Structure of the oral tentacles of early ontogeny stage in brachiopod Hemithiris psittacea (Rhynchonelliformea, Rhynchonellida).

Brachiopods have the most complex lophophore in comparison with other lophophorates, i.e., phoronids and bryozoans. However, at early ontogenetic stages, brachiopods have a lophophore of simple morphology, which consists of the oral tentacles. Data on the ultrastructure of the oral tentacles is mostly missing. Nonetheless, it has recently been suggested that the structure of oral tentacles is ancestral for all lophophorates in general, and for brachiopods in particular. The fine structure of the oral tentacles in the brachiopod Hemithiris psittacea is studied using light microscopy, transmission and scanning electron microscopy, cytochemistry and confocal laser scanning microscopy. The oral tentacles have a round shape in transverse section, and four ciliary zones, i.e., one frontal, two lateral, and one abfrontal. Latero-frontal sensory cells occur among the frontal epithelium. Four basiepithelial nerves in the ciliary epithelium are colocalized with ciliary zones. Lophophores of simple morphology in phoronids and brachiopods are characterized by non-specified round forms of tentacles. In phoronids and bryozoans, tentacles have additional latero-frontal ciliary zones that function as a sieve during filtration. In most brachiopods, lateral cilia are involved in the capture of food particles, whereas latero-frontal cells are retained in the frontal zone as sensory elements. The oral tentacles of H. psittacea contain a coelomic canal and have distinct frontal and abfrontal longitudinal muscles, which are separated from each other by peritoneal cells. A similar structure of tentacle muscles occurs in all bryozoans, whereas in phoronids, the frontal and abfrontal tentacle muscles are not separated by peritoneal cells. We suggest that the lophophorates' ancestor had tentacles, which were similar to the tentacles of some phoronids with lophophore of simple morphology. We also assume that the structure of the oral tentacles is ancestral for all brachiopods and the specialization of brachiopod tentacles correlates with the appearance of the double row of tentacles.

Morphology of ctenostome bryozoans: 7. Hislopia, Echinella and Timwoodiellina.

Ctenostome bryozoans are a small group of gymnolaemates comprising less than 400 recent species. They are paraphyletic and ctenostome-grade ancestors gave rise to Cheilostomata, the most dominant and speciose taxon of Bryozoa in the present day. Investigations into ctenostomes are important for reconstructing character evolution among Gymnolaemata. As a continuation of studies on a morphological series of ctenostome bryozoans, we herein investigate six species of hislopiids, a small clade of three genera occurring in freshwater habitats. The general morphology of all species is similar in having primarily uniserial chains of encrusting zooids, which are mostly oval to ellipsoid and have a flattened frontobasal axis. Hislopia prolixa and Echinella placoides often have more slender zooids with a higher frontobasal axis. Apertures of hislopiids are quadrangular, lined by a thickened cuticle. Apertural spines are present in various lengths in E. placoides, Hislopia lacustris and Hislopia corderoi. The remaining cuticle is rather thin except at lateral areas, close to the pore-plates which are prominent in hislopiids because of abundant special and limiting cells. All species except H. corderoi and Timwoodiellina natans have a prominent collar obstructing the vestibulum, whereas the latter two species instead have an 'external collar' as cuticular, outer folds projecting over the aperture. Hislopiid lophophores carry eight, or more commonly 12-18 tentacles. The digestive tract is distinguished by an often highly elongated esophagus and/or cardia, with the latter always having a prominent bulbous part in the form of a proventriculus-or gizzard in E. placoides. The caecum is extensive in all species. In Hislopia the intestine is characteristically two-chambered with a proximal and distal part before entering an anal tube of various length. The latter is present in all species except T. natans and terminates in mid-lophophoral area. Oocytes in E. placoides are large and macrolecithal indicating brooding and the production of lecithotrophic larvae. Hislopia species produce small, oligolecithal ones, which suggests zygote spawning and planktotrophy. In general, the morphology is similar among the different hislopiids with characters of the gut aiding in delineating the genera Echinella and Timwoodiellina.

The unplumatellid Plumatella fruticosa found its home: Hirosella gen. nov. morphological arguments for the systematic placement of a freshwater bryozoan.

Bryozoans are colonial, suspension-feeding lophotrochozoans. The phylum consists of the large group of chiefly marine Myolaemata and the exclusively limnic Phylactolaemata. Each colony consists of individual zooids that comprise the protective cystid and the retractable polypide. Phylactolaemates are a small group of approximately 90 species in 6 families. They feature a body wall, that can either be gelatinous, as in the families Stephanellidae, Lophopodidae, Cristatellidae and Pectinatellidae, or encrusted, as in Plumatellidae and Fredericellidae. Morphological investigations of the most specious plumatellids are rare and focus on few species. Plumatella fruticosa is of particular interest in this regard, as it shows a mosaic of plumatellid and fredericellids characters. The most recent phylogeny clusters P. fruticosa with cristatellids and pectinatellids as sister groups to fredericellids. Hence, there is considerable doubt, whether P. fruticosa is truly a plumatellid. Therefore, this study aims to reinvestigate the morphology of P. fruticosa with confocal microscopy and section-based three-dimensional reconstruction. The new data show that P. fruticosa has numerous conspicuous stumps from fragmented proliferation buds, which are otherwise only known from fredericellids. Like fredericellids, P. fruticosa grows erect, but in contrast, has a horseshoe-shaped lophophore and floatoblasts. Besides the proportions of the lophophore, the tentacle sheath and digestive tract resemble a fredericellid-like situation. Myoanatomical details like the pronounced longitudinal muscles of the vestibular wall and tentacle sheath differ from plumatellids and favour the recently proposed scenario, which places P. fruticosa next to Pectinatellidae and Cristatellidae. In addition, the intertentacular membrane of P. fruticosa shows structural similarity to cristatellids as it is attached to the tentacles via lamellae. Taking all aspects into account, we erect a new family: Hirosellidae fam. nov. including the new genus Hirosella gen. nov.

Protomelission is an early dasyclad alga and not a Cambrian bryozoan.

The animal phyla and their associated body plans originate from a singular burst of evolution occurring during the Cambrian period, over 500 million years ago1. The phylum Bryozoa, the colonial 'moss animals', have been the exception: convincing skeletons of this biomineralizing clade have been absent from Cambrian strata, in part because potential bryozoan fossils are difficult to distinguish from the modular skeletons of other animal and algal groups2,3. At present, the strongest candidate4 is the phosphatic microfossil Protomelission5. Here we describe exceptionally preserved non-mineralized anatomy in Protomelission-like macrofossils from the Xiaoshiba Lagerstätte6. Taken alongside the detailed skeletal construction and the potential taphonomic origin of 'zooid apertures', we consider that Protomelission is better interpreted as the earliest dasycladalean green alga-emphasizing the ecological role of benthic photosynthesizers in early Cambrian communities. Under this interpretation, Protomelission cannot inform the origins of the bryozoan body plan; despite a growing number of promising candidates7-9, there remain no unequivocal bryozoans of Cambrian age.

Unusual lophophore innervation in ctenostome Flustrellidra hispida (Bryozoa).

Since ctenostomes are traditionally regarded as an ancestral clade to some other bryozoan groups, the study of additional species may help to clarify questions on bryozoan evolution and phylogeny. One of these questions is the bryozoan lophophore evolution: whether it occurred through simplification or complication. The morphology and innervation of the ctenostome Flustrellidra hispida (Fabricius, 1780) lophophore have been studied with electron microscopy and immunocytochemistry with confocal laser scanning microscopy. Lophophore nervous system of F. hispida consists of several main nerve elements: cerebral ganglion, circumoral nerve ring, and the outer nerve ring. Serotonin-like immunoreactive perikarya, which connect with the circumoral nerve ring, bear the cilium that directs to the abfrontal side of the lophophore and extends between tentacle bases. The circumoral nerve ring gives rise to the intertentacular and frontal tentacle nerves. The outer nerve ring gives rise to the abfrontal neurites, which connect to the outer groups of perikarya and contribute to the formation of the abfrontal tentacle nerve. The outer nerve ring has been described before in other bryozoans, but it never contributes to the innervation of tentacles. The presence of the outer nerve ring participating in the innervation of tentacles makes the F. hispida lophophore nervous system particularly similar to the lophophore nervous system of phoronids. This similarity allows to suggest that organization of the F. hispida lophophore nervous system may reflect the ancestral state for all bryozoans. The possible scenario of evolutionary transformation of the lophophore nervous system within bryozoans is suggested.

Reconstructing the neuromuscular ground pattern of phylactolaemate bryozoans: new data from the Lophopodidae.

BACKGROUND: The solely freshwater inhabiting Phylactolaemata is a sister taxon to all other bryozoans. Among phylactolaemates, Lophopodidae represents an early branching clade that is therefore crucial for ground pattern reconstruction. While more recent morphological data of most phylactolaemate families are present, data of lophopodids are scarce. The genus Asajirella especially, which was previously assigned to the family Pectinatellidae, lacks any detailed analysis with more recent morphological methods. RESULTS: This study provides the first morphological analyses of three lophopodid species using serial-sectioning histology and 3D reconstruction, but also immunocytochemical stainings and confocal laserscanning microscopy. There are several lophopodid-specific traits in the nervous system such as the large ganglion with extensive lumen and two prominent protrusions referred to as epistomial horns. The epistome in all lophopodids is rather small and dome-shaped. Contrary to previous reports, we can confirm that duplicature bands insert at the tentacle sheath rather than the diaphragmatic sphincter in all phylactolaemates. The morphology of the digestive tract of lophopodids is identical to other phylactolaemates and possesses exclusively circular muscles. CONCLUSIONS: Altogether, this study fills significant gaps in our knowledge on phylactolaemate neuromuscular systems and general morphology. It shows that the insertion of the duplicature bands at the tentacle sheath and the circular musculature of the digestive tract to be the ground pattern in phylactolaemates. In addition, we found apomorphic characters for lophopodids such as the dome-shaped epistome with its musculature and the voluminous ganglion with its epistomial horns, which aid in defining and delineating the family.

Morphology of ctenostome bryozoans: 6. Amphibiobeania epiphylla.

Ctenostome bryozoans are unmineralized and mostly marine. Their lack of calcified skeletal features requires other characters to be considered for systematic and phylogenetic considerations. As a continuation of an ongoing series of studies, we herein investigate the morphology of Amphibiobeania epiphylla, a unique bryozoan inhabiting mangrove leaves that are highly exposed to tidal cycles and regular dry events according to the tidal cycle. Besides this interesting mode of life, the species was originally interpreted to be a weakly mineralized cheilostome bryozoan, whereas molecular data place it among ctenostome bryozoans. To elucidate the systematic and phylogenetic position of the genus and also find morphological adaptations to an extreme habitat, we investigated the morphology of A. epiphylla in detail. Zooids show a lophophore with eight tentacles and a simple gut with a prominent caecum, lophophoral anus and most notably a distinct gizzard in the cardiac region. Gizzard teeth are multiple, simple homogeneous cuticular structures. The cuticle of the zooid is rather uniform and shows no respective thickenings into opercular flaps or folds. Likewise, apertural muscles are represented by a single pair of muscles. There are no specific closing muscles in the apertural area like the operculum occlusors of cheilostomes. Most prominent within zooids is a spongiose tissue filling most of the body cavity. Although not properly understood, this tissue may aid in keeping animals moist and hydrated during prolonged dry times. In summary, all morphological characters support a ctenostome rather than a cheilostome affinity, possibly with Vesicularioidea or Victorelloidea. In addition, we provide new molecular data that clearly supports such a closer relationship.

New species and new records of Reteporella (Bryozoa: Cheilostomatida) from Greenland waters.

New species and new records of the cheilostome bryozoan family Phidoloporidae are reported from the poorly studied shelf and continental slope of Greenland. Reteporella vitta n. sp. differs from congeners in the shape and size of fenestrulae, the size of autozooids and their arrangement within the colony, features of the orificial complex, including the indistinctly denticulate distal rim, shape of the condyles and height of the peristome, as well as the shape and location of the suboral avicularium, and the morphology of the ovicell. Reteporella obscura n. sp. differs from other Reteporella species in a unique combination of the following characters: colony surface texture, shape of orifice and condyles, shape of the suboral avicularium, and the morphology of the ovicell. Other specimens of Reteporella from Greenland were identified as R. watersi (Nordgaard, 1907), firstly recorded in southern and western parts of the study area, and as R. grimaldii (Jullien, 1903) and R. beaniana (King, 1846), previously known only from some localities of western and eastern Greenland and recorded abundantly in 2016. Reteporella watersi specimens from Greenland differ from conspecific material from the Faroe Islands in lacking oral spines and in the size of zooidal characters.

Morphology of ctenostome bryozoans: 5. Sundanella, with description of a new species from the Western Atlantic and the Multiporata concept.

Ctenostome bryozoans are a small group of gymnolaemates that comprise only a few hundred described species. Soft-tissue morphology remains the most important source for analysing morphological characters and inferring relationships within this clade. The current study focuses on the genus Sundanella, for which morphological data is scarce to almost absent. We studied two species of the genus, including one new to science, using histology and three-dimensional reconstruction techniques and confocal laser scanning microscopy. Sundanella generally has a thick, sometimes arborescent cuticle and multiporous interzooidal pore plates. The lophophore is bilateral with an oral rejection tract and generally has 30 or 31 tentacles in both species. The digestive tract shows a large cardia in S. floridensis sp. nov. and an extremely elongated intestine in Sundanella sibogae. Both terminate via a vestibular anus. Only parietodiaphragmatic muscles are present and four to six duplicature bands. Both species show a large broad frontal duplicature band further splitting into four individual bands. The collar is vestibular. Sundanella sibogae shows highly vacuolated cells at the diaphragm, whereas S. floridensis sp. nov. has unique glandular pouches at the diaphragmal area of the tentacle sheath. Such apertural glands have never been encountered in other ctenostomes. Both species of Sundanella are brooders that brood embryos either in the vestibular or cystid wall. Taken together, the current analysis shows numerous characteristics that refute an assignment of Sundanella to victorellid ctenostomes, which only show superficial resemblance, but differ substantially in most of their soft-body morphological traits. Instead, a close relationship with other multiporate ctenostomes is evident and the families Pherusellidae, Flustrellidrae and Sundanellidae should be summarized as clade 'Multiporata' in the future.

Composite branch construction by dual autozooidal budding modes in hornerids (Bryozoa: Cyclostomatida).

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-computed tomography and scanning electron microscopy 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 centered 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.

The epithelial layers of the body wall in hornerid bryozoans (Stenolaemata: Cyclostomatida).

Bryozoans are small colonial coelomates. They can be conceptualised as "origami-like" animals, composed of three complexly folded epithelial layers: epidermis of the zooidal/colonial body wall, gut epithelium and coelothelium. We investigated the general microanatomy and ultrastructure of the hornerid (Cyclostomatatida) body wall and polypide in four taxa, including three species of Hornera and one species belonging to an undescribed genus. We describe epithelia and their associated structures (e.g., ECM, cuticle) across all portions of the hornerid body wall, including the terminal membrane, vestibular wall, atrial sphincter, membranous sac and polypide-skeletal attachments. The classic coelomate body wall composition (epidermis-ECM-coelothelium) is only present in an unmodified form in the tentacle sheath. Deeper within a zooid it is retained exclusively in the attachment zones of the membranous sac: [skeleton]-tendon cell-ECM-coelothelium. A typical invertebrate pattern of epithelial organisation is a single, continuous sheet of polarised cells, connected by belt desmosomes and septate junctions, and resting on a collagenous extracellular matrix. Although previous studies demonstrated that polypide-specific epithelia of Horneridae follow this model, here we show that the body wall may show significant deviations. Cell layers can lose the basement membrane and/or continuity of cell cover and cell contacts. Moreover, in portions of the body wall, the cell layer appears to be missing altogether; the zooidal orifice is covered by a thin naked cuticle largely devoid of underlying cells. Since epithelium is a two-way barrier against entry and loss of materials, it is unclear how hornerids avoid substance loss, while maintaining intracolonial metabolite transport with imperfect, sometimes incomplete, cell layers along large portions of their outer body surface.

Tentacle muscles in brachiopods: Ultrastructure and relation to peculiarities of life style.

Although the morphology of the brachiopod tentacle organ, the lophophore, is diverse, the organization of tentacles has traditionally been thought to be similar among brachiopods. We report here, however, that the structure of the tentacle muscles differs among brachiopod species representing three subphyla: Lingula anatina (Linguliformea: Linguloidea), Pelagodiscus atlanticus (Linguliformea: Discinoidea), Novocrania anomala (Craniiformea), and Coptothyris grayi (Rhynchonelliformea). Although the tentacle muscles in all four species are formed by myoepithelial cells with thick myofilaments of different diameters, three types of tentacle organization were detected. The tentacles of the first type occur in P. atlanticus, C. grayi, and in all rhynchonelliforms studied before. These tentacles have a well-developed frontal muscle and a small abfrontal muscle, which may reflect the ancestral organization of tentacles of all brachiopods. This type of tentacle has presumably been modified in other brachiopods due to changes in life style. Tentacles of the second type occur in the burrowing species L. anatina and are characterized by the presence of equally developed smooth frontal and abfrontal muscles. Tentacles of the third type occur in N. anomala and are characterized by the presence of only well-developed frontal muscles; the abfrontal muscles are reduced due to the specific position of tentacles during filtration and to the presence of numerous peritoneal neurites on the abfrontal side of the tentacles. Tentacles of the first type are also present in phoronids and bryozoans, and may be ancestral for all lophophorates.

Fossil evidence unveils an early Cambrian origin for Bryozoa.

Bryozoans (also known as ectoprocts or moss animals) are aquatic, dominantly sessile, filter-feeding lophophorates that construct an organic or calcareous modular colonial (clonal) exoskeleton1-3. The presence of six major orders of bryozoans with advanced polymorphisms in lower Ordovician rocks strongly suggests a Cambrian origin for the largest and most diverse lophophorate phylum2,4-8. However, a lack of convincing bryozoan fossils from the Cambrian period has hampered resolution of the true origins and character assembly of the earliest members of the group. Here we interpret the millimetric, erect, bilaminate, secondarily phosphatized fossil Protomelission gatehousei9 from the early Cambrian of Australia and South China as a potential stem-group bryozoan. The monomorphic zooid capsules, modular construction, organic composition and simple linear budding growth geometry represent a mixture of organic Gymnolaemata and biomineralized Stenolaemata character traits, with phylogenetic analyses identifying P. gatehousei as a stem-group bryozoan. This aligns the origin of phylum Bryozoa with all other skeletonized phyla in Cambrian Age 3, pushing back its first occurrence by approximately 35 million years. It also reconciles the fossil record with molecular clock estimations of an early Cambrian origination and subsequent Ordovician radiation of Bryozoa following the acquisition of a carbonate skeleton10-13.

A broadly resolved molecular phylogeny of New Zealand cheilostome bryozoans as a framework for hypotheses of morphological evolution.

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.

Mitochondrial gene order of the freshwater bryozoan Cristatella mucedo retains ancestral lophotrochozoan features.

Bryozoans are aquatic colonial suspension-feeders abundant in many marine and freshwater benthic communities. At the same time, the phylum is under studied 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 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 cladesshowing 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.

Peculiarities of Tentacle Innervation of Flustrellidra hispida and Evolution of Lophophore in Bryozoa.

The study of the lophophore organization is of great importance for the reconstruction of lophophorate phylogeny and for understanding the evolutionary transformation in each phylum of Lophophorata. The innervation of the lophophore in ctenostome bryozoan Flustrellidra hispida was studied using immunocytochemistry and confocal laser scanning microscopy. It has been demonstrated that this species has an outer nerve ring giving rise to the tentacle nerves. The outer nerve ring was earlier described in some ctenostomates and cyclostomates, but not as connected with nerves. The discovered feature of lophophore innervation in F. hispida suggests the evolutionary transformation from a hypothetical phoronida-like ancestor lophophore bearing a prominent outer nerve ring with numerous tentacle nerves emanating from it, to the complex bell-shaped lophophore of F. hispida with a well-pronounced outer nervous ring bearing a few tentacle nerves. The next one in this hypothetical row is the lophophore of the other ctenostomates and some cyclostomates with no ring-nerve connection and cheilostomates lophophore with no outer nerve ring at all.

First evidence of virus-like particles in the bacterial symbionts of Bryozoa.

Bacteriophage communities associated with humans and vertebrate animals have been extensively studied, but the data on phages living in invertebrates remain scarce. In fact, they have never been reported for most animal phyla. Our ultrastructural study showed for the first time a variety of virus-like particles (VLPs) and supposed virus-related structures inside symbiotic bacteria in two marine species from the phylum Bryozoa, the cheilostomes Bugula neritina and Paralicornia sinuosa. We also documented the effect of VLPs on bacterial hosts: we explain different bacterial 'ultrastructural types' detected in bryozoan tissues as stages in the gradual destruction of prokaryotic cells caused by viral multiplication during the lytic cycle. We speculate that viruses destroying bacteria regulate symbiont numbers in the bryozoan hosts, a phenomenon known in some insects. We develop two hypotheses explaining exo- and endogenous circulation of the viruses during the life-cycle of B. neritina. Finally, we compare unusual 'sea-urchin'-like structures found in the collapsed bacteria in P. sinuosa with so-called metamorphosis associated contractile structures (MACs) formed in the cells of the marine bacterium Pseudoalteromonas luteoviolacea which are known to trigger larval metamorphosis in a polychaete worm.

Artificial habitats induce plasticity in colonies of the marine bryozoan Schizoporella errata.

Modular organization provides flexibility for colonial animals to deal with variable and unpredictable environmental conditions since each module has specific tasks within the colony, such as feeding, defending or reproducing. Depending on the selecting pressures, sessile organisms may phenotypically adjust the morphology of each module or modify their density, increasing individual fitness. Here we used the marine bryozoan Schizoporella errata (Cheilostomata, Schizoporellidae) to test how the divergent conditions between two artificial habitats, the location inside a marina (IM) and the external wall of the breakwater (BW), affect colony size and the density of the distinct modules. The density of avicularia and ovicells, modules related to defense and reproduction, respectively, did not differ between habitats. However, colonies growing in the turbulent waters of BW were, in general, larger and had higher density of feeding autozooids than those at IM. Reciprocal transplants of bryozoan clones indicated that trait variation is genotype-dependent but varies according to the environmental conditions at the assigned location. The occurrence of larger colonies with more zooids in BW is probably linked to the easier feeding opportunity offered by the small diffusive boundary layer around the colony at this location. Since in colonial polymorphic organisms each module (zooid) performs a specific function, the phenotypic response is not uniform across colonies, affecting only those modules that are susceptible to variations in the main selective pressures. Understanding the importance of colony-level plasticity is relevant to predict how modularity will contribute to organisms to deal with human-induced environmental changes in coastal habitats.

Morphology of ctenostome bryozoans: 1. Arachnidium fibrosum.

The morphology of ctenostome bryozoans remains little investigated with only few species having been subject to more detailed studies. From all the seven main different superfamilies, only few representatives have been studied. The superfamily Arachnidioidea has particularly been neglected concerning detailed morphological and histological details. So far, not a single analysis specifically studied a representative of the family Arachnidiidae. Arachnidium-like forms have, however, often been regarded as potential cheilostome ancestors, the most successful group of bryozoans to date. The lack of any morphological data on this family called for a detailed investigation of one of its representatives. Hence, we analysed the general morphology and histology of Arachnidium fibrosum. Most striking morphological features previously unrecognized are a cardiac constrictor, previously almost unknown in the family, a single pair of apertural muscles consisting of proximal parieto-diaphragmatic and distal parieto-vestibular muscles, six pairs of duplicature bands, a lophophoral anus and retractor muscles attaching to the foregut. Although comparative data are limited, there seem to be two distinct different clades of arachnidiid ctenostomes that are characterized by their aperture and details of gut morphology. Further analysis of additional arachnidioidean species are required to confirm this.

Morphology of ctenostome bryozoans: 2. Haywardozoon pacificum, with implications of the phylogenetic position of the genus.

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.