Wanninger, Andreas; Koop, Demian; Degnan, Bernard M.
The development of gymnolaemate Ectoprocta includes a larval stage of either the coronate or the cyphonautes type. Herein, we provide the first description of the larval neural anatomy of a coronate larva using immunocytochemical methods. We used antibodies against the neurotransmitters serotonin...... that the larval neuroanatomy and the processes that underlie the reorganization of larval organ systems during metamorphosis may vary much more among lophotrochozoan taxa than previously thought....... and FMRFamide and followed the fate of immunoreactive cells through metamorphosis. The larval serotonergic nervous system of Triphyllozoon mucronatum consists of an apical commissure, one pair of lateral axons, a coronate nerve net, an internal nerve mesh, and one pair of axons innervating the frontal organ....... FMRFamide is only found in the larval commissure and in the lateral axons. The entire serotonergic and FMRFamidergic nervous system is lost during metamorphosis and the adult neural structures form independent of the larval ones. In the postlarval zooid, both neurotransmitters are detected in the cerebral...
Full Text Available Abstract Background The phylogenetic position of Bryozoa is one of the most controversial issues in metazoan phylogeny. In an attempt to address this issue, the first bryozoan mitochondrial genome from Flustrellidra hispida (Gymnolaemata, Ctenostomata was recently sequenced and characterized. Unfortunately, it has extensive gene translocation and extremely reduced size. In addition, the phylogenies obtained from the result were conflicting, so they failed to assign a reliable phylogenetic position to Bryozoa or to clarify lophophorate phylogeny. Thus, it is necessary to characterize further mitochondrial genomes from slowly-evolving bryozoans to obtain a more credible lophophorate phylogeny. Results The complete mitochondrial genome (15,433 bp of Bugula neritina (Bryozoa, Gymnolaemata, Cheilostomata, one of the most widely distributed cheliostome bryozoans, is sequenced. This second bryozoan mitochondrial genome contains the set of 37 components generally observed in other metazoans, differing from that of F. hispida (Bryozoa, Gymnolaemata, Ctenostomata, which has only 36 components with loss of tRNAser(ucn genes. The B. neritina mitochondrial genome possesses 27 multiple noncoding regions. The gene order is more similar to those of the two remaining lophophorate phyla (Brachiopoda and Phoronida and a chiton Katharina tunicate than to that of F. hispida. Phylogenetic analyses based on the nucleotide sequences or amino acid residues of 12 protein-coding genes showed consistently that, within the Lophotrochozoa, the monophyly of the bryozoan class Gymnolaemata (B. neritina and F. hispida was strongly supported and the bryozoan clade was grouped with brachiopods. Echiura appeared as a subtaxon of Annelida, and Entoprocta as a sister taxon of Phoronida. The clade of Bryozoa + Brachiopoda was clustered with either the clade of Annelida-Echiura or that of Phoronida + Entoprocta. Conclusion This study presents the complete mitochondrial genome of a
Taylor, P.D.; McKinney, F.K.
Turnerella Taylor & McKinney, 2006, p. 164, introduced for a new genus of cribrimorph Cheilostomata (Bryozoa), is preoccupied by Turnerella Cockerell, 1910, a genus of Hymenoptera, and two other introductions of the same name for new insect genera. We propose Turnerellina as a new name to replace
Ostrovsky, Andrew N; Grischenko, Andrei V; Taylor, Paul D; Bock, Phil; Mawatari, Shunsuke F
The anatomical structure of internal sacs for embryonic incubation was studied using SEM and light microscopy in three cheilostome bryozoans-Nematoflustra flagellata (Waters,1904), Gontarella sp., and Biflustra perfragilis MacGillivray, 1881. In all these species the brood sac is located in the distal half of the maternal (egg-producing) autozooid, being a conspicuous invagination of the body wall. It consists of the main chamber and a passage (neck) to the outside that opens independently of the introvert. There are several groups of muscles attached to the thin walls of the brood sac and possibly expanding it during oviposition and larval release. Polypide recycling begins after oviposition in Gontarella sp., and the new polypide bud is formed by the beginning of incubation. Similarly, polypides in brooding zooids degenerate in N. flagellata and, sometimes, in B. perfragilis. In the evolution of brood chambers in the Cheilostomata, such internal sacs for embryonic incubation are considered a final step, being the result of immersion of the brooding cavity into the maternal zooid and reduction of the protecting fold (ooecium). Possible reasons for this transformation are discussed, and the hypothesis of Santagata and Banta (Santagata and Banta1996) that internal brooding evolved prior to incubation in ovicells is rejected. J. Morphol. (c) 2006 Wiley-Liss, Inc.
Ostrovsky, Andrew N; Dick, Matthew H; Mawatari, Shunsuke F
We studied by SEM the external morphology of the ooecium in eight bryozoans of the genus Cauloramphus (Cheilostomata, Calloporidae): C. spinifer, C. variegatus, C. magnus, C. multiavicularia, C. tortilis, C. cryptoarmatus, C. niger, and C. multispinosus, and by sectioning and light microscopy the anatomy of the brooding apparatus of C. spinifer, C. cryptoarmatus, and C. niger. These species all have a brood sac, formed by invagination of the non-calcified distal body wall of the maternal zooid, located in the distal half of the maternal (egg-producing) autozooid, and a vestigial, maternally budded kenozooidal ooecium. The brood sac comprises a main chamber and a long passage (neck) opening externally independently of the introvert. The non-calcified portion of the maternal distal wall between the neck and tip of the zooidal operculum is involved in closing and opening the brood sac, and contains both musculature and a reduced sclerite that suggest homology with the ooecial vesicle of a hyperstomial ovicell. We interpret the brooding apparatus in Cauloramphus as a highly modified form of cheilostome hyperstomial ovicell, as both types share 1) a brood chamber bounded by 2) the ooecium and 3) a component of the distal wall of the maternal zooid. We discuss Cauloramphus as a hypothetical penultimate stage in ovicell reduction in calloporid bryozoans. We suggest that the internal-brooding genus Gontarella, of uncertain taxonomic affinities, is actually a calloporid and represents the ultimate stage in which no trace of the ooecium remains. Internal brooding apparently evolved several times independently within the Calloporidae.
Menon, N.R; Nair, N.B.
The species Bugulella clavata collected from the Krusadai island, Gulf of Mannar is redescribed Description is based on the three fragments of a colony at a depth of 5 m Redescription has helped to important features such as nature and shape...
Nielsen, Claus; Worsaae, Katrine
We have studied larvae of the freshwater ctenostome Hislopia malayensis with scanning electron microscopy (SEM), confocal laser scanning microscopy (CLSM), and LM of serial sections. Some additional observations on larvae of M. membranacea using SEM and CLSM are also reported. The overall configu...
Seventy new species were recorded during the study of the ectoproctous bryozoans of Indian waters Of these Alcyonidium erectum is of special interest Several zoaria of A erectum were collected from the appendages and carapace of Portunus pelagicus...
Funch, P.; Kristensen, R.M.
The mouthparts of the Norway lobster Nephrops are colonized by an acoelomate metazoan, Symbion pandora gen. et sp, nov. Sessile stages continually produce inner buds replacing feeding structures, They also produce one of three motile stages: (1) larvae containing new feeding stages, (2) dwarf males......, which settle on feeding stages, or (3) females, which settle onto lobster mouthparts, and eventually degenerate, giving rise to dispersive larvae, All motile stages are short-lived, and do not feed, The structure and function of the cilia suggest a phylogenetic position in Protostomia, while some...
Schwaha, Thomas F; Wanninger, Andreas
Myoanatomical studies of adult bryozoans employing fluorescent staining and confocal laser scanning microscopy (CLSM) have been chiefly conducted on freshwater bryozoans. The diversity of muscular systems in the marine bryozoans is currently not well known with only two species being studied in more detail. The aim of this study is to unravel the diversity of muscle systems of 15 ctenostome bryozoans by phalloidin-coupled fluorescence stainings combined with CLSM. In general, the myoanatomy of the selected ctenostomes shows significant similarities and consists of 1) muscles associated with the body wall, 2) apertural muscles, 3) lophophoral muscles, 4) tentacle sheath muscles, 5) digestive tract muscles and 6) the prominent retractor muscles. Differences are present in the arrangement of the apertural muscles from generally three muscles sets of four bundles, which in some species can be partially reduced or modified into a bilateral arrangement. The cardiac region of the digestive tract shows a distinct sphincter in four of the six studied clades. In some cases the cardiac region forms a prominent proventriculus or gizzard. Tentacle sheath muscles in victorelloideans and walkerioideans are arranged diagonally and differ from the simple longitudinal muscle arrangements common to all other taxa. Lophophoral base muscles consist of four sets that vary in the size of the sets and in the shape of the inner lophophoral ring, which either forms a complete ring or separate, intertentacular muscle bundles. The stolon-forming walkeridiodean ctenostomes show prominent transverse muscles in their stolons. These are always present in the shorter side stolons, but their occurrence in the main stolon seems to depend on the colony form, being present in creeping but absent in erect colony forms. This study represents the first broad survey of muscular systems in adult ctenostome bryozoans and shows a certain degree of conservation in a series of diverse colony forms belonging to five major clades. However, several myoanatomical features such as the cardiac sphincter, basal (possibly transitory) cystid muscles, tentacle sheath muscles or apertural muscle arrangement vary across taxa and thus show a high potential for the assessment of character evolution within ctenostomes. As such, this study represents an essential contribution towards determining and reconstructing the character states of the bryozoan ground pattern once a reliable phylogenetic tree of the whole phylum becomes available.
Full Text Available Ten new species belonging to three new genera (Atlantisina gen. nov., Bathycyclopora gen. nov., Calvetopora gen. nov. of umbonulomorph bryozoans from northeastern Atlantic seamounts, islands, and the continental slope are introduced. We furthermore erect the new family Atlantisinidae fam. nov. for these genera. Eight new species belong to the new genus Atlantisina: Atlantisina atlantis gen. et sp. nov. (type species, A. acantha gen. et sp. nov., A. gorringensis gen. et sp. nov., A. inarmata gen. et sp. nov., A. lionensis gen. et sp. nov., A. meteor gen. et sp. nov., A. seinensis gen. et sp. nov., and A. tricornis gen. et sp. nov. The genus Bathycyclopora gen. nov. is introduced for ?Phylactella vibraculata Calvet from the Azores, and also includes Bathycyclopora suroiti gen. et sp. nov. The type species of Calvetopora gen. nov. is Lepralia inflata Calvet from the Gulf of Cadiz; this genus also includes Calvetopora otapostasis gen. et sp. nov. and another species left in open nomenclature. Of the 13 species described herein, 11 occur on seamounts and islands, and nine species are endemic to a single seamount, island or station. The present results show that bryozoans provide striking examples of the function of seamounts as areas of endemism, most likely intrinsically linked to the low dispersal abilities of bryozoan larvae.
Ana C.S. Almeida
Full Text Available ABSTRACT Bryozoans constitute an important component of marine-fouling communities of anthropogenic substrata. Many species have been reported as exotic or widespread around the world, typically in ports and harbors of non-polar regions. Here we present the first record of a species of the bryozoan Triphyllozoon in the Atlantic Ocean. Triphyllozoon arcuatum (MacGillivray, 1889, described originally from Australia, is reported herein from natural substrata in Singapore and natural and artificial substrata in Brazil. Although easily recognizable, the species has not been previously reported from anywhere else in the Atlantic. In the latter instance, the species was collected during monitoring of the invasive scleractinian corals Tubastraea spp. on an oil platform originally from Singapore and now located at Todos os Santos Bay, northeastern Brazil. Colonies of T. arcuatum were also found associated with three species of sponges, giving evidence that it is also growing in the natural environment. Todos os Santos Bay is characterized by intense commercial shipping traffic and oil exploration and the finding of T. arcuatum on an oil platform provides strong evidence that it represents a non-indigenous species in the Atlantic. Owing to the possible impact of T. arcuatum in Brazil, further studies and monitoring of its bioinvasion are recommended.
Schwaha, Thomas F; Wanninger, Andreas
Serotonin represents an evolutionary ancient neurotransmitter that is ubiquitously found among animals including the lophotrochozoan phylum Bryozoa, a group of colonial filter-feeders. Comparatively little is known on their nervous system, and data on their serotonin-lir nervous system currently are mostly limited to the basal phylactolaemates. Previous investigations indicated a common ground-pattern of the serotonin-lir nervous system in these animals, but in order to assess this on a larger scale, 21 gymnolaemate species from 21 genera were comparatively analysed herein. Twenty-one species from 21 gymnolaemate genera were analysed by immunocytochemical stainings and confocal laser scanning microscopy. In all species the serotonin-lir signal is concentrated in the cerebral ganglion from where a nerve tract emanates laterally and traverses orally to engulf the foregut. Serotonin-lir perikarya are situated at the base of the tentacles that almost always correspond to the number of tentacles minus two. The oral side in almost all species shows three serotonin-lir perikarya followed by a 'serotonergic gap' that to our knowledge is not reflected in the morphology of the nervous system. Some species show additional serotonin-lir signal in tentacle nerves, visceral innervation and pore complexes. Paludicella articulata is exceptional as it shows signal in the latero-visceral nerves with serotonin-lir perikarya in the esophagus, parts of the tentacle sheath nerves as well as the frontal body wall around the parietal muscle bundles. In general, the serotonin-lir nervous system in the Bryozoa shows a consistent pattern among its different clades with few deviations. Preliminary data on phylactolaemates suggest the presence of a 'serotonergic gap' similar to gymnolaemates. Both show a subset of oral tentacles and the remaining tentacles in gymnolaemates which correspond to the lateral tentacles of phylactolaemates. The lophophoral concavity lacks serotonin-lir perikarya indicating that due to their larger sizes and increased tentacle number, the horse-shoe shaped arrangement could represent an apomorphy of phylactolaemates.
Sorensen, M. V.; Funch, P.; Willerslev, E.
lamina. An earlier suggested close relationship between Cycliophora and Ectoprocta is rejected. The rotifer-like animal Micrognathozoa is placed in the clade Gnathifera, which also contains Gnathostomulida, Seisonida, Acanthocephala and Eurotatoria. Gnathifera is supported by the presence of jaws...... with a unique ultrastructure and cross-striated muscles attached to the jaw elements by epithelial cells. The jaws in Acanthocephala are considered secondarily reduced. In the analysis, Rotifera turns out to be paraphyletic with respect to Acanthocephala....
Koletić, Nikola; Novosel, Maja; Rajević, Nives; Franjević, Damjan
Bryozoans are aquatic invertebrates that inhabit all types of aquatic ecosystems. They are small animals that form large colonies by asexual budding. Colonies can reach the size of several tens of centimeters, while individual units within a colony are the size of a few millimeters. Each individual within a colony works as a separate zooid and is genetically identical to each other individual within the same colony. Most freshwater species of bryozoans belong to the Phylactolaemata class, while several species that tolerate brackish water belong to the Gymnolaemata class. Tissue samples for this study were collected in the rivers of Adriatic and Danube basin and in the wetland areas in the continental part of Croatia (Europe). Freshwater and brackish taxons of bryozoans were genetically analyzed for the purpose of creating phylogenetic relationships between freshwater and brackish taxons of the Phylactolaemata and Gymnolaemata classes and determining the role of brackish species in colonizing freshwater and marine ecosystems. Phylogenetic relationships inferred on the genes for 18S rRNA, 28S rRNA, COI, and ITS2 region confirmed Phylactolaemata bryozoans as radix bryozoan group. Phylogenetic analysis proved Phylactolaemata bryozoan's close relations with taxons from Phoronida phylum as well as the separation of the Lophopodidae family from other families within the Plumatellida genus. Comparative analysis of existing knowledge about the phylogeny of bryozoans and the expansion of known evolutionary hypotheses is proposed with the model of settlement of marine and freshwater ecosystems by the bryozoans group during their evolutionary past. In this case study, brackish bryozoan taxons represent a link for this ecological phylogenetic hypothesis. Comparison of brackish bryozoan species Lophopus crystallinus and Conopeum seurati confirmed a dual colonization of freshwater ecosystems throughout evolution of this group of animals.
Metcalfe, Kristin; Gordon, Dennis P; Hayward, Ellie
Amphibiobeania epiphylla is a new, monotypic taxon of Beaniidae (Cheilostomata) from Darwin, Northern Territory. It is unique among the 6,000 living species of Bryozoa in that it encrusts mainly living tree leaves (chiefly the mangrove Rhizophora stylosa). The consequence of living in such a specialized habitat is that colonies are emergent (subaerial) for a significant part of the tidal cycle-around 12 of every 24 hours during spring tides and for several days during neap tides. Desiccation is prevented or minimized by the high humidity of the habitat and a cohesive coating of silt covering the colony. Zooids are weakly calcified and lie alternately on their left and right sides in a lineal series, with opercula displaced to the outer corner of the distal zooidal rim. Organisms associated with A. epiphylla include a colony-damaging ceratopogonid (Diptera) larva and a tarsonemid mite that may use dead zooidal interiors, beneath the silt crust, for shelter.
Schwaha, Thomas F; Handschuh, Stephan; Ostrovsky, Andrew N; Wanninger, Andreas
-muscular system is more reminiscent of the condition found in Gymnolaemata rather than Phylactolaemata, which supports a close relationship between Cyclostomata and Gymnolaemata. Several characters of C. elegans such as the lateral ganglia or loss of the cardia are probably apomorphic for this species. For the first time, oocytes that surprisingly develop in the pharyngeal wall are reported for this species.
Wong, Yue Him
The most recent phylogenomic study suggested that Bryozoa (Ectoprocta), Brachiopoda, and Phoronida are monophyletic, implying that the lophophore of bryozoans, phoronids and brachiopods is a synapomorphy. Understanding the molecular mechanisms of the lophophore development of the Lophophorata clade can therefore provide us a new insight into the formation of the diverse morphological traits in metazoans. In the present study, we profiled the transcriptome of the Bryozoan (Ectoproct) Bugula neritina during the swimming larval stage (SW) and the early (4 h) and late (24 h) metamorphic stages using the Illumina HiSeq2000 platform. Various genes that function in development, the immune response and neurogenesis showed differential expression levels during metamorphosis. In situ hybridization of 23 genes that participate in the Wnt, BMP, Notch, and Hedgehog signaling pathways revealed their regulatory roles in the development of the lophophore and the ancestrula digestive tract. Our findings support the hypothesis that developmental precursors of the lophophore and the ancestrula digestive tract are pre-patterned by the differential expression of key developmental genes according to their fate. This study provides a foundation to better understand the developmental divergence and/or convergence among developmental precursors of the lophophore of bryozoans, branchiopods and phoronids.
Full Text Available Macro-fouling communities developed on acrylic, aluminum, wood and fiberglass panels were investigated after 4 months exposure in Marina Bandar al Rawdah and Marina Shangri La. Wet weight of biofouling was about 2-fold higher in Marina Bandar Rawdah and different communities were formed on the front and back sides of the panels. Differences between communities on different materials were less pronounced. In the second study, wet weight and community composition of macro-fouling communities on ceramic tiles at the depth of 1 m and 5 m in Marina Bandar al Rawdah were investigated. During 2008 – 2010, there were no differences between biomass of communities, while in 2011 biomass of macro-fouling was higher on tiles at 5 m. In December 2008 the minimal weight (0 kg/m2 and in September 2011 the maximal weight (26.3 kg/m2 of macro-fouling communities were recorded. In total, 27 invertebrate fouling species were found, which mostly (33% belonged to phylum Ectoprocta. Three invasive bryozoan (Bugula neritina, Zoobotryon verticillatum and Schizoporella errata and one invasive tunicate (Ciona intestinalis species were observed. Overall, this study indicates high biofouling pressure in Muscat marinas and suggests necessity of future studies of fouling communities in Oman waters.
Kristensen, Reinhardt Møbjerg
Loriciferans, cycliophorans and micrognathozoans are amongst the latest groups of animals to be discovered. Other than all being microscopic, they have very different body plans and are not closely related. Loriciferans were originally assigned to the Aschelminthes. However, both new molecular and ultrastructural researches have shown that Aschelminthes consist of two unrelated groups, Cycloneuralia and Gnathifera. Cycloneuralia may be included in the Ecdysozoa, including all molting invertebrates, and Gnathifera are more closely related to Platyhelminthes. The phylum Loricifera shares many apomorphic characters (e.g., scalids on the introvert) with both Priapulida and Kinorhyncha, and can be included in the taxon Scalidophora, a subgroup of Cycloneuralia. Cycliophora was originally allied to the Entoprocta and Ectoprocta (Bryozoa) based on ultrastructual research. Subsequent molecular data show they may be related to Rotifera and Acanthocephala, within the taxon Gnathifera. The phylogenetic position of Cycliophora is therefore not settled, and more ultrastructural and molecular data are needed. Micrognathozoa is the most recent major group of animals to be described. They show strong affinities with both Rotifera and Gnathostomulida (within the taxon Gnathifera), especially in the fine structure of the pharyngeal apparatus, where the jaw elements have cuticular rods with osmiophilic cores. Furthermore the micrognathozoans have two rows of multiciliated cells that form a locomotory organ, similar to that seen in some gastrotrichs and interstitial annelids. This character is never seen in Rotifera or in the monociliated Gnathostomulida. Rotifera and Acanthocephala always have a syncytial epidermis (Syndermata). Micrognathozoa lack this characteristic feature. Therefore, they are postulated to be placed basally in the Gnathifera, either as a sister-group to Gnathostomulida or as a sister-group to Rotifera + Acanthocephala.
Gawin, Natalie; Wanninger, Andreas; Schwaha, Thomas
Phylactolaemata is commonly regarded the earliest branch within Bryozoa and thus the sister group to the other bryozoan taxa, Cyclostomata and Gymnolaemata. Therefore, the taxon is important for the reconstruction of the bryozoan morphological ground pattern. In this study the myoanatomy of Pectinatella magnifica, Cristatella mucedo and Hyalinella punctata was analysed by means of histology, f-actin staining and confocal laser-scanning microscopy in order to fill gaps in knowledge concerning the myoanatomy of Phylactolaemata. The retractor muscles and muscles of the aperture, gut, body wall, tentacle sheath, lophophore constitute the most prominent muscular subsets in these species. The lophophore shows longitudinal muscle bands in the tentacles, lophophoral arm muscles, epistome musculature and hitherto undescribed muscles of the ring canal. In general the muscular system of the three species is very similar with differences mainly in the body wall, tentacle sheath and epistome. The body wall contains an orthogonal grid of musculature. The epistome exhibits either a muscular meshwork in the epistomal wall or muscle fibers traversing the epistomal cavity. The whole tentacle sheath possesses a regular mesh of muscles in Pectinatella and Cristatella, whereas circular muscles are limited to the tentacle sheath base in Hyalinella. This study is the first to describe muscles of the ring canal and contributes to reconstructing muscular features for the last common ancestor of all bryozoans. The data available suggest that two longitudinal muscle bands in the tentacles, as well as retractor muscles and longitudinal and circular muscles in the tentacle sheath, were present in the last common bryozoan ancestor. Comparisons among bryozoans shows that several apomorphies are present in the myoanatomy of each class- level taxon such as the epistomal musculature and musculature of the lophophoral arms in phylactolaemates, annular muscles in cyclostomes and parietal muscles in