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Ascidian embryos have been studied since the birth of experimental embryology at the end of the 19th century. They represent textbook examples of mosaic development characterized by a fast development with very few cells and invariant cleavage patterns and lineages. Ascidians belong to tunicates, the vertebrate sister group, and their study is essential to shed light on the emergence of vertebrates. Importantly, deciphering developmental gene regulatory networks has been carried out mostly in two of the three ascidian orders, Phlebobranchia and Stolidobranchia. To infer ancestral developmental programs in ascidians, it is thus essential to carry out molecular embryology in the third ascidian order, the Aplousobranchia. Here, we present genomic resources for the colonial aplousobranch Clavelina lepadiformis : a transcriptome produced from various embryonic stages, and an annotated genome. The assembly consists of 184 contigs making a total of 233.6 Mb with a N50 of 8.5 Mb and a L50 of 11. The 32,318 predicted genes capture 96.3% of BUSCO orthologs. We further show that these resources are suitable to study developmental gene expression and regulation in a comparative framework within ascidians. Additionally, they will prove valuable for evolutionary and ecological studies. [ABSTRACT FROM AUTHOR]

An unidentified colonial ascidian called gaikotsu-panda-hoya in Japanese, literally meaning 'skeleton panda ascidian,' has been attracting SCUBA divers' attention for the past few years since its strange appearance was introduced on the Internet by a diving shop in Kumejima Island, Japan. To confirm the taxonomic status of this species, fresh samples were collected from a diving point off the coast of Kumejima Island. Our morphological examination revealed that they represent a new species, herein described as Clavelina ossipandae sp. nov., which can be distinguished from 44 congeners in the genus Clavelina Savigny, 1816 by the combination of the following seven characteristics: i) colony consisting of completely free zooids, ii) zooids up to 20 mm in length, iii) in the living state, zooids transparent, with laterally elongated white patch between oral and atrial siphons, as well as four black markings, one between siphons, one mid-dorsally, and the other two situated laterally in a pair on the anterior part of the body, iv) transverse vessels white, v) endostyle black, vi) 10-14 stigmatal rows, and vii) two longitudinal muscular bands running from the abdomen to the endostyle on each side. Partial sequences (810 bp) of the cytochrome c oxidase subunit I gene from the holotype and one of the paratypes differed at 10 sites from each other (1.26% K2P distance) but were the same when translated into amino acids. A phylogenetic tree supported that this species is included in the genus Clavelina. [ABSTRACT FROM AUTHOR]

Aplousobranchia ascidians from two different families were integrated with morphological characteristics and molecular phylogenetic analysis for the first time. The present study employed morphological descriptions (colony structures, tunic, zooids, spicules stigmata and test) and a molecular approach, using a fragment of the mitochondrial cytochrome oxidase I (COI) gene of four Aplousobranchia colonial ascidians Aplidium conicum (98%), Aplidium elegans (98%), Didemnum fulgens (92%) and Trididemnum cyanophorum (94%) from Andaman and Nicobar Islands. Bar-coded sequences were extracted with BLAST format from NCBI and the heritable diversity of the submitted sequences were compared with associated ascidian species. Study revealed that the evolutionary relationship among the ascidian species exhibited the constant clades, which may help for rapid reassessment of morphological characters of the species distributed worldwide. [ABSTRACT FROM AUTHOR]

Tunicata, a diverse clade of approximately 3000 described species of marine, filter-feeding chordates, is of great interest to researchers because tunicates are the closest living relatives of vertebrates and they facilitate comparative studies of our own biology. The group also includes numerous invasive species that cause considerable economic damage and some species of tunicates are edible. Despite their diversity and importance, relationships among major lineages of Tunicata are not completely resolved. Here, we supplemented public data with transcriptomes from seven species spanning the diversity of Tunicata and conducted phylogenomic analyses on data sets of up to 798 genes. Sensitivity analyses were employed to examine the influences of reducing compositional heterogeneity and branch-length heterogeneity. All analyses maximally supported a monophyletic Tunicata within Olfactores (Vertebrata + Tunicata). Within Tunicata, all analyses recovered Appendicularia sister to the rest of Tunicata and confirmed (with maximal support) that Thaliacea is nested within Ascidiacea. Stolidobranchia is the sister taxon to all other tunicates except Appendicularia. In most analyses, phlebobranch tunicates were recovered paraphyletic with respect to Aplousobranchia. Support for this topology varied but was strong in some cases. However, when only the 50 best genes based on compositional heterogeneity were analysed, we recovered Phlebobranchia and Aplousobranchia reciprocally monophyletic with strong support, consistent with most traditional morphology-based hypotheses. Examination of internode certainty also cast doubt on results of phlebobranch paraphyly, which may be due to limited taxon sampling. Taken together, these results provide a higher-level phylogenetic framework for our closest living invertebrate relatives. [ABSTRACT FROM AUTHOR]

Didemnidae is the largest family of tunicates within Aplousobranchia, with about 578 species. This family comprises eight genera: Atriolum, Clitella, Didemnum, Diplosoma, Leptoclinides, Lissoclinum, Polysyncraton, and Trididemnum. Morphological and molecular data suggest that Didemnidae is monophyletic, but the monophyly of each didemnid genus and their phylogenetic relationships are still poorly understood. This study aimed to evaluate the monophyly of six of the eight didemnid genera and assess their phylogenetic relationships based on mitochondrial (COI) and nuclear (18S) sequences. All genera were recovered as monophyletic except Trididemnum. Didemnum comprises two clades that differ one from the other by the presence of an atrial lip, the number of testicular lobes, and the number of ampullae in larvae. These morphological differences indicate that Didemnum could be split into two genera. Morphological evidence and previous taxonomists have suggested a close relationship between Didemnum and Polysyncraton as well as between Diplosoma and Lissoclinum. We re-evaluate these hypotheses, which are supported by our 18S sequences and concatenated data. [ABSTRACT FROM AUTHOR]

Managers and policymakers in eastern Canada embrace science-based management of nonindigenous species and may benefit from having comprehensive regional species checklists at subnational jurisdictional levels. In this paper, regional checklists provide an account of the richness of ascidians in eastern Canada. Records of 58 ascidians resulted from reviewing 108 published sources, accessing data from two online databases, and collecting some common indigenous ascidian specimens. Analysis comparing the similarity of species among nine regions indicates that there is greater similarity in species composition between contiguous regions than between noncontiguous regions and suggests that there are four zoogeographic clusters in eastern Canada. Our checklists can inform managers and policymakers of the diversity of the ascidian taxa and can minimize taxonomic uncertainties of established nonindigenous and prospective invading species, for example, by identifying indigenous species that are congeners of nonindigenous species. The maintenance of checklists can be a valuable tool for the management of nonindigenous species as baselines to estimate changes in richness and to document the invasion status of nonindigenous species over time. For example, more importance can be placed on the spread of nonindigenous ascidians from one zoogeographic cluster to another than spread within the same cluster. [ABSTRACT FROM AUTHOR]

Background: More than 50 ascidian species distributed in the Palk Bay, Southeast coat of India. Up to a very few molecular work has been performed to determine the evolutionary relationships of ascidians in the world.Objectives: Present study explored the value of mtDNA data in assessing phylogenetic relationships within the family Ascidiacea, Didemnidae, Styelidae, and molecular identification of ascidians from the Palk Bay, Southeast coast of India.Materials and methods: The phylogeny analysis was executed based on mitochondrial cytochrome oxidase subunit I (COI) sequences of ascidian species. A BLASTN search can be run to determine the similarity of an unknown DNA sequence (query) with the collection of all known DNA sequences in GenBank.Result: The BLASTN results showed thatDidemnum candidum, Ascidia ahodoriandStyela clavamatch the other listings for these species in GenBank. Mitochondrial COI gene sequences of collected ascidians were submitted to GenBank and obtained the accession numbers.Conclusion: This study is the first report of molecular identification of ascidian species of the Palk Bay, Southeast coast of India. This information about Palk Bay ascidian communities provides a baseline of general biodiversity of that ecosystem. [ABSTRACT FROM AUTHOR]

The order Aplousobranchia (Chordata, Ascidiacea) contains approximately 1500 species distributed worldwide. Their phylogeny, however, remains unclear, with unresolved family relationships. While most Aplousobranchia are colonial, debates exist concerning the phylogenetic position of families such as the Diazonidae and Cionidae, which exhibit a solitary lifestyle and share morphological characteristics with both Aplousobranchia and Phlebobranchia orders. To clarify the phylogenetic position of the Diazonidae and Cionidae, we determined the complete mitochondrial sequence of the solitary diazonid Rhopalaea idoneta . The phylogenetic reconstruction based on the 13 mitochondrial protein coding genes strongly supports a positioning of Diazonidae well-nested within the Aplousobranchia rather than a positioning as a sister clade of the Aplousobranchia. In addition, we examined the regenerative ability of R. idoneta. Similar to colonial Aplousobranchia, R. idoneta was found to be able to completely regenerate its thorax. Ciona, also known to possess high regenerative abilities, is the Aplousobranchia sister clade rather than a member of the Phlebobranchia. Our results thus indicate that the colonial lifestyle was acquired in the Aplousobranchia, starting from a Ciona -like solitary ancestor and secondarily lost in Diazonidae representatives such as Rhopalaea. The solitary lifestyle of Rhopalaea is thus a derived characteristic rather than an ancestral trait. [ABSTRACT FROM AUTHOR]

Many sessile, suspension-feeding marine invertebrates mate by spermcasting: aquatic sperm are spawned and gathered by conspecific individuals to fertilize eggs that are generally retained during development. In two phylogenetically distant examples, a cheilostome bryozoan and an aplousobranch ascidian, the receipt of allosperm has previously been shown to alter sex allocation by triggering female investment in eggs and brooding. Here we report experiments demonstrating that two species of cyclostome bryozoan also show restrained female investment in the absence of mating opportunity. In Tubulipora plumosa, the production of female zooids and progeny is much reduced in reproductive isolation. In Filicrisia geniculata, development of distinctive female zooids (gonozooids) begins but halts in the absence of mating opportunity, and no completed gonozooids or progeny result. Reduced female investment in the absence of a mate thus occurs in at least two orders of Bryozoa, but significant differences in detail exist and the evolutionary history within the phylum of the mechanism(s) by which female investment is initiated might be complex. The broadening taxonomic spectrum of examples where female investment appears restrained until allosperm becomes available may signify a general adaptive strategy among outcrossing modular animals, analogous to similarly adaptive sex allocation typical of many flowering plants. [ABSTRACT FROM AUTHOR]

We documented changes in the abundance and distribution patterns of tunic cells involved in the allorecognition response of the colonial aplousobranch Didemnum vexillum, whose zooids do not share a common vascular system. A histological examination of the fusion zone of isogeneic (CIAs) and allogeneic (CAAs) fused colony assays revealed that tunic cuticles were rapidly regenerated. The underlying tunic matrix fused readily in all assays and controls. We identified four different types of tunic cells. Phagocytic cells represented the most abundant cell type in allogeneic fusions, followed by morula cells. These cells were more abundant at the immediate fusion junction than at 120 μm or 240 μm from the junction, most likely because they mediate the allorecognition reaction. Elongated filopodial cells also were present, although only at very low abundances, and a layer of bladder cells was located immediately below the cuticle. Our results provide quantitative evidence for the involvement of tunic cells in the allorecognition response of a highly invasive ascidian. [ABSTRACT FROM AUTHOR]

The class Ascidiacea presents fundamental opportunities for research in the fields of development, evolution, ecology, natural products and more. This review provides a comprehensive overview of the current knowledge regarding the global biodiversity of the class Ascidiacea, focusing in their taxonomy, main regions of biodiversity, and distribution patterns. Based on analysis of the literature and the species registered in the online World Register of Marine Species, we assembled a list of 2815 described species. The highest number of species and families is found in the order Aplousobranchia. Didemnidae and Styelidae families have the highest number of species with more than 500 within each group. Sixty percent of described species are colonial. Species richness is highest in tropical regions, where colonial species predominate. In higher latitudes solitary species gradually contribute more to the total species richness. We emphasize the strong association between species richness and sampling efforts, and discuss the risks of invasive species. Our inventory is certainly incomplete as the ascidian fauna in many areas around the world is relatively poorly known, and many new species continue to be discovered and described each year. [ABSTRACT FROM AUTHOR]

Seventy-eight species of Ascidiacea, including 21 new species from deeper waters (about 100 m and occasionally to about 500 m) off Western Australia appear to represent a diverse fauna not previously sampled. The relatively few solitary species taken include two possibly introduced (Styela plicata and S. clava). Large aplousobranch colonies dominate the collections. Aplousobranch species with numerous, small, deeply embedded zooids in massive three-dimensional, vertical and often branched or stalked inflexible colonies supported by embedded sand (Polyclinindae) or calcareous spicules (Didemnidae) are common. Species with flexible stalks (enabling them to respond to prevailing currents) such as Clavelina meridionalis, Sigillina australis and S. cyanea were not taken often. Some known species (e.g. Synoicum chrysanthemum, and Atriolum tubiporum) previously considered rare were taken at these depths in significant numbers. Other unusual taxa (Pseudodiazona longigona and Condominium spp.) are recorded from new locations but do not appear to be common. [ABSTRACT FROM AUTHOR]

Forty-eight species (including nine that are new) of the 288 Didemnidae now known from Australian waters are reported on in the present work. Dredged material is from benthic habitats in inter-reefal areas of the Great Barrier Reef and from deeper water canyons off northeastern and northwestern Tasmania. Collections from shallower waters of Port Davey (western Tasmania) and Kangaroo Island (South Australia) were hand-collected by scuba divers. Seven of the eight known genera of the Didemnidae are represented. Despite this emphasis on sampling in temperate waters, only 14 species (all indigenous) are recorded from around the southern half of the continent. The species diversity of this family is greater in the tropics, with 34 species being recorded. Nearly half of these are known also from the tropical western Pacific and some from the Indian Ocean as well. Tropical indigenous species are relatively few. [ABSTRACT FROM AUTHOR]

Three colonial aplousobranch ascidian species from the central coast of New South Wales were originally described from one of the earliest significant regional collections of Australian ascidians (Herdman, 1899). They have a similar habit, their colonies being long, oval to cylindrical gelatinous heads on relatively short, thick, fleshy stalks and they all respond in the same way to the multidirectional currents to which they are subjected in the habitats they occupy. Nevertheless, they are not closely related phylogenetically. One is the monotypic Euclavella claviformis (Herdman, 1899), known only from the central eastern Australian coast and the North Island (New Zealand). The second species is the indigenous Hypsistozoa distomoides (Herdman, 1899), known from the Great Australian Bight and the central eastern coast of Australia but related closely to the only other species known in this genus, its type species H. fasmeriana (Michaelsen, 1924), known only from New Zealand. The third species discussed, Sigillina cyanea (Herdman, 1899), an indigenous Australian species known from a wide range around the Australian continent, has no closely related species in the indigenous New Zealand fauna although a similar and partly sympatric species Sigillina australis (Savigny, 1816) does occur in New Zealand. A close relationship of Sigillina to the Pseudodistomidae (rather than the Holozoidae) is apparent. Larvae of H. distomoides are found to have characters formerly known only in larvae of the type species (see Brewin 1956, 1959). [ABSTRACT FROM AUTHOR]

Thirty–five species (including eight that are new) of non–didemnid aplousobranch species and 25 phlebobranch (including one new) and stolidobranch species are discussed and a further 42 species in non–didemnid taxa in all suborders are newly recorded. Material was taken by scuba diving in the waters around Kangaroo I. and other southern Australian and Tasmanian locations; and further material was taken by dredge from canyons off the southeastern Continental Slope and from inter–reefal areas in the northern Great Barrier Reef region. The known geographic ranges of many southern Australian species are extended into Tasmanian waters and a new species of Rhopalaea is described from the Tasmanian Canyons. The genera Sigillina and Pseudodistoma are found to be diverse in Australian temperate waters, and they have many characters in common, suggesting a probable phylogenetic relationship that formerly was not recognised. The greatest diversity of species reported is from the vicinity of the northern Great Barrier Reef. Almost without exception these are non–indigenous tropical species often extending into Australian temperate locations where they are sympatric with indigenous species found mainly around the southern half of the continent. Pyura stolonifera has a similar range in temperate Australian waters but appears also to occur on the South African and Chilean coasts in similar habitats and may represent a Gondwana relict. [ABSTRACT FROM AUTHOR]

The majority of the 58 species discussed, including 10 new species, were collected by scuba divers at 5–10?m in waters around Australia. Species are from Leptoclinides (11), Polysyncraton (11), Didemnum (19), Trididemnum (7), Lissoclinum (8), Clitella (1), and Diplosoma (1), and new species are in all except Trididemnum and Diplosoma . Additional characters (including a pyloric vesicle reported previously in the Holozoidae and several unrelated didemnid taxa) have been detected for the monotypic genus Clitella Kott, 2001, which is recorded for only the second time. A review of known Australian ascidian species confirms the Didemnidae as the most speciose ascidian family in these waters. In this family, there appears not to be appreciable gene flow between tropical and temperate waters and few species have a continuous tropical–temperate range. A preponderance of Western Pacific non-indigenous species is in the north, while indigenous species, some probably isolated from related tropical ones, are dominant in the temperate waters of the southern half of the continent. Although intraspecific variation and convergence obscures species differences, some aspects of the living organisms detected in in situ photographs contribute to identification. Keys to Australian didemnid species described since the publication of the Australian Ascidiacea part 4, Didemnidae (Kott 2001) are included. [ABSTRACT FROM AUTHOR]

We explored the usefulness of mtDNA data in assessing phylogenetic relationships within the Ascidiacea. Although ascidians are a crucial group in studies of deuterostome evolution and the origin of chordates, little molecular work has been done to ascertain the evolutionary relationships within the class, and in the studies performed to date the key group Aplousobranchiata has not been adequately represented. We present a phylogenetic analysis based on mitochondrial cytochrome c oxidase subunit I (COI) sequences of 37 ascidian species, mainly Aplousobranchiata (26 species). Our data retrieve the main groups of ascidians, although Phlebobranchiata appeared paraphyletic in some analyses. Aplousobranch ascidians consistently appeared as a derived group, suggesting that their simple branchial structure is not a plesiomorphic feature. Relationships between the main groups of ascidians were not conclusively determined, the sister group of Aplousobranchiata was the Stolidobranchiata or the Phlebobranchiata, depending on the analysis. Therefore, our data could not confirm an Enterogona clade (Aplousobranchiata+Phlebobranchiata). All of the tree topologies confirmed previous ideas, based on morphological and biochemical characters, suggesting that Cionidae and Diazonidae are members of the clade Aplousobranchiata, with Cionidae occupying a basal position within them in our analyses. Within the Aplousobranchiata, we found some stable clades that provide new data on the evolutionary relationships within this large group of ascidians, and that may prompt a re-evaluation of some morphological characters. [Copyright &y& Elsevier]

Eighteen new species of Didemnidae are included in the 71 species discussed. Two hundred and forty-one species of this family now are known from around the Australian continent. Newly recorded material is from the Western Australian Museum (principally from north-western Australia), the South Australian Museum (from Kangaroo I. and Tasmanian waters) and the Queensland Museum (from Darwin and the Northern Territory). Many of the species recorded from north-western, north-eastern and the north of Australia have a range that extends into the western Pacific and sometimes the western Indian Ocean. Tropical didemnid species seldom extend into temperate Australian waters, where the species are largely indigenous. The genus Didemnum dominates the fauna in both tropical and temperate waters. Of the new species, half are in the genus Polysyncraton (five tropical and four temperate species), four are Lissoclinum spp., two are described in each of the genera Didemnum and Leptoclinides , and one in the genus Diplosoma . Spicules are confirmed as reliable, genetically controlled characters showing little intraspecific variation. Amongst the known species, the unusual colonial organization reported for Leptoclinides glauerti Michaelsen, 1930 , with its atrial apertures opening directly on the under surface of the colony, is confirmed. Originally described from a single mutilated part of a colony, the species has been redescribed, and its status in the genus Atriolum validated. Trididemnum pseudodiplosoma ( Kott, 1962 ), formerly known only from South Australia, is found to extend into tropical waters of the Northern Territory and the Coral Sea; and Didemnum incanum ( Herdman, 1899 ) appears to be one of the few known species with a trans-Tasman range. [ABSTRACT FROM AUTHOR]

The transport speed of latex spheres trapped in the mucous feeding filter of 30 specimens of Calvelina lepadiformis (Müller 1776) was measured by microscopical examination and a stop watch. The speed across the gill screen increased from 24 μm s-1 in small animals to 220 μm s-1 in large ones. The net translocation speed of the food roll behind the dorsal lamina was generally much slower but revealed a similar trend (7 – 78 μm s-1). The increased feeding filter speed in larger animals was parallelled by an increased branchial circumference in such a way that the duration of water filtration through each part of the filter remained almost constant (between 38 and 56 s as means for all groups of animals). The amount of filter secreted by large animals approached 150 cm² h-1 or 2.5 m² week-1. Based on available data on water filtration rates it could be calculated that the speed of water through the filter was of the same order of magnitude as the observed translocation speed of the filter. [ABSTRACT FROM AUTHOR]

Allorecognition, the ability to distinguish self or kin from unrelated conspecifics, plays several important biological roles in invertebrate animals. Two of these roles include negotiating limited benthic space for colonial invertebrates, and inbreeding avoidance through self-incompatibility systems. Subphylum Tunicata (Phylum Chordata), the sister group to the vertebrates, is a promising group in which to study allorecognition. Coloniality has evolved many times independently in the tunicates, and the best known invertebrate self-incompatibility systems are in tunicates. Recent phylogenomic studies have coalesced around a phylogeny of the Tunicata as well as the Order Stolidobranchia within the Tunicata, providing a path forward for the study of allorecognition in this group. [ABSTRACT FROM AUTHOR]

Lissoclinum midui was collected from Lyudao (Green Island), Taiwan. This is the first record outside of the type locality, Kumejima Island (Ryukyu Archipelago, Japan), of this species. The colonies in Lyudao were found on non-geniculate coraline algae covering the coral limestones in the reef crest (0.5–1 m deep). They were not rare at the collection sites, and it is uncertain why this species has only been recorded in the two sites in Taiwan–Ryukyus. It may be that the shallow, wave-exposed reef crest is a poorly investigated habitat for small benthic invertebrates. [ABSTRACT FROM PUBLISHER]