Bryozoan Species Research Articles

La collection de bryozoaires du Prof. Dr Ehrhard Voigt (1905–2004) conservée à l'Institut Senckenberg de Francfort. 3e partie - Chéilostomates Ascophores et bibliographie

The bryozoan collection of Prof. Dr Ehrhard Voigt (1905–2004) at the Senckenberg Research Institute in Frankfurt am Main, Germany is a world-renowned collection of great scientific value. It is the world's largest collection of fossil bryozoans from the Upper Cretaceous and Paleocene and a unique archive documenting the evolution of this phylum of marine invertebrates during this time interval in the Boreal Chalk Sea that extended from the British Isles to the Aral Sea in Central Asia. The Voigt Collection contains over 300,000 specimens and was relocated to the Senckenberg Institute in 2005 according to the bequest of Ehrhard Voigt. As a result of a DFG-funded project, we present here a three-part type catalogue of the holotypes and neotypes of 256 bryozoan species in the Voigt Collection, of which this is Part 3. In total over the three parts, 247 species are re-illustrated but the name-bearing type specimens of 20 species are missing and no material could be found for 9 species. Two species, described as ctenostome bryozoans by Ehrhard Voigt, are questionable, while a further three 'ctenostome' species and one 'ctenostome' genus are considered as ichnotaxa.

Marine bryozoans of Gujarat and Maharashtra

Extensive field survey was conducted along the coast of Gujarat and in Maharashtra states during November 2015, 2016 and February 2016 respectively in selected reef localities. A total of 24 species of bryozoans belonging to 18 families were collected and identified. Of which eight species of bryozoans were recorded for first time in Indian coastal waters. More research should be encouraged in reef areas to throw more light on the marine bryozoan diversity in India.

An example of a possible leech-bryozoan association in freshwater.

Associations of various invertebrate species with bryozoans and sponges are a well-known marine phenomenon but such epizooic communities are far less diverse in freshwater environments. Here an occurrence of numerous leeches Alboglossiphoniacf.papillosa (Braun, 1805), in interstitial spaces between zooids of a colony of the freshwater bryozoan species Plumatellaaff.fungosa (Pallas, 1768) in Eastern Siberia is described. To the best of our knowledge, this record appears to be the first known example of a leech-bryozoan association, although such relationships deserve further research.

A checklist of marine bryozoan taxa in Scottish sea regions.

Contemporary and historical bryozoan records were compiled to provide a comprehensive checklist of species in Scottish waters. The checklist comprises 218 species in 58 families, with representatives from each of the extant bryozoan orders. The fauna was relatively sparse compared to other regions for which bryozoan checklists were available e.g. New Zealand and Australia. Six non-indigenous bryozoan species from the Scottish seas region were included in the checklist. Baseline information on species distributions, such as that presented in this checklist, can be used to monitor and manage the impact of human activities on the marine environment, and ultimately preserve marine biodiversity.

Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion-Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining.

This work describes Bryozoa of the orders Cyclostomata and Ctenostomata found associated with polymetallic nodules collected by box-coring in the eastern part of the Russian exploration area of the Clarion-Clipperton Fracture Zone (CCFZ) under contract to Yuzhmorgeologiya. Scanning electron microscopic study of 358 cyclostome colonies and 14 ctenostome colonies from 4510-5280 m depth has resulted in the recognition of two new species of Ctenostomata, and 14 new species, nine new genera and two new families of Cyclostomata; three additional species of Cyclostomata are left in open nomenclature pending the discovery of missing reproductive characters. The taxonomic novelty is thus notable. One of the new Ctenostomata represents the first living example of the previously monotypic Late Cretaceous genus Pierrella. Twelve of the new cyclostome taxa have well-developed gonozooids, indicating that embryonic cloning (polyembryony) is normal in this deep-sea environment. On the other hand, one indeterminate tubuliporine and two rectangulates have dimorphic peristomes. In the latter two cases, enough mature colonies were found to suggest that this feature is normal, and that the dimorphic zooids are possibly female-in other words, capacious incubation chambers are apparently lacking, and therefore polyembryony would also be lacking or reduced. In one of these species, evidence is presented to suggest that the ancestrular zooid can reproduce precociously. Of the species reported here, only one has previously been found outside the exploration area, highlighting both the limited knowledge we have of bryozoans in the deep Pacific and/or a fauna that is largely endemic to the nodule environment. An additional 31 species of Cheilostomata have also been discovered that will be described in a subsequent publication. Most bryozoans are macrofaunal-sized, so are both inadequately determinable and overlooked in images obtained by remotely operated vehicles; yet, with 50 species, Bryozoa is the most speciose sessile macrofaunal phylum on the nodules. Nodules constitute hard substrata in an area otherwise mostly inhospitable for Bryozoa, hence mining would lead to loss of critical habitat. Further, as suspension-feeders, bryozoans are highly susceptible to smothering by suspended sediment, and non-mined areas closely adjacent to extraction zones would likely also be affected and their associated bryozoan fauna obliterated. More data are required on the distribution of the CCFZ bryozoan species elsewhere in the east Central Pacific to determine if mining would lead to local taxon extirpation or global extinction at both low and high taxonomic levels.

Marine Biosecurity Crisis Decision-Making: Two Tools to Aid “Go”/“No Go” Decision-Making

Determining if a newly detected marine species is introduced to an area is an important first step when considering if control or eradication should be attempted. This step is often challenging, especially when data and introduced species expertise is limited: yet decisions about responding to a new invasion needs to occur in a timely manner. The crux is that biosecurity crisis decisions are often made in a vacuum. To improve this process, we consider expanded criteria to determine if a species is native, cryptogenic or introduced and outline application in a rapid response approach that uses a non-probabilistic decision tree to support decision makers. Effective use of the rapid response decision-tree and species criteria requires a multi-disciplinary approach drawing upon biology (taxonomy, phylogeny, genetics, ecology, biogeography) and monitoring. We assessed the expanded criteria against 213 bryozoan species present in Australian waters. A multivariate evaluation highlighted that a weight of evidence approach using the expanded criteria was successful in differentiating between native and introduced status. Our assessment highlighted that five criteria provide a high level of congruence with heuristic assignments, and provide a precautionary assignment of species' status by reducing mis-classifications of introduced species as native species (Type I error) in comparison to the original criteria. However, differentiating between introduced and cryptogenic species remains problematic, especially when using the original criteria. We highlight the critical need for taxonomic identification, appropriate application of assigning cryptogenic status, and monitoring requirements to enable use of the criteria in a rapid response context. Using both the rapid response decision tree and the criteria provides a quantifiable mechanism to aid decision-makers in deciding whether to respond to a marine species introduction.

Bryozoan framework composition in the oddly shaped reefs from Abrolhos Bank, Brazil, southwestern Atlantic: taxonomy and ecology.

Bryozoans are a key group of sessile invertebrates in some reef frameworks but are typically neglected in environmental monitoring programs. Abrolhos Bank (Brazil) is the largest reef complex in the South Atlantic Ocean, encompassing several reef landscapes over an area of 46,000 km2. A transition takes place across the shelf from mangroves to soft sediments, coastal shallow reefs to a volcanic archipelago - surrounded by fringing reefs - and unique mushroom-shaped biogenic structures, with mesophotic pinnacle reefs, rhodolith beds, sink-holes and shelf break deep environments. The taxonomic composition of the bryozoan fauna was studied in 11 core samples taken from shallow to mesophotic mid-shelf reefs (4-25 m deep) on Abrolhos Bank by divers using a submersible drill. Of the 20 bryozoan species sampled, 17 are new records for Abrolhos Bank and seven species are new to science: Crassimarginatella winstonae n. sp., Parasmittina distincta n. sp., Parasmittina abrolhosensis n. sp., Hemismittoidea asymmetrica n. sp., Stylopoma variabilis n. sp., Stylopoma hastata n. sp., and Plesiocleidochasma acuminata n. sp. (described by Ramalho, Taylor Moraes). The most conspicuous species is Celleporaria atlantica. These results increase to 48 the total number of bryozoan species known in this region and reinforce the importance of this group as one of the main components apart from crustose coralline algae and corals of the reef framework-building community of Abrolhos Bank.

Bryozoa from the continental shelf off Tierra del Fuego (Argentina): Species richness, colonial growth-forms, and their relationship with water depth

As bryozoans have a rich fossil record, the analysis of their bathymetric ranges and the occurrence of different colonial growth-forms in living assemblages may be valuable for palaeoecological interpretation. The bryozoan fauna from 58 benthic stations (18–92 m) collected on the Atlantic continental shelf off Tierra del Fuego (Southwest Atlantic) was analyzed to assess its biodiversity and the relationship of species richness and colonial growth-forms with depth. In the 58 stations analyzed here, which covered an area of ∼5600 km2, 90 species were found. Of these, 43 species were already known for this area, whereas the remaining 47 are new records. Therefore, the present study increases by 87% the known biodiversity for the study area, which now reaches 101 species. There is a clear increase in bryozoan species richness with depth, but the ratio of erect-rigid/encrusting species remained unchanged, as all the growth-forms increased with depth. Stations were classified by cluster analysis in three groups differing mainly in species richness. Taxonomic turnover (beta diversity) increased with increasing differences in depth between stations. This was due to the presence of a new set of deeper species, but not to the demise of the shallower fauna in the deeper stations. Twenty-two potentially habitat-forming species were found along the bathymetric range surveyed in this study. They were absent at the shallowest station, its number was relatively low at intermediate depths (31–64 m) and then experienced a sharp increase at 70 m. Palaeoecological interpretations should take into account that fragile, calcareous growth-forms could be expected to occur at shallower depths in more protected areas.

Young species of cupuladriid bryozoans occupied new Caribbean habitats faster than old species

The breadth of habitat occupied by a species, and the rate at which a species can expand into new habitats has important ecological and evolutionary consequences. Here we explore when extant species of free-living cupuladriid bryozoans expanded into new benthic Caribbean habitats that emerged during the final stages of formation of the Isthmus of Panama. Habitat breadth was estimated using the abundances of over 90,000 colonies in ten cupuladriid species, along with the ecological and sedimentary characteristics of the samples in which they occurred. Data reveal that all species expanded their habitat breadths during the last 6 Myr, but did so at a different tempo. ‘Young’ species - those that originated after 5 Ma - expanded relatively quickly, whereas ‘old’ species - those that originated before 9 Ma - took a further 2 Myr to achieve a comparable level of expansion. We propose that, like invasive species, young species are less restrained when expanding their habitat breadths compared to older well-established species. Understanding the mechanism causing this restraint requires further research.

Compositional Differences in the Habitat-Forming Bryozoan Communities of the Antarctic Shelf

In some areas of the Antarctic shelf, bryozoans are abundant, acting as ecosystem engineers creating secondary structures with wide benthic coverage and harboring numerous other species. As the combined forces of global warming and ocean acidification threaten these habitats, we measured the composition of habitat-forming bryozoan communities using two techniques for imaging the sea floor, a YoYo-camera system and the AWI Ocean Floor Observation System (OFOS). YoYo-camera transects of the Bellingshausen, Amundsen, and Ross Seas were conducted during a research cruise on the R/V Nathaniel B. Palmer in 2013. OFOS transects included sites in the northern Palmer Archipelago where it borders the Scotia Sea and the Weddell Sea as part of the DynAMO project during the PS81 and PS96 cruises of R/V Polarstern in 2013 and 2015-16, respectively. Areas of bryozoan colonies were measured from the sea floor images using machine-learning algorithms available through the Trainable Weka Segmentation plugin developed for FIJI software. Habitat-forming bryozoan communities in the Palmer Archipelago and Ross Sea were largely composed of anascan flustrid species with finely mineralized skeletons, and to a lesser extent by other ascophoran lepraliomorph and umbonulomorph species having more robustly mineralized skeletons. Although habitat-forming bryozoan communities in the shallower (200 m) sites of the Weddell Sea also contained flustrid species, percent area and composition of flustrid bryozoans declined with increasing depth. Lepraliomorph and umbonulomorph bryozoan morphotypes were more abundant in the Weddell Sea, maintaining their relative percent area and increasing their percent composition between 200 - 400 m. Moreover, our analyses of species composition based on externally gathered datasets show similar trends among sites, depths, and degrees of colony mineralization to our seabed imaging study. Variation present in the bryozoan species compositions of the Amundsen and Bellingshausen Seas suggest that these areas potentially represent divergent bryozoan communities requiring further validation via remote imaging surveys. Overall, compositional differences among Antarctic habitat-forming bryozoan communities are likely influenced by the combined effects of seasonal ice scour and carbonate chemistry, which in an increasingly acidified and warming ocean may put the communities of the eastern Weddell Sea at greater risk.

SYMBIOSIS OF CORNULITIDS AND BRYOZOANS IN THE LATE ORDOVICIAN OF ESTONIA (BALTICA)

Three species of trepostome bryozoans formed syn vivo associations with the Cornulites in the Late Ordovician of Estonia. Cornulites sp. and Mesotrypa excentrica presumably formed a true symbiotic association. This is the first known case of symbiosis between cornulitids and bryozoans. It is not known whether this symbiotic association was obligatory of facultative for the cornulitid, but it was facultative for the bryozoan. In this association cornulitids may have competed for the food with bryozoans and the association may have been parasitic. The remaining associations between cornulitids and bryozoans were accidental. Most common skeletonized endobionts of the Ordovician bryozoans were not cornulitids, but conulariids and rugosans.

Freshwater bryozoans of Lithuania (Bryozoa).

Nine species of freshwater bryozoans were recorded in Lithuania in a survey of 18 various types of freshwater bodies. Eight species were assigned to the Class Phylactolaemata and families Plumatellidae and Cristatellidae (Plumatella repens, Plumatella fungosa, Plumatella fruticosa, Plumatella casmiana, Plumatella emarginata, Plumatella geimermassardi, Hyalinella punctata and Cristatella mucedo). The ninth species, Paludicella articulata, represented the Class Gymnolaemata. Plumatella geimermassardi and P. casmiana were recorded for the first time in Lithuania. For the plumatellids, species identification was achieved partly by analysing statoblasts’ morphological ultrastructures by scanning electron microscopy.

Bryozoa on disarticulated bivalve shells from Todos os Santos Bay, northeastern Brazil, with the description of two new species.

As sessile organisms, bryozoans are among the commonest components of the encrusting communities on hard substrata. On unconsolidated bottoms, benthic animals and their carapaces, including mollusk shells, represent potential surfaces for encrustation. Here we describe the bryozoan fauna growing on disarticulated bivalve shells collected intertidally on Itaparica beach, Todos os Santos Bay, northeast Brazil. We examined a total of 382 valves representing seven species of bivalves, among which we found 82 valves of three bivalve species to be encrusted by bryozoans, with 109 bryozoan colonies detected overall. We identified 11 cheilostome bryozoan species, among which Phidoloporidae and Smittinidae were the most diverse, with four and two species, respectively. We redescribed two bryozoan species, Parasmittina loxoides and Rhynchozoon phrynoglossum, and describe another two, Pleurocodonellina marcusi n. sp. and Rhynchozoon itaparicaensis n. sp., as new. A new combination for Cribella triangulata Canu Bassler, 1928 is proposed, including Calyptotheca tenuata Harmer, 1957 as a new junior subjective synonym of Calyptotheca triangulata n. comb. All but one of the 11 species have encrusting colonies; Licornia aff. diadema, forms erect branching colonies. For seven of the bryozoans, ours are the first records of these species growing on shells. Although the majority of bryozoans we studied have also been reported on other substrata, Rhynchozoon brasiliensis and Parasmittina loxoides were predominant in the fauna, comprising 79 (72%) of all colonies inventoried. Due to the lack of information on the distribution of these species in northeastern Brazil, further studies are needed to determine whether they show any substratum preference.

Skeletal carbonate mineralogy of Scottish bryozoans.

This paper describes the skeletal carbonate mineralogy of 156 bryozoan species collected from Scotland (sourced both from museum collections and from waters around Scotland) and collated from literature. This collection represents 79% of the species which inhabit Scottish waters and is a greater number and proportion of extant species than any previous regional study. The study is also of significance globally where the data augment the growing database of mineralogical analyses and offers first analyses for 26 genera and four families. Specimens were collated through a combination of field sampling and existing collections and were analysed by X-ray diffraction (XRD) and micro-XRD to determine wt% MgCO3 in calcite and wt% aragonite. Species distribution data and phylogenetic organisation were applied to understand distributional, taxonomic and phylo-mineralogical patterns. Analysis of the skeletal composition of Scottish bryozoans shows that the group is statistically different from neighbouring Arctic fauna but features a range of mineralogy comparable to other temperate regions. As has been previously reported, cyclostomes feature low Mg in calcite and very little aragonite, whereas cheilostomes show much more variability, including bimineralic species. Scotland is a highly variable region, open to biological and environmental influx from all directions, and bryozoans exhibit this in the wide range of within-species mineralogical variability they present. This plasticity in skeletal composition may be driven by a combination of environmentally-induced phenotypic variation, or physiological factors. A flexible response to environment, as manifested in a wide range of skeletal mineralogy within a species, may be one characteristic of successful invasive bryozoans.

Mineralogical and Elemental Composition of Pectinatella magnifica and Its Statoblasts

Several massive colonies of Pectinatella magnifica have been observed during the summer almost every year since 1974 in agricultural reservoir ponds and lakes with dirty freshwater environments in Ishikawa, Japan, which has posed serious environmental problems on the shores of Hokuriku District. We collected Pectinatella magnifica during the summer at Kahokugata Lake and Makiyama agricultural reservoir pond in June and July 2016. However, scientific data for the statoblasts of Pectinatella magnifica are limited. Our results for scanning electron microscopy equipped with energy-dispersive spectroscopy (SEM-EDS), inductively coupled plasma-mass spectrometry (ICP-MS), and X-ray powder diffraction (XRD) analyses of Pectinatella magnifica indicated immobilization of the chemical elements that were involved in the mass during the summer. We also reported the characterization of an invasive species of bryozoan (Pectinatella magnifica) in lakes and ponds in Ishikawa, Japan, based on field observations in 2016. We studied the microstructure, mineralogy, chemical composition, and radioactivity associated with these organisms, using a combination of micro-techniques, SEM-EDS, associated with ICP-MS, and XRD. This study aims to illustrate the capability of Pectinatella magnifica to produce minerals within statoblasts and gelatinous material. Obtained results may indicate forming quartz, palygorskite, dolomite, bischofite, pyrolusite, and pyrite, associated with native sulfur and copper in the statoblast. The mass of gelatinous material contains talc and vermiculite as well as non-crystalline phase. The mechanism of biomineral formation has important implications for water–mineral–organism or microorganism interactions both in lower drainage basin systems, such as Kahokugata Lake, and upper water areas, such as Makiyama agricultural reservoir pond. Many types with variety of sizes and shapes of bryozoan (Pectinatella magnifica) were found in lakes and ponds in Japan. The biomineralization systems will be made available for use not only in researching bryozoans (Pectinatella magnifica), but also for environmental change systems from upstream to downstream of the lake. To date, there have been no reports on related electron microscopy observations, including the real-life occurrence of “bioremediation”. These observations could lead to simple methods of removing statoblasts of the invasive alien species Pectinatella magnifica from agricultural and reservoir environments, because there was limited microbial immobilization of the ions during the winter.

Experimental cultivation of the invasive freshwater bryozoan Pectinatella magnifica

The freshwater bryozoan Pectinatella magnifica (Leidy, 1851) is an invasive species in many countries all over the world. Although native to North America, it has been found in many countries of Europe and Asia. Pectinatella magnifica forms the largest colonial masses from all recently known bryozoan species. Culturing this organism in an aquarium has never been achieved for more than few days so far. Colonies from laboratory culture are important for various studies on its biology and life cycle of this species in experimental conditions. Young colonies successfully hatched from germinating statoblasts of P. magnifica in the laboratory and were maintained over eight weeks. Moreover, this was the first time when the compound colonies of this species were carried from its natural habitat to the laboratory, into a special aquarium system, and kept alive for more than three weeks. In both experiments the physicochemical parameters of the water (temperature, concentration of dissolved oxygen, electrolytic conductivity and pH) and changes in weight of compound colonies of P. magnifica in laboratory conditions were checked. The results found in this study are essential for understanding the invasiveness of this species and identifying methods for elimination of its ecological risks because these are closely resembling those of other invasive species.

Mediterranean non-indigenous bryozoans: an update and knowledge gaps

This paper provides an inventory of non-indigenous bryozoan species (NIB) recorded in the Mediterranean Sea. Taking into account previous NIB lists and updated non-indigenous concept, a careful literature review was carried out, reporting data on 33 NIB from 14 Mediterranean countries (Algeria, Croatia, Cyprus, Egypt, France, Greece, Israel, Italy, Lebanon, Malta, Spain, Syria, Tunisia and Turkey). A total of 296 valid records were listed, including 6 yet unpublished. Other 62 records were labelled as uncertain and listed separately, until further analyses on taxonomic identity, non-indigenous status and distribution patterns of the species involved will clarify their status. The countries with the higher numbers of NIB and single records are Lebanon (20 and 78, respectively) and Italy (12 and 112, respectively). The distribution of NIB in the Mediterranean Sea appears scattered and no data are available at all for eight countries. Studies and monitoring efforts on bryozoans in the Mediterranean Sea are not evenly distributed, probably as a consequence of the decrease in number of expert taxonomists and the poor allocation of funds on biodiversity studies. The coordination among Mediterranean countries on standardised sampling methods is needed in order to create a comprehensive baseline knowledge on NIB distribution in the Mediterranean Sea and prioritise taxa and countries that are still poorly investigated.

Sixteen new generic records of Korean Bryozoa from southern coastal waters and Jeju Island, East China Sea: evidence of tropical affinities.

The first records of 16 genera of cheilostome Bryozoa not previously reported for Korea are given here. All are from southern coastal waters, especially Jeju Island, which appears to be Korea's marine-biodiversity 'hotspot'. Jodoella koreensis n. gen. et sp. (Robertsonidridae) is new to science. Seven additional new species are described for the genera Copidozoum, Reptadeonella, Schizosmittina, Saevitella, Torquatella, Stephanollona and Plesiocleidochasma. The other genera new to the Korean fauna are Dibunostoma, Corbulella, Onychocella, Stephanotheca, Bryopesanser, Calloporina, Hippaliosina and Pleuromucrum. The Japanese Pleistocene species Calloporina hayamiae Arakawa is newly reported alive from Jeju Island. Four of the species were previously known only from the tropical Indo-West Pacific, three others are found in semitropical/warm-temperate water, and the other known genera have warm-water species, supporting data from other taxa that Korean's southern waters, especially in the vicinity of Jeju Island, are profoundly influenced by the warm Kuroshio Current and global warming trends. Currently, at least 112 species of Bryozoa are known from Jeju Island, 101 of them Cheilostomata, representing c. 70% of all Korean cheilostome species. Four new combinations are introduced: Copidozoum canui (Sakakura) n. comb., Schizo pedicellata (Soule, Soule Chaney) n. comb., Torquatella ensenada (Tilbrook) n. comb. and Torquatella longiuscula (Harmer) n. comb. Torquatellidae is subsumed in Celleporidae and Predanophora in Torquatella.

Early Pleistocene and Holocene bryozoans from Indonesia.

This paper describes 40 bryozoan species, comprising one cyclostome and 39 cheilostomes (8 anascan- and 31 ascophoran-grade), obtained from early Pleistocene and Holocene samples from two localities in Indonesia. Five of the cheilostomes are described as new species: Acanthodesia variegata n. sp., Pleurocodonellina javanensis n. sp., Calyptotheca sidneyi n. sp., Characodoma wesselinghi n. sp., and Turbicellepora yasuharai n. sp. All of the remaining species are extant and characterized by a tropical to subtropical Indo-West Pacific distribution.

The Madagascan Maastrichtian bryozoans of Ferdinand Canu – Systematic revision and scanning electron microscopic study

In 1922, the first taxonomic work on Maastrichtian bryozoans from Madagascar was published in the Annales de Paléontologie by Ferdinand Canu. Canu described 25 species, 17 of which were regarded as new but the material has never been revised until now. Here we employ scanning electron microscopy (SEM) to illustrate Canu's material and revise the taxonomy. Based on this revision, 23 species, comprising eight cyclostomes and 15 cheilostomes, are recognized. A new genus, Fehiborypora, is introduced for the cribrimorph Cribilina (?) labiatula, and two new species are described: ‘Plagioecia’ antanihodiensis sp. nov. and ‘Escharoides’ charbonnieri sp. nov. Galeopsis parvipora is synonymized with Crustoporina prona Stoliczka. All bryozoan species found are encrusters on echinoids or oysters.