Article

Phylum Bryozoa Ehrenberg, 1831

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Abstract

This paper is an invited contribution to the Zootaxa series ‘Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness.’ (Zhang 2011). This series pertains to living biodiversity and the species numbers given here are more or less derived from the World Register of Marine Species (WoRMS), for which the two authors of this paper are listed as editors (PEB hands-on editor; DPG advisory editor). Thus circumscribed, Bryozoa includes 3 classes, 4 orders, 187 families, 808 genera and 5869 species [Phylactolaemata 86 species, Stenolaemata 543 species, Gymnolaemata 5240 species (Ctenostomata 319 species, Cheilostomata 4921 species)]. Although all Phylactolaemata and several species of Ctenostomata are freshwater organisms, the full list of bryozoan species is accessible on WoRMS. The WoRMS list must be understood to be provisional, owing to the need for taxonomic revisions of many genera. Nevertheless, even if not all species in the list are accurately attributed to genera, the names represent operational taxonomic units (OTUs) that are indicative of living species diversity as known to date.

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... Our target organisms are cheilostomes, the most species-rich order of the phylum Bryozoa, with ca. 6,500 described extant species, representing about 80% of the living species diversity of the phylum (Bock & Gordon, 2013). Cheilostomes are lightly to heavily calcified, sessile, colonial metazoans common in benthic marine habitats. ...
... Systematic relationships among cheilostome bryozoans remain largely based on morphological characters (Bock & Gordon, 2013) with molecular phylogenies being restricted to recently collected, ethanol-preserved samples where genetic data were obtained using PCR-based methods (Fuchs et al., 2009;Knight et al., 2011;Waeschenbach et al., 2012), or more recently, by a HTS genome-skimming approach . While these studies have improved our understanding of the phylogenetic relationships among cheilostomes, many key taxa, potentially filling important phylogenetic positions, are hard to procure and remain unfeatured in sequencing projects. ...
... We utilized de novo and iterative methods to assemble our cheilostome mitochondrial sequence data to overcome the challenges of reference-based assemblies, due to the high degree of observed sequence variability among cheilostomes. This is especially important because systematic relationships remain largely based on morphological characters for cheilostomes (Bock & Gordon, 2013) which we know can sometimes be misleading ; hence, reference choice is fraught with difficulties. By using de novo assembled sequences from the target colony itself as a reference for iterative mapping, we circumvent the difficulties of reference choice. ...
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Article
Resolution of relationships at lower taxonomic levels is crucial for answering many evolutionary questions, and as such, sufficiently varied species representation is vital. This latter goal is not always achievable with relatively fresh samples. To alleviate the difficulties in procuring rarer taxa, we have seen increasing utilization of historical specimens in building molecular phylogenies using high throughput se-quencing. This effort, however, has mainly focused on large-bodied or well-studied groups, with small-bodied and under-studied taxa under-prioritized. Here, we utilize both historical and contemporary specimens, to increase the resolution of phylo-genetic relationships among a group of under-studied and small-bodied metazoans, namely, cheilostome bryozoans. In this study, we pioneer the sequencing of air-dried cheilostomes, utilizing a recently developed library preparation method for low DNA input. We evaluate a de novo mitogenome assembly and two iterative methods, using the sequenced target specimen as a reference for mapping, for our sequences. In doing so, we present mitochondrial and ribosomal RNA sequences of 43 cheilos-tomes representing 37 species, including 14 from historical samples ranging from 50 to 149 years old. The inferred phylogenetic relationships of these samples, analyzed together with publicly available sequence data, are shown in a statistically well-supported 65 taxa and 17 genes cheilostome tree, which is also the most broadly sampled and largest to date. The robust phylogenetic placement of historical samples whose contemporary conspecifics and/or congenerics have been sequenced verifies the appropriateness of our workflow and gives confidence in the phylogenetic placement of those historical samples for which there are no close relatives sequenced. The success of our workflow is highlighted by the circularization of a total of 27 mi-togenomes, seven from historical cheilostome samples. Our study highlights the potential of utilizing DNA from micro-invertebrate specimens stored in natural history collections for resolving phylogenetic relationships among species.
... Among the extant bryozoan suborders, Cheilostomata Busk, 1852a accounts for about 90% of all species, which are classified into about 139 families and 700 genera (e.g. Bock & Gordon 2013). It is the most diverse clade, dominating in respect of both zooidal and colony morphologies and being among the most abundant marine epibiotic invertebrates colonizing a wide range of substrata. ...
... Several classification concepts for Cheilostomata have been proposed more recently (e.g. d 'Hondt 1985a, Viskova & Morozova 1988, Bock & Gordon 2013, d'Hondt 2016, but none of these concepts could gain universal acclaim, while Voigt (1991) showed that a systematic separation relying on the traditionally used groups to subdivide Cheilostomata is arbitrary as the characteristic morphological features associated with each group may be convergencies. ...
... Therefore, combined molecular and morphological works are needed to better understand phylogenetic significance of morphological characters and homologies among structures. In the light of a wanting comprehensive and stable family-level molecular phylogeny of cheilostome bryozoans, we decided to not push another morphology-based classification but largely follow the classifications of Bock and Gordon (2013), Ostrovsky (2013), andCook et al. (2018) for convenience. However, Skylonina Viskova in Viskova and Morozova (1988), a valid but neglected name for an enigmatic suborder comprising three cheilostome families with mainly (or exclusively) fossil representatives, is reintroduced here. ...
Chapter
The most recent review on cheilostome morphology, ecology and systematics. Chapter includes the following sections: General morphology (General morphology, General structure, Structure of frontal walls and mechanisms of polypide eversion, Polymorphism, A short summary of cheilostome phylogeny); Ecology of cheilostome bryozoans (Habitat and substratum selection, Diet of cheilostomes, Predation on Cheilostomata); Systematics of Cheilostomata & Literature.
... The great majority of bryozoans grow calcified zooids, and in such cases taxonomy is based primarily on the morphology of the skeleton. Approximately 6600 living species have been described, as well as over 15000 extinct species, dating back as far as the early Paleozoic (McKinney & Jackson, 1989;Bock & Gordon, 2013;Orr et al., 2018). The extensive, diverse and character-rich fossil record of the Bryozoa has made this phylum an important group for stratigraphic studies, palaeoenvironmental analyses and for testing ecological hypotheses and evolutionary theories, Biological Reviews (2020) 000-000 © 2020 Cambridge Philosophical Society such as punctuated equilibrium and clade replacement (Cuffey, 1967;Nelson et al., 1988;Jackson & Cheetham, 1994;McKinney et al., 1998;Liow & Taylor, 2019). ...
... Six of these orders are extinct, collectively comprising the superorder Paleostomata (Ma, Buttler & Taylor, 2014), with~10000-15000 species. Notably, the only extant order, the Cyclostomata, contains about 550 living species (Bock & Gordon, 2013). All stenolaemates secrete robust skeletons composed of calcite, and are well calcified, maintaining their shape and integrity following immersion in strong bleach (NaOCl). ...
... The class Gymnolaemata contains the order Cheilostomata, with~5000 living species (Bock & Gordon, 2013). The cheilostomes are a relatively modern group, first appearing in the Late Jurassic, and radiating in the Late Cretaceous (Jablonski, Lidgard & Taylor, 1997). ...
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Article
Skeletal resorption-the physiological removal of mineralised parts by an organism-is an important morphoge-netic process in bryozoans. Reports of its occurrence and function across the phylum are patchy, however, and have not previously been synthesised. Here we show that resorption occurs routinely across a wide range of bryozoan clades, colony sizes, growth forms, ontogenetic stages, body wall types, skeletal ultrastructures and mineralogies. Beginning in the early Paleozoic, different modes and functions of resorption have evolved convergently among dis-parate groups, highlighting its utility as a morphogenetic mode in this phylum. Its functions include branch renovation , formation of branch articulations, excavation of reproductive chambers, part-shedding, and creation of access portals for budding beyond previously formed skeletal walls. Bryozoan skeletons can be altered by resorption at microscopic, zooidal and colony-wide scales, typically with a fine degree of control and coordination. We classified resorption patterns in bryozoans according to the morphology and function of the resorption zone (window formation, abscission or excavation), timing within the life of the skeletal element resorbed (primary or secondary), and scale of operation (zooidal or multizooidal). Skeletal resorption is probably greatly underestimated in terms of its utility and role in bryozoan life history, and its prevalence across taxa, especially in fossil forms. It is reported proportionally more frequently in stenolaemates than in gymnolaemates. Some modes of resorption potentially alter or remove the spatial-temporal record of calcification preserved within a skeleton. Consequently, knowledge that resorption has occurred can be relevant for some common applications of skeletal analysis, such as palaeoen-vironmental interpretation, or growth and ageing studies. To aid recognition we provide scanning electron micros-copy, backscattered electron scanning electron microscopy and transmission electron microscopy examples of skeletal ultrastuctures modified by resorption.
... Bryozoa (Ectoprocta) is an aquatic phylum that comprises more than 6000 described recent and 15000 fossil species of epibiotic, active suspension-feeding coelomate invertebrates (Gordon, Taylor & Bigey, 2009;Bock & Gordon, 2013). Bryozoans are known from the beginning of the Ordovician and represent major components of most benthic ecosystems from the intertidal to abyssal depths exceeding 8000 m. ...
... In fact, the high diversity of bryozoans through geological time is an important indicator of their evolutionary success. Three larger taxa are commonly recognized among Bryozoa: (i) the solely freshwaterinhabiting Phylactolaemata; (ii) the marine Stenolaemata with the only surviving taxon Cyclostomata; and (iii) the predominantly marine and morphologically most diverse Gymnolaemata (Bock & Gordon, 2013). Allman's (1856) original classification divided Bryozoa into Phylactolaemata [phylassoto guard, laimosthroat (referring to the epistome, a ciliated flap 'guarding' or covering the mouth opening] and Gymnolaemata ['naked throats' (gymnosnaked) with respect to the missing epistome] which initially included three groups defined by Busk (1852) as Cyclostomata (cycloring, stomamouth), Biological Reviews (2020) 000-000 © 2020 The Authors. ...
... influenced the diversity of these groups. However, two ctenostome superfamilies (Alcyonidioidea, Walkerioidea) without such connections show higher diversity with respect to their number of families and species than ctenostomes whose stolonal and autozooidal funicular elements are interconnected (Vesicularioidea, Victorelloidea, Nolellidae of Arachnidioidea) (Jebram, 1986b;Bock & Gordon, 2013). Thus, the evolution of funicular connectivity between zooids appears not to have a pivotal role in ctenostome diversification. ...
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Article
Molecular techniques are currently the leading tools for reconstructing phylogenetic relationships, but our understanding of ancestral, plesiomorphic and apomorphic characters requires the study of the morphology of extant forms for testing these phylogenies and for reconstructing character evolution. This review highlights the potential of soft body morphology for inferring the evolution and phylogeny of the lophotrochozoan phylum Bryozoa. This colonial taxon comprises aquatic coelomate filter‐feeders that dominate many benthic communities, both marine and freshwater. Despite having a similar bauplan, bryozoans are morphologically highly diverse and are represented by three major taxa: Phylactolaemata, Stenolaemata and Gymnolaemata. Recent molecular studies resulted in a comprehensive phylogenetic tree with the Phylactolaemata sister to the remaining two taxa, and Stenolaemata (Cyclostomata) sister to Gymnolaemata. We plotted data of soft tissue morphology onto this phylogeny in order to gain further insights into the origin of morphological novelties and character evolution in the phylum. All three larger clades have morphological apomorphies assignable to the latest molecular phylogeny. Stenolaemata (Cyclostomata) and Gymnolaemata were united as monophyletic Myolaemata because of the apomorphic myoepithelial and triradiate pharynx. One of the main evolutionary changes in bryozoans is a change from a body wall with two well‐developed muscular layers and numerous retractor muscles in Phylactolaemata to a body wall with few specialized muscles and few retractors in the remaining bryozoans. Such a shift probably pre‐dated a body wall calcification that evolved independently at least twice in Bryozoa and resulted in the evolution of various hydrostatic mechanisms for polypide protrusion. In Cyclostomata, body wall calcification was accompanied by a unique detachment of the peritoneum from the epidermis to form the hydrostatic membraneous sac. The digestive tract of the Myolaemata differs from the phylactolaemate condition by a distinct ciliated pylorus not present in phylactolaemates. All bryozoans have a mesodermal funiculus, which is duplicated in Gymnolaemata. A colonial system of integration (CSI) of additional, sometimes branching, funicular cords connecting neighbouring zooids via pores with pore‐cell complexes evolved at least twice in Gymnolaemata. The nervous system in all bryozoans is subepithelial and concentrated at the lophophoral base and the tentacles. Tentacular nerves emerge intertentacularly in Phylactolaemata whereas they partially emanate directly from the cerebral ganglion or the circum‐oral nerve ring in myolaemates. Overall, morphological evidence shows that ancestral forms were small, colonial coelomates with a muscular body wall and a U‐shaped gut with ciliary tentacle crown, and were capable of asexual budding. Coloniality resulted in many novelties including the origin of zooidal polymorphism, an apomorphic landmark trait of the Myolaemata.
... Their skeletal features are finely preserved over large timescales, to the point where fossil taxa can often be identified at the species level (Jackson & Cheetham, 1990;Simpson & Jackson, 2022). Order Cheilostomatida, in particular, represents the most abundant and diverse order within Bryozoa, containing around 80% of the phylum's living species diversity plus as many as 7,900 described fossil species (Bock & Gordon, 2013). ...
... Most notably, DeepBryo currently focuses on Cheilostomatida. While cheilostomes are the most diverse order within Bryozoa (Bock & Gordon, 2013;Taylor, 2020), containing around 80% of extant species, other orders are also mineralized and could be included in future model retraining. Therefore, future work will concentrate on increasing taxonomic coverage and expanding the app to other bryozoan orders. ...
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Preprint
Bryozoans are becoming an increasingly popular study system in macroevolutionary, ecological, and paleobiological research. Members of this colonial invertebrate phylum are notable for displaying an exceptional degree of division of labor in the form of specialized modules (polymorphs), which allow for the inference of individual allocation of resources to reproduction, defense, and growth using simple morphometric tools. However, morphometric characterizations of bryozoans are notoriously labored, due to the high number of structures often captured per image, as well as the need for specialized knowledge necessary for classifying individual skeletal structures within those images. We here introduce DeepBryo, a web application for deep learning-based morphometric characterization of cheilostome bryozoans. DeepBryo requires a single image as input and performs measurements automatically using instance segmentation algorithms. DeepBryo is capable of detecting objects belonging to six classes and outputting fourteen morphological shape measurements for each object based on the inferred segmentation maps. The users can visualize the predictions, check for errors, and directly filter model outputs on the web browser. Measurements can then be downloaded as a comma-separated values file. DeepBryo has been trained and validated on a total of 72,412 structures, belonging to six different object classes in 935 SEM images of cheilostome bryozoans belonging to 109 different families. The model shows high (>0.8) recall and precision for zooid-level structures. Its misclassification rate is low (~4%) and largely concentrated in a single object class (opesia). The model’s estimated structure-level area, height, and width measurements are statistically indistinguishable from those obtained via manual annotation (r ² varying from 0.89 to 0.98) and show no detectable bias. DeepBryo reduces the person-hours required for characterizing the zooids in individual colonies to less than 1% of the time required for manual annotation at no significant loss of measurement accuracy. Our results indicate that DeepBryo enables cost-, labor,- and time-efficient morphometric characterization of cheilostome bryozoans. DeepBryo can greatly increase the scale of macroevolutionary, ecological, taxonomic, and paleobiological analyses, as well as the accessibility of deep learning tools for this emerging model system. Finally, DeepBryo provides the building blocks necessary for adapting the current application to other study groups.
... Bryozoa is a phylum of aquatic, sessile, modular suspension feeders, comprising over 6500 Recent species distributed across all latitudes and oceans (Bock and Gordon, 2013;Schwaha, 2020), with over 300 species known globally from waters deeper than 700 m (Gordon, 1987(Gordon, , 2014. Bryozoans occur across a wide spectrum of deep-sea environments, including abyssal plains (Grischenko and Chernyshev, 2015;d'Hondt, 1975d'Hondt, , 1985d'Hondt, , 1987d'Hondt and Hayward, 1981;d'Hondt and Schopf, 1985), deep-sea trenches (Grischenko et al., 2019;Hayward, 1981), ridges (Gordon, 1984(Gordon, , 1985(Gordon, , 2021Grischenko et al., 2020;Matsuyama et al., 2014;Souto and Albuquerque, 2019), enclosed deep-water basins (Grischenko, 2013;Grischenko and Chernyshev, 2018), cold seeps (Jollivet et al., 1990), hot hydrothermal vent fields associated with active submarine volcanoes (Gordon, 2012), massive seabed sulphides (Gerdes et al., 2021), seamounts (Berning et al., 2017;Gordon, 2014Gordon, , 2021Gordon and Taylor, 2010;Harmelin, 1977;Hayward, 1979;Ramalho et al., 2020;Souto et al., 2016), and polymetallic-nodule bottoms (Amon et al., 2017;Grischenko et al., , 2021Jones et al., 2021;Schwaha et al., 2020aSchwaha et al., , 2021. ...
... Holotype specimens are deposited in the Zoological Institute of the Russian Academy of Sciences (ZIRAS), Saint Petersburg, Russia. The classification used follows Bock and Gordon (2013). The benthic depth zonation follows Watling et al. (2013). ...
Article
The sessile benthic fauna on ferromanganese (Fe-Mn) crusts is poorly known, with many groups of invertebrates still barely examined. Collections made during Yuzhmorgeologiya expeditions on RV Gelendzhik in October 2016 (YMG 6–16) and October 2020 (YMG 6–19) to the Magellan Seamounts, tropical western Pacific, have yielded two new species and a new genus of cheilostome bryozoans. These are Allantocallopora makarikhini sp. nov., obtained at Stn 14B135 from the Kotzebue Guyot, 17.3800°N, 153.0800°E, at 1505 m depth, and Melnikum magellanum gen. et sp. nov., collected at Stn 08D108 from the Govorov Guyot, 17.8984°N, 151.2415°E, at 2036 m depth. Colonies of these new species were attached to the surface of cobalt-rich Fe-Mn crusts, representing a rare example of detection of Bryozoa on Fe-Mn crusts in the deep-sea. A literature review revealed 23 reports documenting bryozoans on the surfaces of Fe-Mn crusts or the deposition of Fe-Mn oxides on the surface of bryozoan colonies, defined as ferrugination. Fe-Mn crust-associated Bryozoa have been found at 29 localities in ten regions of the world’s oceans, in a depth range of 400–5850 m, with the majority of finds made in the central to southwestern Pacific. Most of these reports were based on preliminary data; they identified nine bryozoan taxa to the species level. Fe-Mn crust-associated bryozoans show low local species richness, up to four species per locality, and generally low abundance. Analyses of published data suggest that this apparent low richness might be due to the short history of investigations of crusts and strongly affected by the sampling method. Thorough inventories of Fe-Mn crust-associated bryozoans are urgently needed; bryozoans are a component of the sessile biota, which is sensitive and vulnerable to the proposed mining of Fe-Mn crusts as an important mineral resource in the deep sea.
... In this contribution, we begin to rectify the paucity of large and/or taxonomically broadly sampled molecular phylogenies for marine invertebrates, targeting a phylum whose rich fossil record can be subsequently integrated for evolutionary analyses. Our focal group is Cheilostomatida, the dominant living order of the colonial metazoan phylum Bryozoa, with c. 5200 described extant species, corresponding to >80% of the living species diversity of the phylum (Bock and Gordon, 2013). Cheilostomes first appeared in the fossil record in the Late Jurassic (c. ...
... Most species are encrusting, while fewer are erect, with some forming robust structures whereas many are small and inconspicuous (Fig. 1). Although a number of cheilostome bryozoans have been sequenced and placed in a molecular phylogenetic context (Fuchs et al., 2009;Knight et al., 2011;Orr et al., 2019a;Waeschenbach et al., 2012) the systematics of cheilostome bryozoans aimed at reflecting their evolutionary relationships still remain largely based on morphological characters (Bock and Gordon, 2013;Martha et al., 2020;Taylor and Waeschenbach, 2015). This is in part because assumed cheilostome phylogenetic relationships have only recently benefited from high-throughput sequencing (HTS) techniques and the increased phylogenetic support it provides (Orr et al., 2019a(Orr et al., , b, 2020. ...
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Article
Larger molecular phylogenies based on ever more genes are becoming commonplace with the advent of cheaper and more streamlined sequencing and bioinformatics pipelines. However, many groups of inconspicuous but no less evolutionarily or ecologically important marine invertebrates are still neglected in the quest for understanding species- and higher-level phylogenetic relationships. Here, we alleviate this issue by presenting the molecular sequences of 165 cheilostome bryozoan species from New Zealand waters. New Zealand is our geographic region of choice as its cheilostome fauna is taxonomically, functionally and ecologically diverse, and better characterized than many other such faunas in the world. Using this most taxonomically broadly-sampled and statistically-supported cheilostome phylogeny comprising 214 species, when including previously published sequences, and 17 genes (2 nuclear and 15 mitochondrial) we tested several existing systematic hypotheses based solely on morphological observations. We find that lower taxonomic level hypotheses (species and genera) are robust while our inferred trees did not reflect current higher-level systematics (family and above), illustrating a general need for the rethinking of current hypotheses. To illustrate the utility of our new phylogeny, we reconstruct the evolutionary history of frontal shields (i.e., a calcified body-wall layer in ascus-bearing cheilostomes) and ask if its presence has any bearing on the diversification rates of cheilostomes.
... The phylogenetic placement and systematic affinity of Haywardozoon is not clear. Originally assigned to the alcyonidioidean Flustrellidridae (Hayward, 1978), it was subsequently shifted to a separate genus, family and superfamily, presumably closely related to hislopioidean ctenostomes (see Bock & Gordon, 2013;d'Hondt, 1983). However, no detailed morphological analysis has been undertaken for any of these three species and their systematic position remains poorly defined. ...
... However, only few alcyonidiid species have been properly analyzed with respect to their apertural musculature. The genus Hislopia, of the Hislopioidea, to which Haywardozoon has often been associated to (e.g., Bock & Gordon, 2013), differ in their apertural muscles. Besides vestibular wall muscles that are, due to the thick cuticle, apparently missing in H. pacificum, they possess four prominent apertural bundles inserting at the diaphragm and large areas of the vestibular wall. ...
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Article
The genus Haywardozoon represent a little known genus of ctenostome bryozoans that has only been found in the deep‐sea. It forms small, mostly uniserial colonies lacking polymorphs. Zooids have a conspicuous apertural closure mechanism consisting of a cuticular lower lip that closes the aperture. The systematic placement of the genus remains uncertain, detailed morphological studies that include soft‐body morphological traits are missing. Consequently, this is the first study analyzing H. pacificum by means of histological serial sections and 3d‐reconstruction. Zooids are ovoid and in some cases solitary, that is, showing no interconnected zooids. Most prominent is the large vestibular wall that can be more than half of the total length of the zooid. Its vestibular wall is particularly lined by a complex, multilayered and branched cuticle. A single pair of lateral parieto‐diaphragmatic muscles is present. The polypide is small and comprises about 17 tentacles. The digestive tract is short, has an elongated cardia, a vestigial caecum and a vestibular anus. An ovipositor/intertentacular organ and several oligolecithal oocytes were detected. Several aspects of zooidal morphology, including the structure of the bilateral aperture, parieto‐diaphragmatic muscles, general structure of the gut and the thick cuticle, clearly indicate an association to the ctenostome superfamily Alcyonidioidea. Therefore, we reject the previous placement into Hislopioidea and suggest a possible association to pherusellid ctenostomes. New reproductive characters show that H. pacificum is a broadcaster contrary to some other deep‐sea forms that are brooding. Research Highlight Morphology of ctenostome bryozoans remain little investigated. This contribution is the second of a series of detailed morphological analyses of this understudied clade of bryozoans. The morphological investigation of Haywardozoon pacificum revealed numerous characters that show a closer relationship to Flustrellididrae rather than Hislopiidae as previously assumed.
... The lophotrochozoan phylum Bryozoa comprises about 5900 extant and more than 15,000 extinct species (Bock and Gordon 2013). As aquatic sessile animals, they colonize various substrates in predominantly marine environments with a small fraction restricted to freshwater habitats. ...
... Within the Gymnolaemata, the cheilostomes account for the majority of bryozoan species. With~5000 described species, cheilostomes are the most diverse and abundant group of bryozoans and possess a calcareous skeleton and a characteristic operculum (Ryland 2005;Bock and Gordon 2013). In contrast, only~350 species have been described from the uncalcified ctenostome Gymnolaemata (Schwaha 2019b). ...
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Article
The epiphytic community on the endemic seagrass Posidonia oceanica from the Mediterranean Sea is well studied, but still harbors some little investigated epiphytic bryozoans. Numerous, yet always small colonies of Pherusella sp. were recently encountered in the Northern Adriatic Sea. The aim of this study was to generate data on the life history, colonial development, and reproduction of the Mediterranean population of this Pherusella species in order to gain a better understanding of the biology of this understudied species. The morphology of adult zooids was also studied due to the lack of recent data on the family with state-of-the-art techniques. Long-term observation shows that this species is highly adapted to an epiphytic life cycle with short generation time throughout the year. First laboratory cultures appear promising in establishing a reliable model system for developmental and ecological studies. Larvae are easily obtainable, and metamorphosis and colonial growth patterns are documented here for the first time. The morphology of adults shows distinct similarities with other pheruselllids and, along with the neuromuscular system, is similar to other alcyonidioideans supporting the close relationship of these taxa. This study constitutes one of the first long-time observations of the life cycle and colonial growth of a pherusellid bryozoan, including morphological data about the neuromuscular system of an otherwise incompletely known group of bryozoans. Pherusella sp. appears to be a promising candidate for future studies since it is easy to collect and maintain under laboratory conditions as well as to obtain different developmental stages.
... All bryozoans are suspension feeders, which means their lophophores capture organic particles out of the water column. Bryozoans comprise a comparatively poorly studied group despite their diversity (~6000 described extant species) [13]). They are ecologically important suspension feeders found in freshwater, brackish, and marine environments. ...
... Bryozoans are lophotrochozoan animals which are composed of three classes: Phylactolaemata, Stenolaemata, and Gymnolaemata, the latter comprising the orders Ctenostomatida and Cheilostomatida [21,22]. The least diverse of these is the Phylactolaemata, which comprises ~86 species, all of which are uncalcified and live in freshwater habitats [13]. The remaining species live either exclusively (Stenolaemata) or mostly (Gymnolaemata) in marine habitats, and most produce a calcium carbonate skeleton. ...
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Article
Less than one percent of marine natural products characterized since 1963 have been obtained from the phylum Bryozoa which, therefore, still represents a huge reservoir for the discovery of bioactive metabolites with its ~6000 described species. The current review is designed to highlight how bryozoans use sophisticated chemical defenses against their numerous predators and competitors, and which can be harbored for medicinal uses. This review collates all currently available chemoecological data about bryozoans and lists potential applications/benefits for human health. The core of the current review relates to the potential of bryozoan metabolites in human diseases with particular attention to viral, brain, and parasitic diseases. It additionally weighs the pros and cons of total syntheses of some bryozoan metabolites versus the synthesis of non-natural analogues, and explores the hopes put into the development of biotechnological approaches to provide sustainable amounts of bryozoan metabolites without harming the natural environment.
... Bryozoans are a phylum of aquatic invertebrates containing almost 6000 described species (Bock and Gordon, 2013) and are one of the most common and most abundant marine invertebrates in the benthic ecosystems (Bock, 1982;. They are suspension-feeding colonial invertebrates that are widely distributed across the globe, inhabiting most types of marine ecosystems . ...
... They are suspension-feeding colonial invertebrates, widely distributed, inhabiting most types of aquatic ecosystems. represented in the fossil record; and Gymnolaemata are predominantly marine and comprise the majority of extant species (Bock and Gordon, 2013). ...
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Thesis
Marine ecosystems are directly threatened by multiple and interactive human stressors at global and local scales. Hence, it is vital to study biodiversity and ecological patterns through a multi-disciplinary approach, from understanding global diversity patterns to evaluating the ecological responses of species to different impacts in order to protect marine ecosystems. In this thesis, we focused on bryozoans, an abundant group of sessile marine invertebrates distributed worldwide, but generally understudied. Accordingly, in this thesis we provide different approaches to understand discovery and macroecological patterns at global scales, and the response of species to different stressors at local scales, combining the use of open databases, the in situ monitoring of natural populations, experiments in aquaria and the development of restoration techniques. At global scale, in Chapter I we unraveled discovery patterns of fossil and extant bryozoans and showed the highest number of fossil species described, highlighting that the current biodiversity represents only a small proportion of Earth’s past biodiversity. Beyond these differences, both groups showed an increase in the taxonomic effort during the past century. Despite this progress, future projections of discovery patterns of both groups showed a large proportion of species remaining to be discovered by the final of this century. In Chapter II, most of the global diversity patterns of marine sessile groups, including bryozoans, showed a non-unimodal latitudinal pattern with a dip in the number of species at the equator and a higher diversity in the Southern ocean. Moreover, this region will be less affected by global warming at the final of this century. In contrast, our analyses showed that the most sampled region for both marine sessile species and bryozoans was North Temperate Atlantic, highlighting the importance to quantify environmental drivers considering sampling effort biases. For this reason, we tested the effect of using the popular method of rarefaction (ES50) vs the incorporation of a frequency index of sampling effort as co-variate in quantitative models. Despite we obtained the same best predictors for both approaches (depth, nitrate, and SST), the models using the correction of sampling biases through frequency index showed better fitting, encouraging to incorporate this methodology in future studies. Focusing on the Mediterranean Sea, in Chapter 3 we studied the responses of bryozoans to different stressors. First, we showed that two abundant and common bryozoans, Pentapora fascialis and Myriapora truncata, displayed different tolerances to warming through the combination of in situ monitoring and experiments in aquaria. Moreover, in Chapter 4 the in situ monitoring of Pentapora fascialis populations revealed its fast population dynamics, with high recruitment and growth rates, and a high capacity of recovery. Accordingly, we observed an increase in the density of its populations in the Medes Island Marine Reserve since the 1990s. However, we evidenced that diving can impact on the density, recruitment, survival, and size of the colonies, registering lower values in frequented localities. Our results highlight that the over frequentation of divers compromises the future viability of populations, highlighting the need to explore other active management strategies. For this reason, in Chapter 5, we tested and developed different restoration techniques for P. fascialis, focusing on the recruitment enhancement through the installation of recruitment surfaces and the transplantation of adult colonies. The successful results and the affordable and economic cost of tested techniques aim to encourage the managers of Marine Protected Areas to apply similar methodologies. The results presented in this thesis show the importance to combine different approaches to understand the global and local ecological patterns of understudied but abundant groups, such as bryozoans. Our findings enlarge the current ecological knowledge of bryozoans at different scales, and highlight that more effort is needed to protect vulnerable populations.
... Cheilostome bryozoans, though less well-studied than several metazoan clades of similar size, are 56 ubiquitous in benthic marine habitats. They are the most diverse order of Bryozoa with 83% of a 57 conservatively estimated 5869 extant described species (Bock & Gordon, 2013). Bryozoans are 58 ecologically important habitat builders (Wood, Rowden, Compton, Gordon, & Probert, 2013) and are 59 vital components of the marine food chain (Lidgard, 2008). ...
... have species-location information from 3323 species or 68% of 4921 described cheilostome species 172 (Bock & Gordon, 2013). 173 ...
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Aim: We compare and combine data from a public georeferenced biodiversity database and from the published literature in order to identify strengths and gaps in global marine biogeographic knowledge. Using these data, we estimate the latitudinal species diversity distribution for a commonly occurring but under-studied clade, cheilostome Bryozoa, which has long been hypothesized to show a non-canonical latitudinal diversity gradient (LDG). Location: Global. Major taxa studied: Cheilostomata, Bryozoa, with around 5000 described extant species. Methods: We use natural language processing (NLP) to retrieve location data of cheilostome species (text-mined occurrences [TMO]) in an automated procedure. We compare and combine these results with data from the Ocean Biogeographic Information System (OBIS). Using OBIS and TMO data separately and in combination, we present latitudinal species richness curves using standard estimators (Chao2 and the Jackknife) and range-through approaches. Results: Our combined OBIS and TMO species richness curves quantitatively document a bimodal global latitudinal diversity gradient for cheilostomes for the first time, with peaks in the temperate zones. 79% of the georeferenced species we retrieved from TMO (N = 1780) and OBIS (N = 2453) are non-overlapping and underestimate known species richness, even in combination. Main conclusions: Despite clear indications that global location data compiled for cheilostome bryozoans should be improved with concerted effort, our study supports the view that latitudinal species richness patterns deviate from the canonical LDG. Moreover, combining online biodiversity databases with automated information retrieval from the published literature is a promising avenue for expanding taxon-location datasets.
... Several views on the higher taxa within this group had also been expressed before the 2000s (Kluge, 1962;Cuffey, 1973;Ryland, 1982;Kubanin, 1984;d'Hondt, 1985;Viskova and Morozova, 1988). By the early 2010s, the available information had been streamlined (Bock and Gordon, 2013); great efforts have been undertaken to consolidate synonymy of taxa of different ranks in recent years (Bock, 2020). Presently, the systematic affiliation and classification of the higher taxa can be considered to be well established (Phylum Bryozoa…, 2020), allowing comparative analysis of the taxonomic structure of regional faunas of Arctic bryozoans. ...
... Bryozoans are primarily sessile colonial invertebrates (5869 extant species), and most common fouling organisms encountered in modern oceans (Gordon et al., 2007;Bock and Gordon, 2013;Rouse et al., 2013) whilst a few species are even found in freshwaters. However, they show profound abundance in the shelf waters up to 500m of water depths (Cook, 1981;Gordon, 1986Gordon, , 1999Wood et al., 2012;Smith, 2014). ...
Article
The ecological record of bryozoans from the continental shelf-slope region of the southwestern Bay of Bengal is very sparse, prompting this investigation. Forty-five sea-floor sediment samples were collected along eight transects during the cruise on R/V Sagar Paschimi from the offshore region between Chennai and Cuddalore in the southwestern Bay of Bengal. The distribution of bryozoans was assessed based on the fragments of bryozoan colonies from these samples, which were collected at depths ranging from 6–308m. The goal was to determine the relationship between environmental parameters measured from the study area and the abundances of various bryozoan species. Results revealed the presence of 29 species of recent bryozoans in 24 genera pertaining to eight morphotypes. It is evident from the present investigation that the major control on the spatial distribution of bryozoan colonies is the type of substrate available, as sample sites dominated by coarse sediments showed high species richness though other factors such as ocean dynamics and rate of sedimentation also seem to regulate their abundance.
... With an estimated diversity of more than 4,900 species in 2013 (Bock & Gordon, 2013), the Cheilostomatida is the most successful group among the Recent bryozoans. Its high species-diversity seems to be related to their variability in the use of substrates and the acquisition of special features such as zooidal polymorphism and complex reproductive patterns (Ryland, 1970). ...
Article
Knowledge of the marine bryozoan fauna of Uruguay is mostly based on scattered records found in local faunistic surveys and the taxonomic results of two oceanographic cruises to the Southwest Atlantic, but a comprehensive study has not yet been published for this area. This paper aims to compile an updated checklist, bringing together all the published information about the Uruguayan bryozoan fauna. Of the 73 recorded taxa, 30 (41%) are known only from deep waters off the Río de la Plata. Even considering undetermined species, these results show the high degree of endemism as it was already shown for several other benthic groups such as Bivalvia, Ascidiacea and Pycnogonida. The absence of local taxonomists on bryozoans has resulted in the unprecedented situation that the deep-sea bryozoan fauna of Uruguay is better known than the coastal and shelf representatives of the phylum. The main conclusion of this faunal compilation is that efforts should be made to coordinate the gathering and taxonomic study of shallow and shelf bryozoan collections to bridge the present knowledge gap about the biodiversity of this important group of marine benthic invertebrates.
... Horneridae is moderately diverse, with~100 described living and fossil species, ranging from the Eocene to the present-day (Harmelin, 2020;Mongereau, 1972;Smith et al., 2008). Five living genera containing~25 valid species are described (see Bock & Gordon, 2013;Grischenko et al., 2018;Smith et al., 2008), although this is likely a significant underestimate of true diversity. ...
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Article
Horneridae (Cyclostomatida: Cancellata) is a family of marine bryozoans that forms tree‐like colonies bearing functionally unilaminate branches. Colony development in this clade is not well understood. We used micro‐CT and SEM to trace zooidal budding in Hornera from the ancestrula onwards. Results show that hornerid branches are constructed by dual zooidal budding modes occurring synchronously at two separate budding sites at the growing tips. Frontal autozooids bud from a multizooidal budding lamina. Lateral autozooids bud from discrete abfrontal budding loci by ‘exomural budding', a previously undescribed form of frontal budding centred on hypostegal pores in interzooidal grooves on the colonial body wall. These two budding modes are integrated during primary branch morphogenesis, forming composite, developmentally bilaminate, branches. Patterns of exomural budding vary among hornerid taxa, and future studies of Cancellata taxonomy and phylogeny may benefit from morphological concepts presented here. This article is protected by copyright. All rights reserved.
... Cheilostomes are also the most species-rich order within Bryozoa, with more than 6000 described extant species and likely about the same number yet to be formally described (11). They represent c. 80% of the phylum's living species diversity (12). Likewise, there are c. ...
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Article
Phylogenetic relationships and the timing of evolutionary events are essential for understanding evolution on longer time scales. Cheilostome bryozoans are a group of ubiquitous, species-rich, marine colonial organisms with an excellent fossil record but lack phylogenetic relationships inferred from molecular data. We present genome-skimmed data for 395 cheilostomes and combine these with 315 published sequences to infer relationships and the timing of key events among c. 500 cheilostome species. We find that named cheilostome genera and species are phylogenetically coherent, rendering fossil or contemporary specimens readily delimited using only skeletal morphology. Our phylogeny shows that parental care in the form of brooding evolved several times independently but was never lost in cheilostomes. Our fossil calibration, robust to varied assumptions, indicates that the cheilostome lineage and parental care therein could have Paleozoic origins, much older than the first known fossil record of cheilostomes in the Late Jurassic.
... It is in fact representatives of the phylum Bryozoa the most commonly found attached to plastic debris, being the most species-rich and the most abundant in specimen number, as resulted in this study. This is expected as bryozoans are abundant and diverse worldwide (>6000 living species; Bock and Gordon, 2013), including the Mediterranean (>550 species; Rosso and Di Martino, 2016), almost exclusively sessile and ubiquitous, being present from the tropics to the poles and from the intertidal to the deep sea (e.g. Figuerola et al., 2012Figuerola et al., , 2018Almeida et al., 2021). ...
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Article
Plastic debris provides long-lasting substrates for benthic organisms, thus acting as a potential vector for their dispersion. Its interaction with these colonizers is, however, still poorly known. This study examines fouling communities on beached, buoyant and benthic plastic debris in the Catalan Sea (NW Mediterranean), and characterizes the plastic type. We found 14 specimens belonging to two phyla (Annelida and Foraminifera) on microplastics, and more than 400 specimens belonging to 26 species in 10 phyla (Annelida, Arthropoda, Brachiopoda, Bryozoa, Chordata, Cnidaria, Echinodermata, Mollusca, Porifera and Sipuncula) on macroplastics. With 15 species, bryozoans are the most diverse group on plastics. We also report 17 egg cases of the catshark Scyliorhinus sp., and highlight the implications for their dispersal. Our results suggest that plastic polymers may be relevant for distinct fouling communities, likely due to their chemical structure and/or surface properties. Our study provides evidence that biofouling may play a role in the sinking of plastic debris, as the most abundant fouled plastics had lower densities than seawater, and all bryozoan species were characteristic of shallower depths than those sampled. More studies at low taxonomic level are needed in order to detect new species introduction and potential invasive species associated with plastic debris.
... Type specimens, comprising the newly discovered mature colony (holotype) and the two-largest earlier-described colonies (designated paratypes; Grischenko et al. 2018, p. 55, figs 36, 37), are deposited in the Zoological Institute, Russian Academy of Science (ZIRAS), Saint Petersburg, Russia. The classification below is adapted from that provided by Bock & Gordon (2013). Etymology. ...
Article
A new abyssal cyclostome bryozoan genus and species, Vasopora ceramica n. gen., n. sp., is described from the eastern Russian exploration area of the Clarion-Clipperton Fracture Zone based on newly collected material from Yuzhmorgeologiya GLD4-19 station 421 (13.23408° N, 134.22180° W, 4809 m depth). Generic characters include an erect pedunculate colony with a distinct boundary between column and flared capitulum, short autozooidal peristomes in a single whorl, numerous alveoli, a central unidirectional sac-like gonozooid covered by a surficial network of crossed ridges continuous with adjacent rims of alveoli, a laterally opening ooeciopore, and the entire capitulum surface being minutely densely granular to subspinulate. Whereas the skeletal microstructure of the capitulum surface comprises irregular imbricated crystallites, the column has a planar-spherulitic fabric of acicular crystallites in fan-like arrays, and there are no pseudopores. The sharp boundary between capitulum and column, with their different microstructure separates Vasopora n. gen. from the two existing genera of Alyonushkidae that are found in the same environment. Vasopora n. gen. has a stalk formed of calcified exterior wall, whereas it is interior-walled in Alyonushka and Calyssopora.
... Bryozoans are a group of colonial, aquatic suspensionfeeders consisting of over 6.000 recent and over 15.000 fossil described species [1]. Along with other phyla such as mollusks, nemerteans or phoronids, they belong to the protostome Lophotrochozoa, and lately are reunited with phoronids and brachiopods as monophyletic lophophorates (e.g. ...
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Background Bryozoans are sessile aquatic suspension feeders in mainly marine, but also freshwater habitats. Most species belong to the marine and calcified Cheilostomata. Since this taxon remains mostly unstudied regarding its neuroanatomy, the focus of this study is on the characterization and ground pattern reconstruction of the autozooidal nervous system based on six representatives. Results A common neuronal innervation pattern is present in the investigated species: a cerebral ganglion is located at the base of the lophophore, from where neurite bundles embrace the mouth opening to form a circumoral nerve ring. Four neurite bundles project from the cerebral ganglion to innervate peripheral areas, such as the body wall and parietal muscles via the tentacle sheath. Five neurite bundles comprise the main innervation of the visceral tract. Four neurite bundles innervate each tentacle via the circumoral nerve ring. Mediofrontal tentacle neurite bundles emerge directly from the nerve ring. Two laterofrontal- and one abfrontal tentacle neurite bundles emanate from radial neurite bundles, which originate from the cerebral ganglion and circumoral nerve ring in between two adjacent tentacles. The radial neurite bundles terminate in intertentacular pits and give rise to one abfrontal neurite bundle at the oral side and two abfrontal neurite bundles at the anal side. Similar patterns are described in ctenostome bryozoans. Conclusions The present results thus represent the gymnolaemate situation. Innervation of the tentacle sheath and visceral tract by fewer neurite bundles and tentacular innervation by four to six tentacle neurite bundles support cyclostomes as sister taxon to gymnolaemates. Phylactolaemates feature fewer distinct neurite bundles in visceral- and tentacle sheath innervation, which always split in nervous plexus, and their tentacles have six neurite bundles. Thus, this study supports phylactolaemates as sistergroup to myolaemates.
... Vieira et al. 2008), classification (e.g. Bock & Gordon 2013), bryozoans as associates of other organisms (e.g. Rudman 2007;, metazoan phylogeny (e.g. ...
Article
This short account is an invited contribution to the Zootaxa special volume ‘Twenty years of Zootaxa.’ Zootaxa was first published on 28 May 2001. Between this date and December 2020, 116 papers were published in Zootaxa that mention Bryozoa, comprising mostly descriptions of new species and higher taxa, but also including molecular sequencing (e.g. Fehlauer-Ale et al. 2011; Taylor et al. 2011; Franjevic et al. 2015), invasive-species research (e.g. Ryland et al. 2014; Vieira et al. 2014), checklists (e.g. Vieira et al. 2008), classification (e.g. Bock & Gordon 2013), bryozoans as associates of other organisms (e.g. Rudman 2007; Chatterjee & Dovgal 2020; Chatterjee et al. 2020), metazoan phylogeny (e.g. Giribet et al. 2013), biographies of historical figures who worked on bryozoans (e.g. Calder & Brinkmann-Voss 2011; Calder 2015) and a catalogue of the fossil invertebrate taxa described by William Gabb (including 67 bryozoan species) (Groves & Squires 2018). Of the 116 papers, 15 (13%) were open-access.
... Bryozoa is a phylum of suspension-feeding, almost exclusively colonial lophophorates comprising over 6000 recent species (Bock & Gordon, 2013). Individuals of a colony are termed zooids and comprise a protective body wall and cuticle (cystid) and a tentacle crown and associated digestive tract (polypide), which is characteristically retractable by retractor muscles (Ryland, 1970). ...
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Article
The genus Pierrella was originally created for a single fossil ctenostome bryozoan species from the Late Cretaceous, which is characterized by runner‐like colonies, with zooids possessing a distinctive radial, folded aperture. Not long ago, a few specimens of a recent deep‐sea congener, P. plicata, were discovered and described from the Russian exploration area of the Clarion‐Clipperton Fracture Zone, eastern Central Pacific Ocean. Owing to the lack of data on the internal morphology of this species, we investigated the soft‐body morphology of P. plicata using serial sectioning and 3D‐reconstruction in order to compare it to other more recently investigated ctenostome bryozoans and to infer the systematic position of the genus. The most striking peculiarity of the examined species is the radial aperture formed by multiple cuticular, pleated folds of the cystid wall. The cuticle is thickened into triangular‐shaped folds in this area. An orifical sphincter underlies the folded aperture. Apertural muscles are present as a single pair of parieto‐diaphragmatic muscles and four duplicature bands. The remaining polypide anatomy is mainly characterized by its miniature design: the lophophore has eight short tentacles and the digestive tract is one of the shortest and most compact ever observed in any bryozoan. A small intertentacular organ was detected at the lophophoral base. Taken together the genus Pierrella shows unique characters, such as the radial apertural folds that are closed by a series of orificial sphincter muscles, and its particularly small polypide. The general colony morphology resembles arachnidioidean ctenostomes whereas its internal morphology resembles alcyonidioidean species.
... Bryozoans comprise marine Stenolaemata (with single order Cyclostomata), predominantly marine Gymnolaemata (orders Ctenostomata and Cheilostomata), and exclusively freshwater Phylactolaemata. This phylum, reportedly containing more than 6000 extant species (Ryland, 2005;Bock and Gordon, 2013), is among the least studied animal groups where complete mitochondrial genomes until recently were sequenced in only eight species, seven of which are marine (Jang and Hwang, 2009;Nesnidal et al., 2011;Shen et al., 2012;Sun et al., 2009, Waeschenbach et al., 2006, 2012a. Recently, new mitogenomes of other 33 bryozoan species have been added, but all of them belong to Gymnolaemata (Orr et al., 2019a(Orr et al., , 2021. ...
Article
Bryozoans are aquatic colonial suspension-feeders abundant in many marine and freshwater benthic communities. This phylum is under-investigated, however, on both morphological and molecular levels, and its position on the metazoan tree of life is still disputed. Bryozoa include the exclusively marine Stenolaemata, predominantly marine Gymnolaemata and exclusively freshwater Phylactolaemata. Here we report the mitochondrial genome of the phylactolaemate bryozoan Cristatella mucedo. This species has the largest (21,008 bp) of all currently known bryozoan mitogenomes, containing a typical metazoan gene compendium (except for not found trnY) as well as a number of non-coding regions, three of which are longer than 1500 bp. The trnS1/trnG/nad3 region is presumably duplicated in this species. Comparative analysis of the gene order in C. mucedo and another phylactolaemate bryozoan, Pectinatella magnifica, confirmed their close relationships, and revealed a stronger similarity to mitogenomes of phoronids and other lophotrochozoan species than to marine bryozoans, indicating the ancestral nature of their gene arrangement. We suggest that the ancestral gene order underwent substantial changes in different bryozoan clades showing mosaic distribution of conservative gene blocks regardless of their phylogenetic position. Altogether, our results support the early divergence of Phylactolaemata from the rest of Bryozoa.
... Other genera: Ca 10 genera: Filisparsa d 'Orbigny, 1853, Microeciella Taylor & Sequeiros, 1982, Proboscina Audouin, 1826 Pallas, 1766). Other genera: More than 20 genera of which half are extant (Bock & Gordon 2013), among them such genera as Exidmonea David et al., 1972, Fenestulipora Taylor & Gordon, 1997, Harmelinopora Brood, 1976, and Idmidronea Canu & Bassler, 1920 Diagnosis: Encrusting or erect colonies of various morphology, fixed-walled, formed by long zooecia commonly grouped into longitudinal or oblique connate rows. Peristomes usually long. ...
... The phylum Bryozoa is moderately diverse, estimated in 2013 to contain 5869 living species (Bock and Gordon 2013), and has a good fossil record dating back to the Early Ordovician (Taylor and Ernst 2004). Most extant bryozoan species are marine and belong to the order Cyclostomata in the class Stenolaemata or the dominant order Cheilostomata in the class Gymnolaemata. ...
Chapter
Bryozoan constructions have been present in all major climatic zones of marine ecosystems for the past 450 million years, since the Early Ordovician. Some fossil species possessed large bioconstructional colonies that would have provided habitats for other marine animals and plants, just as similar colonies do at the present day. Today, biogenic calcareous structures vary greatly in size, ranging from a few centimetres to many kilometres, and in complexity, forming composite structures that are distributed globally. Despite the role of bioconstructional bryozoan species in promoting marine biodiversity worldwide, they have been excluded from several protection strategies. Information emerging from the literature provides only a very incomplete picture of the role of bryozoans as bioconstructors at the global level. The ability of bryozoans to form long-lasting carbonate structures makes them important carbonate producers, with a significant role in the carbon cycle. Bryozoan reefs are facing the challenges of climate change, which will be detrimental for some species and their associated biota. Better knowledge of the contribution of bryozoans to the carbon stock and the ecosystem services they provide will be of great importance to ensure their protection and to understand their potential in adaptation strategies under future ocean scenarios.
... A systematic study of Bryozoan is primarily based on the highly calcified zooecia [1]. Circumscribed bryozoans include three classes, four orders, 187 families, 808 genera and about 6000 species [4]. A total of 257 marine species have been identified from Indian EEZ so far. ...
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Article
Aiming to collate the distribution and the attachment preference of intertidal bryozoan of Andaman waters, a study was carried out in three intertidal sites (Burmanallah, Kodiyaghat, and Chidiytapu) of southeastern coasts of the Andaman Islands between June and August 2016. The present study is the first exclusive report on bryozoans from the Andaman Islands after a long research gap of nine decades. During our investigation , a total of twelve genera were identified from both calcareous and non-calcareous substratum. Out of the twelve genera, eight genera were new records from the island. The present study showed that the attachment affinity of the Bryozoans is more towards natural substratum particularly on the rocks. The Thalamoporella sp. reported the most abundant species with maximum average colony length of 3.5 cm from the rock substratum.
... The holotype specimen is deposited in the Zoological Institute Russian Academy of Science (ZIRAS), Saint Petersburg, Russia. The classification we use follows Bock & Gordon (2013). Pandanipora fragilis n. sp. ...
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Article
A new cyclostome bryozoan, Pandanipora fragilis n. sp., is described from 3453 m depth on the subequatorial Mid-Atlantic Ridge. It differs from the type species, P. helix Grischenko, Gordon & Melnik, 2018, by a combination of colonial and zooidal characters. These include regular branching of a uniserial stem along the entire colony length; a straight or just slightly elevated and gently curved distal autozooidal peristomial component, forming a sharp angle with the frontal wall of the next zooid; the pattern of zooidal budding, achieved via development of a partition from the floor of the parent zooid in its distal quarter to third, leaving the proximal portion of daughter zooids not overlapping, so that preceding and subsequent zooids are not appressed along their proximal segments; props are uniformly straight and filiform, with their diameter much smaller than in P. helix; the exceptionally rare presence of slit-like pseudopores, restricted to props; and wedge-shaped crystallites on the internal surface of developing zooids, with irregular, ragged edges. The discovery of P. fragilis suggests that Pandanipora is most likely a relict element of a more-widespread ancient distribution that existed in tropical and boreal zones of both the Atlantic and Pacific Oceans prior to formation of the Isthmus of Panama in the Pliocene.
... The phylum is monophyletic and can be divided into three monophyletic classes: Gymnolaemata, Stenolaemata and Phylactolaemata (Fuchs et al. 2009;Taylor and Waeschenbach 2015). Members of Phylactolaemata occur exclusively in fresh water, while members of Stenolaemata (order Cyclostomatida) and Gymnolaemata (orders Ctenostomatida and Cheilostomatida) inhabit marine and/or estuarine environments (Hayward and Ryland 1999;Bock and Gordon 2013). Most of our knowledge about bryozoans around South Africa is taxonomic in nature, though a few works have been published on biology and ecology (e.g. ...
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Article
The zoogeography of marine Bryozoa around South Africa was investigated using published distribution records, museum catalogues, and an examination of previously unworked bryozoan material in (mostly) museum collections. Although a total of 276 valid species are recognised, it was not possible to unambiguously assess geographic patterns of diversity. At all depth zones examined (shore and inner-shelf, 0–30 m; mid- and outer-shelf, 31–350 m; bathyal, >500 m), there was a clear geographic structure to communities that mirrored established regional patterns of biogeography. Too few samples were collected from the shelf edge (351–500 m) and they were consequently excluded from zoogeographic analysis. Communities on the shore and inner-shelf and on the mid- and outer-shelf were more similar to each other than they were to bathyal communities, and the pronounced structure in bathyal communities suggests heterogeneity in the deep sea around South Africa.
... By contrast, Stenolaemata have not received nearly as much attention from researchers (Smith et al. 2017). It is a relatively large class with a rich fossil history (thousands of species from six extinct orders, dating back to the Ordovician) and about 543 living species in 98 genera in the extant order Cyclostomatida (Ryland 1970;Bock and Gordon 2013). Yet accounts of morphology of their soft parts and interrelationships with skeletal dimensions are extremely uncommon. ...
Article
Marine bryozoans have been members of benthic skeletal faunas since the Ordovician. These small suspension feeders collect particles in the range of 10 to 100 μm. Specific details of their feeding depend on the morphology of the feeding apparatus, which may be reflected in skeletal characters. While several studies have described the link between the skeletal and soft-body traits of gymnolaemate bryozoans, stenolaemates have received less attention. To fill this gap, we conducted a detailed analysis of morphometry within and across species and attempted to develop robust predictive models that can be used to infer the soft-body morphology from skeletal data. This, in turn, will help with extracting data on ecology of Paleozoic communities of suspension feeders from the extensive bryozoan fossil record. Characters of polypide morphology among New Zealand cyclostomates (single Recent order in Stenolaemata) displayed staggering variability and almost without exception were not connected to skeletal characters at the species level. When this variability is reduced to its central tendency, interspecific trends are more apparent. The relationship is positive, linear, and moderately strong, but the resulting models have wide predictive intervals (plus/minus hundreds of micrometers). A precise estimate of the characters of the feeding apparatus of modern, and especially fossil, stenolaemates may be difficult to attain, at least on the basis of the skeletal traits used here.
... In this study, we present a detailed characterization of the internal structure and biomineral architecture of colonies of recent Anoteropora latirostris, a bryozoan that is free-living and does not encrust with their basal colonial surface. The genus Anoteropora ( Canu and Bassler, 1927) belongs to the suborder Cheilostomata which is the most diverse one of recent bryozoans, represented by about 5000 extant species (Bock and Gordon, 2013), and showing the highest developed polymorphism among all Bryozoa. ...
Thesis
Bivalve shells are nano-composite materials consisting of crystalline calcium carbonate phases (i.e. aragonite and calcite) that are intimately arranged with organic constituents into different hierarchical architectures. This explains their enhanced mechanical properties that outperform their monolithic geological counterparts. Shells grow incrementally throughout their lifetime and record environmental conditions in their trace element and isotopic signatures. Our understanding of how shells grow has seen a recent paradigm change away from classical ion-by-ion crystallization models to non-classical crystallization pathways via colloid attachment and transformation of nanogranular amorphous calcium carbonate precursor phases. These findings complicate any paleo-environmental reconstructions and lead to fundamental questions, such as: which impact do the new crystallization pathways have on trace element partitioning? How are these pathways affected by physiological effects on sub-micron growth processes? These questions are addressed in this thesis by using living shells in controlled aquaculture conditions. Four different bivalve species with different architectural ultrastructures were studied here: Anadara trapezia (crossed-lamellar and complex crossed-lamellar architectures), Katelysia rhytiphora (compound composite prismatic and crossed-acicular), Mytilus galloprovincialis (simple prismatic and nacre) and Pinctada imbricata fucata (simple prismatic and nacre). These bivalves were periodically subjected to pulse-chase labelling experiments with strontium-enriched seawater to create “snapshots” of submicron shell growth processes correlatively visualized with micro- to nano-analytical instrumentation such as EPMA, FEG-SEM, Nano-SIMS, Micro-Raman, EBSD, HR-(S)TEM, and Atom Probe Tomography. As non-nacreous architectures are virtually not studied, pulse-chase labelling experiments combined with architectural investigations provide a powerful tool for detailed characterization at the micron scale: the Sr-label in the shell is shown to cut across all architectural units independent of hierarchy and local orientation, indicating growth rates across a uniform growth front during shell mineralization. While this growth process may appear intuitive, this thesis presents here the first direct evidence compared to previous inferences from natural environments without rigorous time-resolution. This growth process across all architectural hierarchies in the shell is fundamentally different to that of nacroprismatic shells, which are thought to grow in a two-step process of forming organic compartments prior to calcification. Material properties of non-nacreous shells are investigated via stereographic visualization of Young’s moduli and reveal a girdle-like arrangement of elastically stiffer orientations that result in quasi-isotropic planes perpendicular to the growth direction across the entire shell. For nacroprismatic shells at the nano-scale, the thesis presents new insights into the toughening processes investigated via innovative TEM-based nanoindentation experiments. At low compression, it is found that organic interfaces confine strain propagation within each tablet and nanogranules as well as organic inclusions rotate and deform, while at high compression tablets interlock and fully recover to their original state. Nacre is by far the best-studied biomineral, but its insulating properties and high organic content presents significant challenges when taking geochemical investigations to the nano-scale. Therefore, a best practice methodological protocol for Atom Probe Tomography is developed for nacre that minimises the generation of artefacts in the mass spectrum and allows for correlative analysis of the insulating material. This protocol paves the way for innovative studies focussing on the growth dynamics within individual nacre tablets. The correlated, multianalytical approach is further used to study the ultrastructure and composition of bimineralic cheilostome bryozoan skeletons that consist of primary, calcitic lateral walls that are covered with fine-grained fibrous aragonite on their distal side while the colony’s frontal and basal walls are fully aragonitic. This arrangement of mineral phases was previously unknown in cheilostome bryozoa. Further, an organic membrane is situated between both mineral phases that may serve as a template for biomineralisation. The last part of the thesis investigates another facet of nacre – lustrous pearls by mapping out the knowledge gaps existing in the area of provenance identification and presenting the world’s only source of untreated Akoya cultured pearls. These untreated pearls are an invaluable source to investigate the natural colour-palette that are linked to pigments within the nacre.
... They create habitats that support diverse communities [15], and contribute significantly to shelf carbonate dynamics [16]. This family is currently divided into 10 extant genera with 106 described extant species [17]. Previously, the Adeonidae were separated into two distinct families by Gregory (1893): Adeonidae (including Adeona, Adeonellopsis, Bracebridgia, Cucullipora and Reptadeonella) and Adeonellidae, Gregory 1893 (comprising the genera Adeonella and Laminopora). ...
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Background: Understanding the phylogenetic relationships among species is one of the main goals of systematic biology. Simultaneously, credible phylogenetic hypotheses are often the first requirement for unveiling the evolutionary history of traits and for modelling macroevolutionary processes. However, many non-model taxa have not yet been sequenced to an extent such that statistically well-supported molecular phylogenies can be constructed for these purposes. Here, we use a genome-skimming approach to extract sequence information for 15 mitochondrial and 2 ribosomal operon genes from the cheilostome bryozoan family, Adeonidae, Busk, 1884, whose current systematics is based purely on morphological traits. The members of the Adeonidae are, like all cheilostome bryozoans, benthic, colonial, marine organisms. Adeonids are also geographically widely-distributed, often locally common, and are sometimes important habitat-builders. Results: We successfully genome-skimmed 35 adeonid colonies representing 6 genera (Adeona, Adeonellopsis, Bracebridgia, Adeonella, Laminopora and Cucullipora). We also contributed 16 new, circularised mitochondrial genomes to the eight previously published for cheilostome bryozoans. Using the aforementioned mitochondrial and ribosomal genes, we inferred the relationships among these 35 samples. Contrary to some previous suggestions, the Adeonidae is a robustly supported monophyletic clade. However, the genera Adeonella and Laminopora are in need of revision: Adeonella is polyphyletic and Laminopora paraphyletically forms a clade with some Adeonella species. Additionally, we assign a sequence clustering identity using cox1 barcoding region of 99% at the species and 83% at the genus level. Conclusions: We provide sequence data, obtained via genome-skimming, that greatly increases the resolution of the phylogenetic relationships within the adeonids. We present a highly-supported topology based on 17 genes and substantially increase availability of circularised cheilostome mitochondrial genomes, and highlight how we can extend our pipeline to other bryozoans.
... One of the biggest gaps in genome-based phylogeny is Bryozoa -medium-sized phylum of microscopic aquatic invertebrates comprising about 6000 recent and more than 15000 fossil species and having a long fossil history beginning in the early Ordovician 1 . The phylum consists of three classes: exclusively freshwater, non-skeletal Phylactolaemata, exclusively marine, calcified Stenolaemata and predominantly marine, but occasionally brackish-and freshwater, Gymnolaemata. ...
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The Southern Ocean is showing one of the most rapid responses to human-induced global change, thus acting as a sentinel of the effects on marine species and ecosystems. Ocean warming and acidification are already impacting benthic species with carbonate skeletons, but the magnitude of these changes to species and ecosystems remains largely unknown. Here we provide the largest carbonate mineralogical dataset to date for Southern Ocean bryozoans, which are diverse, abundant and important as carbonate producers, thus making them excellent for monitoring the effects of ocean warming and acidification. To improve our understanding of how bryozoans might respond to ocean warming and acidification, we assess latitudinal and seafloor temperature patterns of skeletal mineralogy using bryozoan species occurrences together with temperature data for the first time. Our findings, combining new mineralogical data with published data from warmer regions, show that the proportions of high-Mg calcite and bimineralic species increase significantly towards lower latitudes and with increasing seawater temperature. These patterns are consistent with the hypothesis that seawater temperature is likely a significant driver of variations in bryozoan mineralogy at a global scale.
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We use natural language processing (NLP) to retrieve location data for cheilostome bryozoan species (text-mined occurrences (TMO)) in an automated procedure. We compare these results with data combined from two major public databases (DB): the Ocean Biodiversity Information System (OBIS), and the Global Biodiversity Information Facility (GBIF). Using DB and TMO data separately and in combination, we present latitudinal species richness curves using standard estimators (Chao2 and the Jackknife) and range-through approaches. Our combined DB and TMO species richness curves quantitatively document a bimodal global latitudinal diversity gradient for extant cheilostomes for the first time, with peaks in the temperate zones. A total of 79% of the georeferenced species we retrieved from TMO ( N = 1,408) and DB ( N = 4,549) are non-overlapping. Despite clear indications that global location data compiled for cheilostomes should be improved with concerted effort, our study supports the view that many marine latitudinal species richness patterns deviate from the canonical latitudinal diversity gradient (LDG). Moreover, combining online biodiversity databases with automated information retrieval from the published literature is a promising avenue for expanding taxon-location datasets.
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Animal mitogenomes are typically devoid of introns. Here, we report the largest number of mitochondrial introns ever recorded from bilaterian animals. Mitochondrial introns were identified for the first time from the phylum Bryozoa. They were found in four species from three families (Order Cheilostomatida). A total of eight introns were found in the complete mitogenome of Exechonella vieirai, and five, 17 and 18 introns were found in the partial mitogenomes of Parantropora penelope, Discoporella cookae and Cupuladria biporosa, respectively. Intron-encoded protein domains reverse transcriptase and intron maturase (RVT-IM) were identified in all species. Introns in E. vieirai and P. penelope had conserved Group II intron ribozyme domains V and VI. Conserved domains were lacking from introns in D. cookae and C. biporosa, preventing their further categorization. Putative origins of metazoan introns were explored in a phylogenetic context, using an up-to-date alignment of mitochondrial RVT-IM domains. Results confirmed previous findings of multiple origins of annelid, placozoan and sponge RVT-IM domains and provided evidence for common intron donor sources across metazoan phyla. Our results corroborate growing evidence that some metazoans with regenerative abilities (i.e. placozoans, sponges, annelids and bryozoans) are susceptible to intron integration, most likely via horizontal gene transfer.
Article
Twenty-four Recent species of the boreal-Arctic and Pacific cheilostome bryozoan genus Rhamphostomella are described. The species R. tatarica and R. pacifica are transferred to Rhamphostomella from Posterula and Porella, respectively. Eight species are new: R. aleutica n. sp., R. aspera n. sp., R. commandorica n. sp., R. echinata n. sp., R. microavicularia n. sp., R. morozovi n. sp., R. multirostrata n. sp. and R. obliqua n. sp. Neotypes are selected for six species, and lectotypes for eight species. Mixtoscutella n. gen. is established for several Rhamphostomella-like species, including M. androsovae [formerly Smittina androsovae Gontar], M. cancellata [formerly Escharella porifera forma cancellata Smitt], M. harmsworthi [formerly Schizoporella harmsworthi Waters], M. ovata [formerly Cellepora ovata (Smitt)], and M. ussowi [formerly Schizoporella ussowi (Kluge)]. In addition to taxonomic revision, the morphology (frontal shields, ovicells and multiporous septula), ecology and zoogeography of these cheilostomes are discussed, and identification keys are presented. Most species of Rhamphostomella have broad bathymetric distributions. Some have long protuberances on their basal walls that allow them to grow elevated above allelopathically active substrates such as sponges. The diversity of Rhamphostomella peaks in the northwestern Pacific.
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Among marine invertebrates, bryozoans are small, not well known, and complex to identify. Nevertheless, they offer unique opportunities for whole-body generation research, because of their colonial, modular mode of growth. Here, we describe detailed methods for collection of bryozoans from a range of environments, sample preparation and identification, culture and feeding, spawning and breeding, marking colonies for growth studies, and histological preparation.Key wordsBryozoansCultureCollection Growth Feeding Whole-body regeneration Budding Regression Brown bodies Anatomy Histology Larvae Spawning Settlement
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Based on records dating from 1859 to 2021, we provide an overview of the marine animal diversity reported for Galiano Island, British Columbia, Canada. More than 650 taxa are represented by 20,000 species occurrence records in this curated dataset, which includes dive records documented through the Pacific Marine Life Surveys, museum voucher specimens, ecological data and crowd-sourced observations from the BC Cetacean Sightings Network and iNaturalist. We describe Galiano Island's marine animal diversity in relation to the Salish Sea's overall biodiversity and quantify the proportional contributions of different types of sampling effort to our current local knowledge. Overviews are provided for each taxonomic group in a format intended to be accessible to amateur naturalists interested in furthering research into the region's marine biodiversity. In summary, we find that the Pacific Marine Life Surveys, a regional community science diving initiative, account for 60% of novel records reported for Galiano Island. Voucher specimens account for 19% and crowd-sourced biodiversity data 18% of novel records, respectively, with the remaining 3% of reports coming from other sources. These findings shed light on the complementarity of different types of sampling effort and demonstrate the potential for community science to contribute to the global biodiversity research community. We present a biodiversity informatics framework that is designed to enable these practices by supporting collaboration among researchers and communities in the collection, curation and dissemination of biodiversity data.
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This paper summarizes the trends in nematode species description and systematics emerging from a comparison of the latest comprehensive classification and census of Phylum Nematoda (Hodda 2022a, b) with earlier classifications (listed in Hodda 2007). It also offers some general observations on trends in nematode systematics emerging from the review of the voluminous literature used to produce the classification. The trends in nematodes can be compared with developments in the systematics of other organisms to shed light on many of the general issues confronting systematists now and into the future.
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There is a lack of knowledge regarding the modern status of intertidal bryozoan communities in the coastal Barents Sea. Here, we studied species composition, richness, and biomass of bryozoans in Yarnyshnaya and Dalnezelenetskaya Bays, both located in the eastern part of the Kola Peninsula (Barents Sea), in summer. Species composition and biodiversity were consistent with previous research but the record of the ctenostome bryozoan Walkeriauva is the first for the region indicating eastward range expansion of this species associated with climate forcing in the Arctic. Mean biomass was relatively low accounting for 2.25 ± 0.95 g·m−2. The most common species were Eucratea loricata, Harmeria scutulata, Crisia eburnea, and Cribrilina cryptooecium averaging 96% of the total biomass. Cluster analysis delineated two distinct groups of stations, one with true marine conditions and another with brackish water conditions. Redundancy analysis revealed that bryozoan diversity was strongly associated with salinity fluctuations being extremely low at brackish water sites. In contrast, water temperature was found to be a significant contributor to biomass with the lowest values found at warmer waters probably owing to the predominance of Boreo-Arctic species which prefer lower-temperatures. Other hydrological variables (dissolved organic matter, silicates, and oxygen) were consistent with usual summer values and had no significant effects on the bryozoan assemblages. Our study provides a reference point for further biodiversity studies in changing marine ecosystems of the Arctic region.
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We used micro-CT and SEM to trace zooidal budding in Hornera (Cyclostomatida: Cancellata) from the ancestrula onwards. Results show that hornerid branches are constructed by dual zooidal budding modes occurring synchronously at two separate budding sites at the growing tips. Frontal autozooids bud from a multizooidal budding lamina. Lateral autozooids bud from discrete abfrontal budding loci by 'exomural budding', a previously undescribed form of budding centred on hypostegal pores in interzooidal grooves on the colonial body wall. These two budding modes are integrated during primary branch morphogenesis, forming composite, developmentally bilaminate, branches.
Article
Cheilostomata is the most diverse and ecologically dominant order of bryozoans living today. We apply a Bayesian framework to estimate macroevolutionary rates of cheilostomes since the Late Jurassic across four datasets: (I) manually curated genus ranges; (II) published text-mined genus ranges; (III) non-revised Paleobiology Database (PBDB) records; (IV) revised and augmented PBDB records. All datasets revealed increased origination rates in the Albian, and a twin K–Pg and Danian extinction rate peak. High origination rates in the Late Selandian–Ypresian in Dataset I indicate the onset of an ascophoran-grade radiation. Lineage-through-time plots confirm the macroevolutionary lag preceding the radiation of cheilostomes in the middle Cretaceous, and their renewed diversification in the late Paleocene and Eocene. A multivariate birth–death model indicates that origination rates are shaped by diversity-dependent dynamics coupled with a positive correlation with sea surface temperature, while extinction rates negatively correlate with sea level. Text-mined data provide broadly similar rate dynamics as manually curated data, although discrepancies could be attributed to the omission of key literature in Dataset II, and the inclusion of new published and unpublished data, and revised ranges in Dataset I. Revision and augmentation of PBDB occurrences were necessary to generate rate profiles akin to those of Datasets I and II and highlight the risks of using unedited occurrence data. Our results support the widely held assumption that diversification dynamics are controlled by both biotic and abiotic factors, and pave the way for integrating fossils with molecular phylogenies to study these processes in more detail.
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Bryozoans are epibenthic suspension-feeders and use their ciliated tentacles to generate feeding currents. Modern bryozoan mouth size limits the size of the particles that can be ingested, and lophophore diameter is correlated with water pumping rates. In fossil bryozoans these soft parts are absent, however mouth and lophophore dimensions can be inferred from preserved skeletons. Gondwanan Permian palaeostomate bryozoans show distinct order-level trophic partitioning across warm to cold-water faunas. In diverse warm-water faunas of southern Thailand, fenestrate bry-ozoans consumed the smallest food particles, cryptostomes and trepostomes consumed mid-size particles, and cystopo-rates consumed the largest, and widest range, of particles. In contrast, in low diversity cold-water faunas of eastern Aus-tralia, where cystoporates and cryptostomes are uncommon, fenestrate bryozoans again consumed the smallest food particles , however the abundant trepostomes had much larger mouths than their warm-water counterparts and consumed the largest food particles. This variation in mouth size, especially in the trepostomes, suggests that within a regional fauna mouth size is not controlled only by higher level sys-tematics and that sessile benthic organisms are trophically structured to utilize different sized food particles from a suspended food source. Gondwanan Permian palaeostomates had much larger mouth and lophophore sizes than modern stenolaemates (cyclostomes) and are more comparable to gymnolaemate (cheilostome) bryozoans. This suggests that palaeostomates were able to access and consume the same range of food resources as modern gymnolaemate bryozoans, and feed at similar rates, prior to the marine environmental changes at the close of the Palaeozoic.
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Pherusellid ctenostomes are a little‐investigated clade of bryozoans comprising three described species that are usually distinguished by their different peristome sizes. In this study, the entire family of Pherusellidae is revised with description of two new species, Pherusella minima from the Mediterranean Sea and Pherusella liowae from Singapore. Serial semithin sections and 3‐D reconstruction are used for the first time on an entire family of bryozoans. Soft‐tissue morphology yields numerous characters such as details of the digestive tract, duplicature bands, and pore‐plate structure that are useful and necessary for species identification. Cystid characters alone are regarded as insufficient for proper species recognition. Numerous previous studies have reported Pherusella tubulosa from Asia, but probably have misidentified this species. Hence, it appears to be restricted to the Atlantic Ocean and Mediterranean Sea. Furthermore, we suggest Pherusella brevituba only occurs in the Pacific Ocean and is not an alien species to the Mediterranean Sea, as previously suggested. We conclude that previous records most likely belong to the new species P. minima, which itself is endemic to the Mediterranean Sea. This study revises the ctenostome bryozoan family Pherusellidae by analyzing soft‐body morphological characters. Two new species Pherusella minima sp. nov. from the Mediterranean Sea and Pherusella liowae sp. nov. from Singapore are described.
Article
In bryozoans (phylum Bryozoa), representative colonial animals mostly found in marine environments, some species possess different types of individuals (heterozooids) specialized in different functions such as defense or structural support for their colonies. Among them, the best-known heterozooids are the avicularia, known to function as defenders. The differentiation processes of heterozooids, including avicularia, should be important keys to understand the evolutionary significance of bryozoans. However, the developmental process of avicularium formation remains to be fully understood. In this study, therefore, in order to understand the detailed developmental process and timing of avicularium formation, extensive observations were carried out in a bryozoan species, Bugulina californica (Cheilostomata, Bugulidae), that possesses adventitious avicularia, by performing stereomicroscopy on live materials, in addition to scanning electron microscopy and histological observations. The whole process can be divided into seven stages based on developmental events. Especially notably, at the earlier stages, there are three major budding events that produce proliferating cell masses corresponding to primordial tissues: (1) budding of the peduncle cushion at the outer margin of the distal part of a young autozooid, (2) budding of the head-part primordium from the peduncle cushion, and (3) budding of the polypide inside the head part. Experimental control of temperature showed that 20°C would be the best to maintain B. californica colonies.
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Bryozoans are major fouling organisms and include some of the most invasive marine species globally. Hull fouling of transoceanic vessels is a major vector of non-indigenous bryozoans. One genus known to be important in this regard is Conopeum, but its occurrence in the coastal waters of South Korea has yet to be established. We sorted bryozoan samples from the collection of Park et al. (2017) and carried out surveys for marine organisms in 2013 and 2019–2020 in coastal waters of South Korea. We found two non-indigenous bryozoans: Conopeum reticulum (Linnaeus, 1767) and C. seurati (Canu, 1928). These two species and a third in the genus, C. hexagonum Seo, 1996, have distinctive morphologies and distributions along environmental gradients of the sampling sites. Gymnocyst and cryptocyst development and spine presence were used to identify each species morphologically. According to the salinity and turbidity of the sites, the inhabiting species appeared differently. The two non- indigenous species occurred in association with other sessile organisms such as oysters, mussels, and serpulid polychaetes. In particular, C. reticulum was associated with an invasive species of the Mediterranean mussel (Mytilus galloprovincialis Lamarck, 1819) and C. seurati was associated with the serpulid polychaetes Hydroides ezoensis Okuda, 1934 and Ficopomatus enigmaticus (Fauvel, 1923), as well as with the mytilid bivalve Xenostrobus securis (Lamarck, 1819), and the Pacific oyster Crassostrea gigas (Thunberg, 1793). Conopeum seurati and H. ezoensis completely encrusted all surfaces in artificial canal docks seemingly negatively affecting richness of native species.
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Larger molecular phylogenies based on ever more genes are becoming commonplace with the advent of cheaper and more streamlined sequencing and bioinformatics pipelines. However, many groups of inconspicuous but no less evolutionarily or ecologically important marine invertebrates are still neglected in the quest for understanding species-and higher-level phylogenetic relationships. Here, we alleviate this issue by presenting the molecular sequences of 165 cheilostome bryozoan species from New Zealand waters. New Zealand is our geographic region of choice as its cheilostome fauna is taxonomically, functionally and ecologically diverse, and better characterized than many other such faunas in the world. Using this most taxonomically broadly-sampled and statistically-supported cheilostome phylogeny comprising 214 species, when including previously published sequences, we tested several existing systematic hypotheses based solely on morphological observations. We find that lower taxonomic level hypotheses (species and genera) are robust while our inferred trees did not reflect current higher-level systematics (family and above), illustrating a general need for the rethinking of current hypotheses. To illustrate the utility of our new phylogeny, we reconstruct the evolutionary history of frontal shields (i.e., a calcified body-wall layer in ascus-bearing cheilostomes) and asked if its presence has any bearing on the diversification rates of cheilostomes.
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Non-studied museum collections are hidden treasures—a source of information for various research fields. The novel taxa presented here were discovered during taxonomic examination of the backlogs of Bryozoa (Cheilostomata) from the Iziko South African Museum. We describe one new genus, Khulisa n. gen., and nine new species of bryozoans from South Africa. The new species are: Biflustra adenticulata n. sp., Aspidostoma sarcophagus n. sp., ?Micropora erecta n. sp., Trypostega richardi n. sp., Khulisa carolinae n. gen. et n. sp., Adeonella assegai n. sp., Hippomonavella lingulata n. sp., Phidolopora chakra n. sp. and Reteporella ilala n. sp. Three genera, Biflustra, Phidolopora and Triphyllozoon, are recorded for the first time from South Africa. This study highlights the importance of examining existing backlogged material lodged in museum collections.
Chapter
Bryozoans are common animals in marine benthic (seabed) ecosystems, and also occur in freshwater rivers, with 6063 and 108 extant species in each environment respectively. There are 17,867 fossil species described. They are distributed across the oceans with a relatively high diversity on the Antarctic shelf. They are colonial suspension filter feeders with a calcified skeleton, a wide variety of life cycles and growth forms, and are well represented in the fossil record.
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We identified nine species of phylactolaemate bryozoans collected from ponds and dams on Okinawa Island, Japan. This is the first report on freshwater bryozoans from Okinawa and includes the first record of Plumatella javanica from Japan. Phylactolaemate species were identified on the basis of colony and statoblast morphology and are illustrated by light mi-croscopy and scanning electron microscopy (SEM). We detected two new plumatellid species that, like Plumatella minuta and Plumatella vorstmani, have a transparent, weakly chitinized colony and hypertubercles on the floatoblast fenestra. A previous molecular study showed these four species to form a clade separate from Plumatella and Hyalinella. We here establish the new genus Rumarcanella to accommodate these species and describe Rumarcanella gusuku n. sp. and R. yan-baruensis n. sp. We discuss the dispersal of bryozoans to and from Japan by birds carrying statoblasts and provide a key to the Phylactolaemata of Japan.
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Article
The present study considers 88 bryozoan species occurring in freshwater: 69 phylactolaemate and 19 gymnolaemate species. Roughly 49% of these species are confined to one zoogeographical region. The cosmopolitan status of species like Fredericella sultana, Plumatella repens or P. emarginata has to be reconsidered. Among the Phylactolaemata, which are phylogenetically older than the Gymnolaemata, the gelatinous species (Lophopodidae, Pectinatellidae, Cristatellidae) are more primitive than the branching tubular species (Plumatellidae, Fredericellidae).
Article
Recent research on the ultrastructure of the calcareous skeleton in the bryozoan order Cyclostomata is summarized and updated, based on field emission SEM studies of 87 species. Six fundamental ultrastructural fabrics are recognized which differ in the crystallographic orientations, shapes and prevailing growth directions of the constituent crystallites. During the growth of individual walls a succession of fabrics is secreted, defining a fabric suite. Five fabric suites are described in interior walls and four in exterior walls. Nine ultrastructural characters were combined with 37 other skeletal characters in a PAUP analysis of the relationships between 28 post-Palaeozoic cyclostomes chosen to include representatives of all suborders. A single tree of length 142 steps was found. Comparison of tree statistics for three categories of characters showed ultrastructural characters to be more homoplastic than zooidal characters, and the latter more homoplastic than colonial characters. Rooting the tree on the paleotubuliporine Cuffeyella gave four transitions from fixed- to free-walled organization and no reversals. With respect to the five extant suborders of cyclostomes, this first, preliminary analysis implies that Rectangulata and Cancellata are monophyletic groups, whereas Articulata are diphyletic, and both Tubuliporina and Cerioporina paraphyletic. (C) 2000 The Linnean Society of London.
Article
Soft-bodied ctenostome bryozoans, preserved as bioimmurations following overgrowth by encrusting organisms with hard skeletons, are described from the Middle and Upper Jurassic of England and Normandy. They include one new genus, Cardoarachnidium, and three new species, C. bantai, C. voigti and Arachnoidella abusensis. These and new bioimmured specimens of Arachnidium smithii (Phillips) show fine-scale details of original morphology as well as artefacts caused by partial collapse of zooids during overgrowth. The D-shaped zooidal orifices present in Cardoarachnidium probably indicate that the zooids were operculate. Opercula are one of the two apomorphies of the closely-related cheilostomes, and therefore the Oxfordian Cardoarachnidium is placed in the stem-group of the Cheilostomata, which first appear in the Tithonian. -Author
Article
Recent research on the ultrastructure of the calcareous skeleton in the bryozoan order Cyclostomata is summarized and updated, based on field emission SEM studies of 87 species. Six fundamental ultrastructural fabrics are recognized which differ in the crystallographic orientations, shapes and prevailing growth directions of the constituent crystallites. During the growth of individual walls a succession of fabrics is secreted, defining a fabric suite. Five fabric suites are described in interior walls and four in exterior walls. Nine ultrastructural characters were combined with 37 other skeletal characters in a PAUP analysis of the relationships between 28 post-Palaeozoic cyclostomes chosen to include representatives of all suborders. A single tree of length 142 steps was found. Comparison of tree statistics for three categories of characters showed ultrastructural characters to be more homoplastic than zooidal characters, and the latter more homoplastic than colonial characters. Rooting the tree on the paleotubuliporine Cuffeyella gave four transitions from fixed- to free-walled organization and no reversals. With respect to the five extant suborders of cyclostomes, this first, preliminary analysis implies that Rectangulata and Cancellata are monophyletic groups, whereas Articulata are diphyletic, and both Tubuliporina and Cerioporina paraphyletic.
Article
The relative appearance of the parietal muscles in the development of the zooids has been studied in several ctenostomatous and “cheilostomatous” species. A comparison of the different on-togenetical sequences demonstrated that a “cheilostomatous” type of organization of the zooids with a great probability has been achieved in minimum three times independentl and originated from different ctenostomatous sub-grous: the Membranidea from plesiomorgic victorelloids (ancestors of Bulbella with not yet developed peristomial tube), the Inoviceiata (Aetea) from advanced forms of victorelloids with reduced primary parietal muscles (perhaps stcies related to Pottsiella), and Penetrantia from arachnidioid or vesicularioid ancestors (?). Therefore, the classical orders “Ctenostomata” and “Cheilostomata” represent only “stage groups” but no monohyletic systematical units. Because of the new concept and interpretation I propose a new name for the united group: Cteno-Cheilostomata, supra-ord. nov.
Article
A new term, “trophon”, has been introduced for the feeding and sexually propagating unit in the stolonate ctenostomes; the “stolon” and the “trophon” represent “sukzooids”, because they are homologous only to parts of the autozooids in the uniserially arranged ctenostomes. – The ontogeny of the muscles, especially the parietal muscles, has been studied in 17 species of ctenostomatous bryozoans belonging to different sub-groups of this order; the main effort was directed toward the details of the differentiations in living animals (14 of the investigated species). In various species of the Ctenostomata two separate sets of parietal muscles develop – probably as a consequence of the elongation of the cystid; the functional parietal muscles in the trophons of the stolonial forms are homologous to the secondary parietal muscles found in several uniserial ctenostomes. Contrary to the supposition by Soule (1954) in the different families of the serially arranged forms the muscles appear in different sequences during the development of the zooid and partly have different ontogenetical fates. Neither the “Carnosa” nor the “Paludicellea” (s. 1.) nor the “Stolonifera” can be maintained as taxa with the argument of a similar ontogeny of the muscles. A detailed comparison regarding the fate of the parietal muscles (and the polypide anlage) showed similarities between certain of the uniserial to either some of the stolonial, the sheet like, or the cystid-fusing forms suggesting a separate evolutionary origin of the advanced forms in those groups of the serially growing forms. – A new family, Pottsiellidae fam. nov., has been introduced and defined.
Article
The Cyclostomata are the only extant representatives of the class Stenolaemata, an ancient group of exclusively marine bryozoans. Previous cladistic analyses of cyclostome bryozoans, based exclusively on skeletal characters, revealed extensive homoplasy amongst morphological traits. This study presents the first molecular phylogeny for Cyclostomata and confirms the previous findings of homoplasy. Almost complete lsr and ssrDNA fragments were sequenced for 22 taxa of cyclostome bryozoans, plus the outgroup (Pectinatella magnifica and Flustrellidra hispida). Three well-supported major clades were found, but their inter relationships are unclear. Suborder Tubuliporina was polyphyletic, with representatives found in all three major clades. The tubuliporine family Plagioeciidae was resolved as polyphyletic; Plagioecia grouped with Lichenoporidae and Densiporidae, whereas Entalophoroecia, Diplosolen and Cardioecia formed a paraphyletic subgroup that included Frondiporidae and Horneridae. The suborder Cerioporina was also polyphyletic; Densiporidae grouped with Plagioecia and Lichenoporidae, whereas Heteroporidae nested in a paraphyletic subgroup of tubuliporines, with the Crisiidae forming the sister-group to this clade. Cinctiporidae could not be placed unambiguously. Morphological character mapping was performed in order to find evidence favouring one of the three possible hypotheses of inter relationships of the major clades, but the results were ambiguous. This study questions the extent to which morphological characters can be used in phylogenetic studies of cyclostome bryozoans, both fossil and extant, and how far their morphology is the result of ecophenotypic plasticity and convergent evolution. The finding of numerous non-monophyletic taxa has implications for extinction rate assessment and for the use of fossil cyclostomes to calibrate molecular trees.
classes) Class Phylactolaemata Allman, 1856 (1 order) Order Plumatellida Pennak, 1953 (6 families) Family Cristatellidae Allman, 1856 (1 genus, 1 species) Family Fredericellidae Hyatt, 1868 (2 genera, 8 species) Family Lophopodidae Rogick, 1935 (3 genera, 8 species) Family Pectinatellidae Lacourt
  • Ehrenberg Phylum Bryozoa
Phylum Bryozoa Ehrenberg, 1831 (3 classes) Class Phylactolaemata Allman, 1856 (1 order) Order Plumatellida Pennak, 1953 (6 families) Family Cristatellidae Allman, 1856 (1 genus, 1 species) Family Fredericellidae Hyatt, 1868 (2 genera, 8 species) Family Lophopodidae Rogick, 1935 (3 genera, 8 species) Family Pectinatellidae Lacourt, 1968 (2 genus, 4 species) Family Plumatellidae Allman, 1856 (6 genera, 64 species) Family Stephanellidae Lacourt, 1968 (1 genus, 1 species)
family) Family Cerioporidae Busk
  • Suborder Cerioporina
  • Von Hagenow
Suborder Cerioporina von Hagenow, 1851 (1 family) Family Cerioporidae Busk, 1859 (5 genera, 19 species)
families) Family Densiporidae Borg, 1944 (5 genera, 17 species) Family Lichenoporidae Smitt, 1867 (9 genera
  • Suborder Rectangulata Waters
Suborder Rectangulata Waters, 1887 (2 families) Family Densiporidae Borg, 1944 (5 genera, 17 species) Family Lichenoporidae Smitt, 1867 (9 genera, 54 species)
families) Family Arachnopusiidae Jullien, 1888 (5 genera, 43 species) Family Exechonellidae Harmer, 1957 (4 genera
  • Jullien Superfamily Arachnopusioidea
Superfamily Arachnopusioidea Jullien, 1888 (2 families) Family Arachnopusiidae Jullien, 1888 (5 genera, 43 species) Family Exechonellidae Harmer, 1957 (4 genera, 20 species)
families) Family Batoporidae Neviani genera, 11 species) Family Conescharellinidae Levinsen genera, 91 species) Family Lekythoporidae Levinsen genera, 23 species) Family Orbituliporidae Canu & Bassler
  • Superfamily Conescharellinoidealevinsen
Superfamily ConescharellinoideaLevinsen, 1909 (4 families) Family Batoporidae Neviani, 1900 (3 genera, 11 species) Family Conescharellinidae Levinsen, 1909 (6 genera, 91 species) Family Lekythoporidae Levinsen, 1909 (8 genera, 23 species) Family Orbituliporidae Canu & Bassler, 1923 (1 genus, 1 species)
Molecular phylogenetic analysis of phylactolaemate bryozoans based on mitochondrial gene sequences
  • M Hirose
  • M H Dick
  • S F Mawatari
Hirose, M., Dick, M.H. & Mawatari, S.F. (2008) Molecular phylogenetic analysis of phylactolaemate bryozoans based on mitochondrial gene sequences. In: Hageman, S.J., Key, M.M. Jr & Winston, J.E. (Eds), Bryozoan Studies 2007. Virginia Museum of Natural History Special Publication, 15, 65-74.
genera, 38 species) Family Thalamoporellidae Levinsen genera
Family Steginoporellidae Hincks, 1884 (3 genera, 38 species) Family Thalamoporellidae Levinsen, 1902 (6 genera, 65 species)
families) Family Haywardozoontidae d'Hondt, 1983 (1 genus, 2 species) Family Hislopiidae Jullien, 1885 (1 genus
  • Jullien Superfamily Hislopioidea
Superfamily Hislopioidea Jullien, 1885 (2 families) Family Haywardozoontidae d'Hondt, 1983 (1 genus, 2 species) Family Hislopiidae Jullien, 1885 (1 genus, 9 species)
1896 (4 families) Family Crisinidae d'Orbigny, 1853 (5 genera, 8 species) Family Horneridae Smitt, 1867 (4 genera, 22 species) Family Pseudidmoneidae Borg, 1944 (1 genus, 2 species) Family Stigmatoechidae Brood, 1972 (1 genus
  • Gregory Suborder Cancellata
Suborder Cancellata Gregory, 1896 (4 families) Family Crisinidae d'Orbigny, 1853 (5 genera, 8 species) Family Horneridae Smitt, 1867 (4 genera, 22 species) Family Pseudidmoneidae Borg, 1944 (1 genus, 2 species) Family Stigmatoechidae Brood, 1972 (1 genus, 2 species)
suborders) Suborder Malacostegina Levinsen superfamily) Superfamily Membraniporoidea Busk, 1852 (3 families) Family Electridae Stach, 1937 (15 genera, 80 species) Family Membraniporidae Busk genera, 45 species) Family Sinoflustridae Gordon
  • Cheilostomata Order
  • Busk
Order Cheilostomata Busk, 1852 (4 suborders) Suborder Malacostegina Levinsen, 1902 (1 superfamily) Superfamily Membraniporoidea Busk, 1852 (3 families) Family Electridae Stach, 1937 (15 genera, 80 species) Family Membraniporidae Busk, 1852 (3 genera, 45 species) Family Sinoflustridae Gordon, 2009 (2 genera, 7 species)
families) Family Cellariidae Fleming, 1828 (16 genera, 106 species) Family Membranicellariidae Levinsen, 1902 (2 genera
  • Fleming Superfamily Cellarioidea
Superfamily Cellarioidea Fleming, 1828 (2 families) Family Cellariidae Fleming, 1828 (16 genera, 106 species) Family Membranicellariidae Levinsen, 1902 (2 genera, 2 species)
family) Family Euthyrisellidae Bassler
Superfamily EuthyriselloideaBassler, 1953 (1 family) Family Euthyrisellidae Bassler, 1953 (5 genera, 6 species)
genera, 9 species) Family Hippothoidae Busk, 1859 (7 genera, 65 species) Family Haplopomidae Gordon in De Blauwe, 2009 (1 genus, 7 species) Family Pasytheidae Davis, 1934 (6 genera, 11 species) Family Trypostegidae Gordon
  • Busk Superfamily Hippothoidea
Superfamily Hippothoidea Busk, 1859 (5 families) Family Chorizoporidae Vigneaux, 1949 (2 genera, 9 species) Family Hippothoidae Busk, 1859 (7 genera, 65 species) Family Haplopomidae Gordon in De Blauwe, 2009 (1 genus, 7 species) Family Pasytheidae Davis, 1934 (6 genera, 11 species) Family Trypostegidae Gordon, Tilbrook & Winston in Winston, 2005 (1 genus, 8 species)
families) Family Adeonidae Busk, 1884 (10 genera, 106 species) Family Inversiulidae Vigneaux genus
  • Superfamily Adeonoidea
  • Busk
Superfamily Adeonoidea Busk, 1884 (2 families) Family Adeonidae Busk, 1884 (10 genera, 106 species) Family Inversiulidae Vigneaux, 1949 (1 genus, 4 species)
family) Family Urceoliporidae Bassler, 1936 (2 genera
  • Superfamily Urceoliporoidea
  • Bassler
Superfamily Urceoliporoidea Bassler, 1936 (1 family) Family Urceoliporidae Bassler, 1936 (2 genera, 3 species)
family) Family Didymosellidae Brown, 1952 (2 genera
  • Superfamily Didymoselloidea Brown
Superfamily Didymoselloidea Brown, 1952 (1 family) Family Didymosellidae Brown, 1952 (2 genera, 7 species)
families) Family Catenicellidae Busk, 1852 (20 genera, 85 species) Family Eurystomellidae Levinsen, 1909 (4 genera, 13 species) Family Petalostegidae Gordon, 1984 (2 genera, 11 species) Family Savignyellidae Levinsen, 1909 (2 genera
  • Busk Superfamily Catenicelloidea
Superfamily Catenicelloidea Busk, 1852 (4 families) Family Catenicellidae Busk, 1852 (20 genera, 85 species) Family Eurystomellidae Levinsen, 1909 (4 genera, 13 species) Family Petalostegidae Gordon, 1984 (2 genera, 11 species) Family Savignyellidae Levinsen, 1909 (2 genera, 2 species)
The Bryozoa home page
  • P E Bock
Bock, P.E. (2013) The Bryozoa home page. http://bryozoa.net/
Tabular keys for identification of the Recent ctenostomatous Bryozoa. Mémoires de l'Institut Océanographique
  • J.-L Hondt
Hondt, J.-L. d' (1983) Tabular keys for identification of the Recent ctenostomatous Bryozoa. Mémoires de l'Institut Océanographique, Monaco, 14, 1-134.
Zeitschrift für Zoologische Systematik und Evolutionsforschung
  • D Jebram
Jebram, D. (1992) The polyphyletic origin of the "Cheilostomata" (Bryozoa). Zeitschrift für Zoologische Systematik und Evolutionsforschung, 30, 46-52.
superfamily) Superfamily Aeteoidea Smitt
  • Suborder Inovicellina Jullien
Suborder Inovicellina Jullien, 1888 (1 superfamily) Superfamily Aeteoidea Smitt, 1868 (1 family)
genera, 66 species) Family Clavoporidae Osburn & Soule, 1953 (5 genera, 6 species) Family Flustrellidridae Bassler, 1953 (6 genera, 21 species) Family Lobiancoporidae Delage & Hérouard, 1897 (1 genus, 4 species) Family Pachyzoontidae d'Hondt, 1983 (1 genus, 1 species) Family
  • Johnston Superfamily Alcyonidioidea
Superfamily Alcyonidioidea Johnston, 1838 (6 families) Family Alcyonidiidae Johnston, 1838 (3 genera, 66 species) Family Clavoporidae Osburn & Soule, 1953 (5 genera, 6 species) Family Flustrellidridae Bassler, 1953 (6 genera, 21 species) Family Lobiancoporidae Delage & Hérouard, 1897 (1 genus, 4 species) Family Pachyzoontidae d'Hondt, 1983 (1 genus, 1 species) Family Pherusellidae Osburn & Soule, 1853 (1 genus, 3 species)
families) Family Pottsiellidae Braem, 1940 (1 genus, 1 species) Family Victorellidae Hincks, 1880 (4 genera, 10 species) Family Sundanellidae Jebram
  • Hincks Superfamily Victorelloidea
Superfamily Victorelloidea Hincks, 1880 (3 families) Family Pottsiellidae Braem, 1940 (1 genus, 1 species) Family Victorellidae Hincks, 1880 (4 genera, 10 species) Family Sundanellidae Jebram, 1973 (1 genus, 1 species)
genus, 2 species) Family Bathyalozoontidae d'Hondt genus, 1 species) Family Farrellidae d'Hondt genus, 1 species) Family Harmeriellidae d'Hondt genus, 1 species) Family Hypophorellidae Prenant & Bobin genus, 1 species) Family Mimosellidae Hincks
  • Hincks Superfamily Walkerioidea
Superfamily Walkerioidea Hincks, 1880 (8 families) Family Aeverrilliidae Jebram, 1973 (1 genus, 2 species) Family Bathyalozoontidae d'Hondt, 1975 (1 genus, 1 species) Family Farrellidae d'Hondt, 1983 (1 genus, 1 species) Family Harmeriellidae d'Hondt, 1983 (1 genus, 1 species) Family Hypophorellidae Prenant & Bobin, 1956 (1 genus, 1 species) Family Mimosellidae Hincks, 1977 (2 genera, 9 species) Family Triticellidae G. O. Sars, 1874 (2 genera, 12 species) Family Walkeriidae Hincks, 1880 (2 genera, 5 species)
family) Family Flustridae Fleming, 1828 (15 genera
  • Fleming Superfamily Flustroidea
Superfamily Flustroidea Fleming, 1828 (1 family) Family Flustridae Fleming, 1828 (15 genera, 80 species)
family) Family Chlidoniopsidae Harmer, 1957 (1 genus
  • Harmer Superfamily Chlidoniopsoidea
Superfamily Chlidoniopsoidea Harmer, 1957 (1 family) Family Chlidoniopsidae Harmer, 1957 (1 genus, 1 species)
genera, 28 species) Family Crepidacanthidae Levinsen, 1909 (1 genus, 14 species) Family Mamilloporidae Canu & Bassler, 1927 (2 genera
  • Family Ascosiidae Jullien
Family Ascosiidae Jullien, 1883 (2 genera, 5 species) Family Cleidochasmatidae Cheetham & Sandberg, 1964 (7 genera, 28 species) Family Crepidacanthidae Levinsen, 1909 (1 genus, 14 species) Family Mamilloporidae Canu & Bassler, 1927 (2 genera, 6 species)
family) Family Siphonicytaridae Harmer, 1957 (1 genus
  • Harmer Superfamily Siphonicytaroidea
Superfamily Siphonicytaroidea Harmer, 1957 (1 family) Family Siphonicytaridae Harmer, 1957 (1 genus, 12 species)
The central role of ctenostomes in bryozoan phylogeny
  • J A Todd
Todd, J.A. (2000) The central role of ctenostomes in bryozoan phylogeny. In: Herrera Cubila, A. & Jackson, J.B.C. (Eds), Proceedings of the 11th International Bryozoology Association Conference. Smithsonian Tropical Research Institute, Balboa, pp. 104-135.
orders) Order Ctenostomata Busk
  • Class Gymnolaemata Allman
Class Gymnolaemata Allman, 1856 (2 orders) Order Ctenostomata Busk, 1852 (8 superfamilies)
family) Family Paludicellidae Allman, 1836 (1 genus, 2 species) Family Panolicellidae Jebram
  • Superfamily Paludicelloidea Allman
Superfamily Paludicelloidea Allman, 1856 (1 family) Family Paludicellidae Allman, 1836 (1 genus, 2 species) Family Panolicellidae Jebram, 1985 (1 genus, 1 species)
family) Family Penetrantiidae Silén, 1946 (1 genus
  • Superfamily Penetrantioidea Silén
Superfamily Penetrantioidea Silén, 1946 (1 family) Family Penetrantiidae Silén, 1946 (1 genus, 8 species)
families) Family Arachnidiidae Hincks genera, 24 species) Family Immergentiidae Silén genus, 10 species) Family Monobryozoontidae Rémane genus, 3 species) Family Nolellidae Harmer
  • Hincks Superfamily Arachnidioidea
Superfamily Arachnidioidea Hincks, 1880 (4 families) Family Arachnidiidae Hincks, 1880 (7 genera, 24 species) Family Immergentiidae Silén, 1946 (1 genus, 10 species) Family Monobryozoontidae Rémane, 1936 (1 genus, 3 species) Family Nolellidae Harmer, 1915 (3 genera, 15 species)
genera, 11 species) Family Spathiporidae Pohowsky, 1978 (1 genus, 7 species) Family Terebriporidae d'Orbigny, 1847 (1 genus, 10 species) Family Vesiculariidae Johnston, 1847 (6 genera
  • Johnston Superfamily Vesicularioidea
Superfamily Vesicularioidea Johnston, 1847 (5 families) Family Buskiidae Hincks, 1880 (2 genera, 11 species) Family Spathiporidae Pohowsky, 1978 (1 genus, 7 species) Family Terebriporidae d'Orbigny, 1847 (1 genus, 10 species) Family Vesiculariidae Johnston, 1847 (6 genera, 68 species)
families) Family Bifaxariidae Busk, 1884 (6 genera, 46 species) Family Mixtopeltidae Gordon
  • Busk Superfamily Bifaxarioidea
Superfamily Bifaxarioidea Busk, 1884 (2 families) Family Bifaxariidae Busk, 1884 (6 genera, 46 species) Family Mixtopeltidae Gordon, 1994 (1 genus, 1 species)
family) Family Pseudolepraliidae Silén, 1942 (1 genus
  • Superfamily Pseudolepralioidea Silén
Superfamily Pseudolepralioidea Silén, 1942 (1 family) Family Pseudolepraliidae Silén, 1942 (1 genus, 1 species)
families) Family Cribrilinidae Hincks, 1879 (26 genera, 151 species) Family Euthyroididae Levinsen, 1909 (1 genus, 3 species) Family Polliciporidae Moyano
  • Hincks Superfamily Cribrilinoidea
Superfamily Cribrilinoidea Hincks, 1879 (3 families) Family Cribrilinidae Hincks, 1879 (26 genera, 151 species) Family Euthyroididae Levinsen, 1909 (1 genus, 3 species) Family Polliciporidae Moyano, 2000 (1 genus, 1 species)
Bryozoa: Cheilostomata. Interim classification for Treatise. Version of 1
  • D P Gordon
Gordon, D.P. (2012) Bryozoa: Cheilostomata. Interim classification for Treatise. Version of 1 November 2012. Unpublished. Available from http://www.bryozoa.net/treat_family_2012.pdf