylepadidae and Neolepadidae. These were implied that the differences in shell forms between Neoverruca and Imbricaverruca are a result of independent divergent evolution in different deep-sea basins.
... All neoverrucid species that have been described until now were found from hydrothermal vent sites. Up to now, a total of two genera and two species have been reported: Neoverruca brachylepadoformis Newman, 1989 and Imbricaverruca yamaguchii Newman, 2000(see Newman, 1989, 2000. ...
... All neoverrucid species that have been described until now were found from hydrothermal vent sites. Up to now, a total of two genera and two species have been reported: Neoverruca brachylepadoformis Newman, 1989 and Imbricaverruca yamaguchii Newman, 2000(see Newman, 1989, 2000. ...
... Family NEOVERRUCIDAE Newman, 1989 Diagnosis.-Distinguished from the extinct Proverrucidae and extant Verrucidae in having a primary wall elevated above the substratum by basal whorls of imbricating plates and an operculum including a median fatus, at least in juveniles (see Newman, 2000). ...
A new species of the neoverrucid barnacle genus NeoverrucaNewman, 1989, N. intermediasp. nov., collected from a hydrothermal vent area in the Okinawa Trough, is described and illustrated. It differs from N. brachylepadoformisNewman, 1989 by the apical end of the inner surface of the movable scutum being in a pocket-form, with a longitudinal groove upwards, and without median ridges; the adductor ridge (or myophore) of the fixed scutum stout and the inner distal part of the scutum without longitudinal ridge.
... However, for most vent-endemic species, migration and dispersal patterns among vent fields have not been well studied. Vent-endemic barnacles belonging to the family Neoverrucidae (Cirripedia: Thoracica) [2] consist of two genera, Neoverruca [2] and Imbricaverruca [3]. The former is abundant in hydrothermal vent fields of the northwestern Pacific [4,5]. ...
Barnacles of the genus Neoverruca are abundant near deep-sea hydrothermal vents of the northwestern Pacific Ocean, and are useful for understanding processes of population formation and maintenance of deep-sea vent faunas. Using next-generation sequencing, we isolated 12 polymorphic microsatellite loci from Neoverruca sp., collected in the Okinawa Trough. These microsatellite loci revealed 2-19 alleles per locus. The expected and observed heterozygosities ranged from 0.286 to 1.000 and 0.349 to 0.935, respectively. Cross-species amplification showed that 9 of the 12 loci were successfully amplified for Neoverruca brachylepadoformis in the Mariana Trough. A pairwise FST value calculated using nine loci showed significant genetic differentiation between the two species. Consequently, the microsatellite markers we developed will be useful for further population genetic studies to elucidate genetic diversity, differentiation, classification, and evolutionary processes in the genus Neoverruca.
... Eoverruca). Newman in Newman & Hessler (1989) subsequently placed both Eoverruca and Proverruca in a new family, Proverrucidae. ...
... In 1989, Newman and Hessler described a new extant cirripede, Neoverruca brachylepadiformis Newman & Hessler, 1989, from hydrothermal vents in the Mariana Basin in the West Pacific, and proposed that this form was a living fossil, a link between the predominantly Cretaceous brachylepadomorphs and Eoverruca. Newman (1989) described the ontogeny of this species, and demonstrated how it commenced life as a pedunculate form which progressively became more asymmetrical through growth. ...
... comm., 2013), but specific assignment of material from other localities is currently uncertain. Neoverruca was interpreted by Newman (1989) and Newman & Hessler (1989) as an intermediate form between brachylepadids and verrucids, and shown as immediately basal to the Late Cretaceous Eoverruca in their phylogeny (Newman & Hessler 1989, fig. 4). ...
Asymmetrical thoracican cirripedes of superfically similar ‘verrucomorph’ morphology are shown to have evolved independently three times from pedunculate ancestors. In each case, the loss of a peduncle was accompanied by preferential attachment on one side or the other, such that the rostrum, carina and one of each paired scuta and terga (fixed) formed a wall-like structure; the opposite scutum and tergum (free or moveable) transforming into an opercular lid closed by the scutal adductor muscle. True verrucomorphs (Eoverruca + Verrucidae) have their origin in the mid-Cretaceous sessile genera Pycnolepas and Faxelepas gen. nov., which demonstrate a marked, but facultative, tendency towards capitular asymmetry, confirmed by strong morphological similarities in plate morphology to the basal verrucomorph genus Eoverruca of Late Cretaceous age. Proverruca, another Late Cretaceous asymmetrical form, is not closely related; it evolved independently from a scalpelliform ancestor. The extant hydrothermal vent genera Neoverruca and Imbricaverruca evolved from symmetrical scalpelliforms of the family Neolepadidae, as demonstrated by both molecular and morphological evidence. Cladistic analysis of 38 characters in 14 living and fossil taxa has yielded a consensus tree showing a monophyletic Verrucomorpha, made up of a stem group (Eoverruca) and crown group (Verrucidae). Faxelepas bruennichi comb. nov. and Pycnolepas rigida are sister taxa to the Verrucomorpha, and evolved from a pedunculate ancestor. The Verrucidae are divided into a basal stem group (Altiverruca, Globosoverruca) and a derived crown group (all other genera). Within the crown group, the monophyletic Priscoverrucinae subfam. nov. is identified. Systematic revision of Cretaceous verrucomorphs and basal sessilians includes description of Pedupycnolepas gen. nov., Faxelepas gen. nov., Youngiverruca gen. nov. and Priscoverruca gen. nov. and two new species, Youngiverruca ruegenensis sp. nov. and Verruca jagti sp. nov. The development of asymmetry was perhaps a feeding adaptation, enabling barnacles to catch crawling benthic prey in deep ocean environments where food is scarce.
... Первый облигатный для хемобиотических сообществ (гидротермальных районов и зон холодных высачиваний) вид усоногих раков был обнаружен в Северной Пацифике (Newman, 1979), однако к настоящему времени число работ по фауне этой группы хемобиотических сообществ Южной Пацифики (Галкин, 1992;Yamaguchi, Newman, 1990;1997a;1997b;Jones, 1993;Newman, Yamaguchi, 1995;Buckeridge, 2000;Newman, 2000;Southward, Jones, 2003;Southward, 2005) заметно превосходит число аналогичных работ по Северной Пацифике (Newman, 1979;1989;Newman, Hessler, 1989;Yamaguchi et al., 2004). Число известных видов усоногих раков -облигатных обитателей хемобиотических сообществ -также существенно выше для Южной Пацифики, чем для северной части Тихого океана. ...
The barnacles (Cirripedia Thoracica) collected during 22 voyage of R/V «Akademic Mstislav Keldysh» taken at depth of 1490–2642 m in hydrothermal vent areas of Northern Pacific – Piipa volcanic seamount (Bering Sea) and the crest of the East Pacifi c Rise, are investigated. Probably new barnacle species, belong to family Chionelasmatidae, was found in the sample collected in the Piipa volcanic seamount region. Neolepas zevinae was found in the samples collected in the crest of the East Pacific Rise more deeply, than the most known depth of dwelling of this species is found.
... Buckeridge & Newman (2006) Remarks. In addition to the Verrucidae, the Verrucomorpha comprises two other families, the earliest of which, the Proverrucidae Newman, 1989 (Newman andHessler, 1989), (Cenomanian-Upper Maastrichtian), is characterised by the presence of extra (lateral) plates in the shell wall, and the extant Neoverrucidae Newman in Newman & Hessler, 1989, which is a true 'link' with early non-verrucomorph barnacles, as it retains several basal whorls of imbricating plates, demonstrating phyletic proximity to a scalpelliform ancestor (Newman, 2000;Buckeridge & Newman, 2006). ...
The discovery of a near-complete shell wall of a small verrucid barnacle from the Lower Danian (Palaeocene) portion of the El Haria Formation as exposed in the El Kef area (northwest Tunisia), permits its description as a new species with characters that, although conforming primarily to Verruca sensu stricto, show some similarities to Altiverruca Pilsbry, 1916, a genus that is not yet known from the fossil record. The present material extends the known geographic distribution of fossil verrucids, and constitutes one of the earliest species of Verruca to be documented subsequent to the Cretaceous/Palaeogene (K/Pg) boundary mass extinction event.
... Remarks. In addition to the Verrucidae, the Verrucomorpha comprises two other families, the earliest of which, the Proverrucidae Newman, 1989 (Newman and Hessler, 1989), (Cenomanian-Upper Maastrichtian), is characterised by the presence of extra (lateral) plates in the shell wall, and the extant Neoverrucidae Newman inNewman & Hessler, 1989, which is a true 'link' with early non-verrucomorph barnacles, as it retains several basal whorls of imbricating plates, demonstrating phyletic proximity to a scalpelliform ancestor (Newman, 2000;Buckeridge & Newman, 2006). ...
The discovery of a near-complete shell wall of a small verrucid barnacle from the Lower Danian (Palaeocene) portion of the El Haria Formation as exposed in the El Kef area (northwest Tunisia), permits its description as a new species with characters that, although conforming primarily to Verruca sensu stricto, show some similarities to Altiverruca Pilsbry, 1916, a genus that is not yet known from the fossil record. The present material extends the known geographic distribution of fossil verrucids, and constitutes one of the earliest species of Verruca to be documented subsequent to the Cretaceous/Palaeogene (K/Pg) boundary mass extinction event.
... The Lau Basin in the southwest Pacific is the site of a backarc spreading center in which hydrothermal geochemistry and ecology are distinct from EPR and other well-studied vent sites in the Pacific Ocean [5]. The Eastern Lau Spreading Center (ELSC), located within the basin (from 19°20'S to 22°45'S) (Figure 1), exhibits a wide-ranging variation in seafloor spreading rates, ridge morphology, ecological communities, and vent fluid chemistry [6][7][8][9]. In particular, large gradients occur over relatively short distances along the north-to-south axis of the ELSC. ...
... However, results from expeditions to the Lau Basin in the 1980's indicated that the chemical composition and biological community diversity near the ELSC vents were uniquely different from well-studied sites at EPR. For example, macrobiological communities containing sulfide-reliant endosymbionts at the Lau Basin vent sites were distinctive for the diversity of snail species, amongst the most diverse known for deepsea hydrothermal communities [9]. Tubeworm species such as Riftia pachytila, which host sulfide-reliant endosymbionts and are ubiquitous in eastern Pacific vent sites, were apparently non-existent in the Lau vent sites [14]. ...
... The ELSC is notable for north-to-south gradients in seafloor geology, ridge morphology and hydrothermal activity [6][7][8][9]. Results of this study indicated that Fe, Zn and sulfide concentrations in the high temperature vent fluids (> 250°C) decreased from north to south at the four sites (Table 1 and Figure 3). In particular, the average total Fe concentration from high temperature vents at Kilo Moana was 1126 ± 597 µM (n = 5), almost ten times greater than vent fluids from the other sites (ranging from 124 to 234 µM total Fe) ( Table 1). ...
The speciation of dissolved sulfide in the water immediately surrounding deep-ocean hydrothermal vents is critical to chemoautotrophic organisms that are the primary producers of these ecosystems. The objective of this research was to identify the role of Zn and Fe for controlling the speciation of sulfide in the hydrothermal vent fields at the Eastern Lau Spreading Center (ELSC) in the southern Pacific Ocean. Compared to other well-studied hydrothermal systems in the Pacific, the ELSC is notable for unique ridge characteristics and gradients over short distances along the north-south ridge axis.
In June 2005, diffuse-flow (< 50 degrees C) and high-temperature (> 250 degrees C) vent fluids were collected from four field sites along the ELSC ridge axis. Total and filtered Zn and Fe concentrations were quantified in the vent fluid samples using voltammetric and spectrometric analyses. The results indicated north-to-south variability in vent fluid composition. In the high temperature vent fluids, the ratio of total Fe to total Zn varied from 39 at Kilo Moana, the most northern site, to less than 7 at the other three sites. The concentrations of total Zn, Fe, and acid-volatile sulfide indicated that oversaturation and precipitation of sphalerite (ZnS(s)) and pyrite (FeS2(s)) were possible during cooling of the vent fluids as they mixed with the surrounding seawater. In contrast, most samples were undersaturated with respect to mackinawite (FeS(s)). The reactivity of Zn(II) in the filtered samples was tested by adding Cu(II) to the samples to induce metal-exchange reactions. In a portion of the samples, the concentration of labile Zn2+ increased after the addition of Cu(II), indicating the presence of strongly-bound Zn(II) species such as ZnS clusters and nanoparticles.
Results of this study suggest that Zn is important to sulfide speciation at ELSC vent habitats, particularly at the southern sites where Zn concentrations increase relative to Fe. As the hydrothermal fluids mix with the ambient seawater, Zn-sulfide clusters and nanoparticles are likely preventing sulfide oxidation by O2 and reducing bioavailability of S(-II) to organisms.
