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Aaron O'Dea and Kimberly García-Méndez introduce the cupuladriids, a group of bryozoans that are atypical for their ability to actively move., Competing Interests: Declaration of interests The authors declare no competing interests., (Published by Elsevier Inc.)

Warming global temperatures have consequences for biological rates. Feeding rates reflect the intake of energy that fuels survival, growth and reproduction. However, temperature can also affect food abundance and quality, as well as feeding behavior, which all affect feeding rate, making it challenging to understand the pathways by which temperature affects the intake of energy. Therefore, we experimentally assessed how clearance rate varied across a thermal gradient in a filter-feeding colonial marine invertebrate (the bryozoan Bugula neritina). We also assessed how temperature affects phytoplankton as a food source, and zooid states within a colony that affect energy budgets and feeding behavior. Clearance rate increased linearly from 18°C to 32°C, a temperature range that the population experiences most of the year. However, temperature increased algal cell size, and decreased the proportion of feeding zooids, suggesting indirect effects of temperature on clearance rates. Temperature increased polypide regression, possibly as a stress response because satiation occurred quicker, or because phytoplankton quality declined. Temperature had a greater effect on clearance rate per feeding zooid than it did per total zooids. Together, these results suggest that the effect of temperature on clearance rate at the colony level is not just the outcome of individual zooids feeding more in direct response to temperature but also emerges from temperature increasing polypide regression and the remaining zooids increasing their feeding rates in response. Our study highlights some of the challenges for understanding why temperature affects feeding rates, especially for understudied, yet ecologically important, marine colonial organisms., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Bryozoan colonies consist of zooids, which can differ in structure and function. Most heteromorphic zooids are unable to feed and autozooids supply them with nutrients. The structure of the tissues providing nutrient transfer is poorly investigated. Here, I present a detailed description of the colonial system of integration (CSI) and communication pores in autozooids and avicularia of the cheilosome bryozoan Terminoflustra membranaceotruncata. The CSI is the nutrient transport and distribution system in the colony. In both autozooids and avicularia it consists of a single cell type, that is, elongated cells, and has a variable branching pattern, except for the presence of a peripheral cord. The general similarity in the CSI structure in avicularia and autozooids is probably due to the interzooidal type of the avicularium. Interzooidal avicularia are likely to consume only a part of the nutrients delivered to them by the CSI, and they transit the rest of the nutrients further. The variability and irregularity of branching pattern of the CSI may be explained by the presence of single communication pores and their varying number. The structure of communication pores is similar regardless of their location (in the transverse or lateral wall) and the type of zooid in contact. Rosette complexes include a cincture cell, a few special cells, and a few limiting cells. Along each zooidal wall, there are communication pores with both unidirectional and bidirectional polarity of special cells. However, the total number of nucleus-containing lobes of special cells is approximately the same on each side of any zooidal wall. Supposing the polarity of special cells reflects the direction of nutrient transport, the pattern of special cells polarity is probably related to the need for bidirectional transport through each zooidal wall. The possibility for such transport is important in large perennial colonies with wide zones of autozooids undergoing polypide degeneration., (© 2024 Wiley Periodicals LLC.)

AbstractThe form of the cyphonautes larva of bryozoans changes little during development. The ciliated band that generates the feeding current increases nearly in proportion to body length, so that the maximum rate of clearing planktonic food from a volume of water becomes increasingly low relative to body protein. This development is unlike the other larvae that produce a feeding current with bands of simple cilia. The cyphonautes' growth rate has therefore been predicted to be unusually low when food is scarce. As predicted, cyphonautes larvae of a species of Membranipora starved at concentrations of food that supported growth of pluteus larvae. Comparisons between the cyphonautes and plutei of a sand dollar were for growth from first feeding to metamorphosis, with a mix of two algal species. Another comparison was for growth of cyphonautes at an advanced stage and plutei of a regular sea urchin at an early stage, with food in seawater at a reduced concentration. The low maximum clearance rate did not prevent rapid growth and development of some cyphonautes from egg through metamorphosis when food was abundant. Twenty-nine days for development to metamorphosis in the laboratory with abundant food was close to Yoshioka's estimate of larval duration from the time lag between adult zooid density and larval abundance in a population in the Southern California Bight. Despite individual variation in growth rates and other physiological and environmental influences, simple measures of larval form predicted the differences in larval performance: scarce food extended larval duration for the cyphonautes more than for plutei.

A bryozoan survey conducted in the Amazon Basin in the vicinities of Manaus and Santarm during the high water season (May, 2018) revealed four new species described here: Fredericella adrianoi n. sp., Plumatella divae n. sp., Plumatella hartikainenae n. sp., and Plumatella spencerjonesae n. sp. Two of these species were encountered only once, suggesting that other undescribed species are likely to occur in the area. Range extensions were determined for two additional species: Plumatella pirassununga and Timwoodiellina natans. In addition, colonies were collected for the first time for two species previously known only by their statoblasts: Plumatella siolii and Plumatella marcusi. Statoblasts of Tapajosella elongata were encountered near Manaus, but the colonies remained elusive. The discovery of new species collected during two expeditions to the Amazon Basin in different seasons and years suggests that further diversity remains undetected in this and other poorly studied regions of the world. With few exceptions, plumatellid colonies described so far from the Amazon Basin are very similar in appearance, with branches wholly attached to the substratum and body walls that are soft, colorless, and transparent.

Metabolism should drive demography by determining the rates of both biological work and resource demand. Long-standing "rules" for how metabolism should covary with demography permeate biology, from predicting the impacts of climate change to managing fisheries. Evidence for these rules is almost exclusively indirect and in the form of among-species comparisons, while direct evidence is exceptionally rare. In a manipulative field experiment on a sessile marine invertebrate, we created experimental populations that varied in population size (density) and metabolic rate, but not body size. We then tested key theoretical predictions regarding relationships between metabolism and demography by parameterizing population models with lifetime performance data from our field experiment. We found that populations with higher metabolisms had greater intrinsic rates of increase and lower carrying capacities, in qualitative accordance with classic theory. We also found important departures from theory-in particular, carrying capacity declined less steeply than predicted, such that energy use at equilibrium increased with metabolic rate, violating the long-standing axiom of energy equivalence. Theory holds that energy equivalence emerges because resource supply is assumed to be independent of metabolic rate. We find this assumption to be violated under real-world conditions, with potentially far-reaching consequences for the management of biological systems., Competing Interests: The authors declare no competing interest.

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., (© 2020 Wiley Periodicals, Inc.)

The reef system off the Amazon River mouth extends from Amapá state to Maranhão state along the Brazilian Equatorial Margin, encompassing more than 10,000 km2 of rhodolith beds and high-relief hard structures on the outer shelf and upper slope. This unique hard bottom mosaic is remarkable for being influenced by the turbid and hyposaline plume from the world's largest river, and also for representing a connectivity corridor between the Caribbean and Brazil. Bryozoans were recently recognized as major reef builders in the Southwestern Atlantic, but their diversity off the Amazon River mouth remained unknown. Here, we report on recent collections obtained from 23 to 120 m depth in Northern Brazil. Sixty-five bryozoan taxa were characterized using scanning electron microscopy, including 57, five and three taxa of Cheilostomatida, Cyclostomatida and Ctenostomatida, respectively. Cribrilaria smitti and three genera (Cranosina, Glabrilaria and Thornelya) are new records for Brazil, and 13 new species are herein described: Antropora cruzeiro n. sp., Cranosina gilbertoi n. sp., Cribrilaria lateralis n. sp., Crisia brasiliensis n. sp., Glabrilaria antoniettae n. sp., Micropora amapaensis n. sp., Parasmittina amazonensis n. sp., Plesiocleidochasma arcuatum n. sp., Poricella bifurcata n. sp., Pourtalesella duoavicularia n. sp., Stephanollona domuspusilla n. sp., Therenia dianae n. sp., and Thornelya atlanticoensis n. sp. Our results highlight the biodiversity significance of the Amazon reefs and the need for more comprehensive sampling to clarify the role of bryozoans in modern turbid-zone reefs and rhodolith beds.

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.

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., (© 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Viable populations of the cheilostome bryozoan Cribrilina mutabilis Ito, Onishi & Dick exist in the NW Pacific (Russian Far East and northern Japan), NE Atlantic (Scandinavia and Scotland), and NW Atlantic (Maine, USA). The first NE and NW Atlantic records are from Norway (2008) and Casco Bay, Maine, USA (2018), respectively, indicating a relatively recent introduction to the region. Mitochondrial COI gene sequences from North Atlantic populations (Sweden, Norway, and Maine) showed two haplotypes differing by one substitution, but differed from two haplotypes from Akkeshi, northern Japan, by 6-8 substitutions. North Atlantic populations differed morphologically from the Akkeshi population in that some zooids formed a suboral projection, and frontal zooids were more common. While C. mutabilis in northern Japan has been found only on natural or artificial eelgrass ( Zostera marina ), across its range it has been found on several species of algae, plastic panels and strips, several species of Zostera , and mollusc shells. Similar frequencies of heteromorphic zooids with differing degree of frontal wall calcification, i.e., R (rib)-, I (intermediate)-, and S (shield)-type zooids, in colonies on eelgrass at comparable times of the season and across populations suggest an innate response to seasonal environmental fluctuations, although zooid frequencies were different on non-eelgrass substrates. The increase in trans-Arctic shipping along the Northern Sea Route in recent decades, and previous documentation of C. mutabilis on ship hulls in the Sea of Japan, indicate a clear mechanism for anthropogenic introduction from the Far East to Europe in recent decades.

Antarctic shallow coastal marine communities were long thought to be isolated from their nearest neighbours by hundreds of kilometres of deep ocean and the Antarctic Circumpolar Current. The discovery of non-native kelp washed up on Antarctic beaches led us to question the permeability of these barriers to species dispersal. According to the literature, over 70 million kelp rafts are afloat in the Southern Ocean at any one time. These living, floating islands can play host to a range of passenger species from both their original coastal location and those picked in the open ocean. Driven by winds, currents and storms towards the coast of the continent, these rafts are often cited as theoretical vectors for the introduction of new species into Antarctica and the sub-Antarctic islands. We found non-native kelps, with a wide range of "hitchhiking" passenger organisms, on an Antarctic beach inside the flooded caldera of an active volcanic island. This is the first evidence of non-native species reaching the Antarctic continent alive on kelp rafts. One passenger species, the bryozoan Membranipora membranacea, is found to be an invasive and ecologically harmful species in some cold-water regions, and this is its first record from Antarctica. The caldera of Deception Island provides considerably milder conditions than the frigid surrounding waters and it could be an ideal location for newly introduced species to become established. These findings may help to explain many of the biogeographic patterns and connections we currently see in the Southern Ocean. However, with the impacts of climate change in the region we may see an increase in the range and number of organisms capable of surviving both the long journey and becoming successfully established.

The evolution of modular colonial animals such as reef corals and bryozoans is enigmatic because of the ability for modules to proliferate asexually as whole colonies reproduce sexually. This reproductive duality creates an evolutionary tension between modules and colonies because selection operates at both levels. To understand how this evolutionary conflict is resolved, we compared the evolutionary potential of module- and colony-level traits in two species of the bryozoan Stylopoma , grown and bred in a common garden experiment. We find quantitatively distinct differences in the evolutionary potential of modular and colony traits. Contrary to solitary organisms, individual traits are not heritable from mother to daughter modules, but colony traits are strongly heritable from parent to offspring colonies. Colony-level evolution therefore dominates because no evolutionary change can accumulate among its modules., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

Based on morphological evidence, Bryozoa together with Phoronida and Brachiopoda are traditionally combined in the group Lophophorata, although this view has been recently challenged by molecular studies. The core of the concept lies in the presence of the lophophore as well as the nature and arrangement of the body cavities. Bryozoa are the least known in this respect. Here, we focused on the fine structure of the body cavity in 12 bryozoan species: 6 gymnolaemates, 3 stenolaemates and 3 phylactolaemates. In gymnolaemates, the complete epithelial lining of the body cavity is restricted to the lophophore, gut walls, and tentacle sheath. By contrast, the cystid walls are composed only of the ectocyst-producing epidermis without a coelothelium, or an underlying extracellular matrix; only the storage cells and cells of the funicular system contact the epidermis. The nature of the main body cavity in gymnolaemates is unique and may be considered as a secondarily modified coelom. In cyclostomes, both the lophophoral and endosaccal cavities are completely lined with coelothelium, while the exosaccal cavity only has the epidermis along the cystid wall. In gymnolaemates, the lophophore and trunk cavities are divided by an incomplete septum and communicate through two pores. In cyclostomes, the septum has a similar location, but no openings. In Phylactolaemata, the body cavity is undivided: the lophophore and trunk coeloms merge at the bases of the lophophore arms, the epistome cavity joins the trunk, and the forked canal opens into the arm coelom. The coelomic lining of the body is complete except for the epistome, lophophoral arms, and the basal portions of the tentacles, where the cells do not interlock perfectly (this design probably facilitates the ammonia excretion). The observed partitioning of the body cavity in bryozoans differs from that in phoronids and brachiopods, and contradicts the Lophophorata concept., (© 2019 Wiley Periodicals, Inc.)

Cheilostome Bryozoa Anoteropora latirostris, a colonial marine invertebrate, constructs its skeleton from calcite and aragonite. This study presents firstly correlated multi-scale electron microscopy, micro-computed tomography, electron backscatter diffraction and NanoSIMS mapping. We show that all primary, coarse-grained platy calcitic lateral walls are covered by fine-grained fibrous aragonite. Vertical lateral walls separating autozooid chambers have aragonite only on their distal side. This type of asymmetric mineralization of lateral walls results from the vertical arrangement of the zooids at the growth margins of the colony and represents a type of biomineralization previously unknown in cheilostome bryozoans. NanoSIMS mapping across the aragonite-calcite interface indicates an organic layer between both mineral phases, likely representing an organic template for biomineralization of aragonite on the calcite layer. Analysis of crystallographic orientations show a moderately strong crystallographic preferred orientation (CPO) for calcite (7.4 times random orientation) and an overall weaker CPO for aragonite (2.4 times random orientation) with a high degree of twinning (45%) of the aragonite grains. The calculated Young's modulus for the CPO map shows a weak mechanical direction perpendicular to the colony's upper surface facilitating this organism's strategy of clonal reproduction by fragmentation along the vertical zooid walls.

Bacteria in corals have been studied in detail in the past decades. However, the biodiversity and bioactivity of fungi in corals are still poorly understood. This study investigated the biodiversity and antifouling activity of fungi in soft corals Cladiella krempfi and Sarcophyton tortuosum from the South China Sea. A high diverse and abundant fungal community was found in the two soft corals. Furthermore, five isolates shared 83-95% similarity with their closest relatives, indicating that they might be novel species in genera Phaeoshaeria and Mucor. In addition, approximately 50% of the representative isolates exhibited distinct antifouling activity. In particular, isolates Fungal sp. SCAU132 and Fungal sp. SCAU133 displayed very strong antifouling activity against Bugula neritina, suggesting they can provide a potential resource for further investigation on isolation of novel antifouling metabolites. To our knowledge, this study is the first report to investigate the biodiversity and antifouling activity of fungi in C. krempfi and S. tortuosum.

Offspring sizes vary within populations but the reasons are unclear. Game-theoretic models predict that selection will maintain offspring-size variation when large offspring are superior competitors (i.e., competition is asymmetric), but small offspring are superior colonizers. Empirical tests are equivocal, however, and typically rely on interspecific comparisons, whereas explicit intraspecific tests are rare. In a field study, we test whether offspring size affects competitive asymmetries using the sessile marine invertebrate, Bugula neritina. Surprisingly, we show that offspring size determines whether interactions are competitive or facilitative-large neighbors strongly facilitated small offspring, but also strongly competed with large offspring. These findings contradict the assumptions of classic theory-that is, large offspring were not superior competitors. Instead, smaller offspring actually benefit from interactions with large offspring-suggesting that asymmetric facilitation, rather than asymmetric competition, operates in our system. We argue that facilitation of small offspring may be more widespread than currently appreciated, and may maintain variation in offspring size via negative frequency-dependent selection. Offspring size theory has classically viewed offspring interactions through the lens of competition alone, yet our results and those of others suggest that theory should accommodate positive interactions in explorations of offspring-size variation., (© 2019 The Author(s). Evolution © 2019 The Society for the Study of Evolution.)

Documented occurrences of fossil taxa are the empirical foundation for understanding large-scale biodiversity changes and evolutionary dynamics in deep time. The fossil record contains vast amounts of understudied taxa. Yet the compilation of huge volumes of data remains a labour-intensive impediment to a more complete understanding of Earth's biodiversity history. Even so, many occurrence records of species and genera in these taxa can be uncovered in the palaeontological literature. Here, we extract observations of fossils and their inferred ages from unstructured text in books and scientific articles using machine-learning approaches. We use Bryozoa, a group of marine invertebrates with a rich fossil record, as a case study. Building on recent advances in computational linguistics, we develop a pipeline to recognize taxonomic names and geologic time intervals in published literature and use supervised learning to machine-read whether the species in question occurred in a given age interval. Intermediate machine error rates appear comparable to human error rates in a simple trial, and resulting genus richness curves capture the main features of published fossil diversity studies of bryozoans. We believe our automated pipeline, that greatly reduced the time required to compile our dataset, can help others compile similar data for other taxa.

Bryozoans are the major component of marine macro-fouling communities. In the study, the relations between bryozoan species and environmental variables were investigated at seven stations along the Aegean coast in August and December 2015. Constant bryozoan species in both sampling periods were Bugula neritina, Amathia verticillata, Shizoporella errata, Cryptosula pallasiana and Celleporaria brunnea. Their relationship with physico-chemical variables (Temperature, salinity, pH, dissolved oxygen, ammonium, nitrite and nitrate nitrogen, orthophosphate phosphorous, total phosphate, chlorophyll-a) were analysed by means of logistic regression analysis. The result showed that temperature with B. neritina; NH 4 -N, oPO 4 -P and TPO 4 -P with A verticillata; dissolved oxygen concentrations with S. errata and C. brunnea were positively related (p < 0.05)., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

The evolution of parental care is a central field in many ecological and evolutionary studies, but integral approaches encompassing various life-history traits are not common. Else, the structure, development and functioning of the placental analogues in invertebrates are poorly understood. Here, we describe the life-history, sexual colony dynamics, oogenesis, fertilization and brooding in the boreal-Arctic cheilostome bryozoan Celleporella hyalina. This placental brooder incubates its progeny in calcified protective chambers (ovicells) formed by polymorphic sexual zooids. We conducted a detailed ultrastructural study of the ovary and oogenesis, and provide evidence of both auto- and heterosynthetic mechanisms of vitellogenesis. We detected sperm inside the early oocyte and within funicular strands, and discuss possible variants of fertilization. We also detail the development and functioning of the placental analogue (embryophore) in the various stages of embryonic incubation as well as embryonic histotrophic nourishment. In contrast to all known cheilostome placentas, the main part of embryophore of C. hyalina is not a single cell layer. Rather, it is a massive "nutritive tissue" whose basal part is associated with funicular strands presumably providing transport function. C. hyalina shows a mixture of reproductive traits with macrolecithal oogenesis and well-developed placenta. These features give it an intermediate position in the continuum of variation of matrotrophic provisioning between lecithotrophic and placentotrophic cheilostome brooders. The structural and developmental differences revealed in the placental analogue of C. hyalina, together with its position on the bryozoan molecular tree, point to the independent origin of placentation in the family Hippothoidae., (© 2019 The Authors. Journal of Morphology published by Wiley Periodicals, Inc.)

The increasing number of anthropogenic areas in the marine environment results in significant impact to adjacent ecosystems. In fact, the presence of marinas modifies the original environmental conditions and ends up disturbing the faunal community. However, despite the essential role displayed by the macrofauna on marinas' fouling biota, certain taxa such as polychaetes have been poorly studied. The present study provides the first spatial characterization of the epibiont polychaete fauna associated with the bryozoan Bugula neritina in marinas along the Iberian Peninsula and the north of Morocco. A total of 32 polychaete species were identified, with Syllidae being the most diverse family. Furthermore, the environmental factors involved in the occurrence and abundance of the dominant species Salvatoria clavata were also analyzed by Generalized Linear Models; results showed that the highest predicted values of S. clavata abundance appeared at marinas with high levels of nutrient enrichment and of heavy metals concentration., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The life histories of modular organisms are complicated, where selection and optimization can occur at both organismal and modular levels. At a modular level, growth, reproduction and death can occur in one module, independently of others. Across modular groups, there are no formal investigations of selection on module longevity. We used two field experiments to test whether selection acts on module longevity in a sessile marine invertebrate and whether selection varies across successional gradients and resource regimes. We found that selection does act on module longevity and that the strength of selection varies with environmental conditions. In environments where interspecific competition is high, selection favours colonies with longer zooid (module) longevity for colonies that initially received high levels of maternal investment. In environments where food availability is high and flow rate is low, selection also favours colonies with longer zooid longevity. These patterns of selection provide partial support for module longevity theory developed for plants. Nevertheless, that selection on module longevity is so context-dependent suggests that variation in module longevity is likely to be maintained in this system., (© 2018 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2018 European Society For Evolutionary Biology.)

Background: Cyclostome bryozoans are an ancient group of marine colonial suspension-feeders comprising approximately 700 extant species. Previous morphological studies are mainly restricted to skeletal characters whereas data on soft tissues obtained by state-of-the-art methods are still lacking. In order to contribute to issues related to cyclostome ground pattern reconstruction, we analyzed the morphology of the neuromuscular system Cinctipora elegans by means of immunocytochemical staining, confocal laser scanning microscopy, histological sections and microCT imaging., Results: Polypides of C. elegans are located in elongated tubular skeletal cystids. Distally, the orifice leads into a prominent vestibulum which is lined by an epithelium that joins an almost complete perimetrical attachment organ, both containing radially arranged neurite bundles and muscles. Centrally, the prominent atrial sphincter separates the vestibulum from the atrium. The latter is enclosed by the tentacle sheath which contains few longitudinal muscle fibers and two principal neurite bundles. These emerge from the cerebral ganglion, which is located at the lophophoral base. Lateral ganglia are located next to the cerebral ganglion from which the visceral neurite bundles emerge that extend proximally towards the foregut. There are four tentacle neurite bundles that emerge from the ganglia and the circum-oral nerve ring, which encompasses the pharynx. The tentacles possess two striated longitudinal muscles. Short buccal dilatators are situated at the lophophoral base and short muscular sets are present at the abfrontal and frontal side of the tentacle base. The pharynx is myoepithelial and triradiate in cross-section. Oocytes are found inside the pharyngeal myoepithelium. The digestive tract contains dense circular musculature and few longitudinal muscles. The membranous sac contains regular, thin, circular and diagonal muscles and neurites in its epithelial lining., Conclusions: The general structure of the neuro-muscular system is more reminiscent of the condition found in Gymnolaemata rather than Phylactolaemata, which supports a close relationship between Cyclostomata and Gymnolaemata. Several characters of C. elegans such as the lateral ganglia or loss of the cardia are probably apomorphic for this species. For the first time, oocytes that surprisingly develop in the pharyngeal wall are reported for this species.

The bryozoan Bugula neritina is a cosmopolitan marine fouling species that causes major fouling problems in sub-tropical waters. Settlement of B. neritina larvae can be triggered without an obvious external cue. Here, the negative regulatory role of nitric oxide (NO) during larval settlement of B. neritina was demonstrated to be mediated by cyclic guanosine monophosphate (cGMP). Although the regulatory role of the NO-p38 MAPK signaling axis in larval settlement was not evident, inhibition of nitric oxide synthase (NOS) led to the deactivation of p38 MAPK. Exclusive localization of NO and NO signaling components in sensory-related organs of the larvae is consistent with its signal transduction function in metamorphosis. Overall, this study provides new insights into the regulatory roles of the NO-p38MAPK/cGMP pathway in B. neritina settlement.

Multivariate analyses were used to investigate the influence of selected surface properties (Owens-Wendt surface energy and its dispersive and polar components, static water contact angle, conceptual sign of the surface charge, zeta potentials) on the attachment patterns of five biofouling organisms (Amphibalanus amphitrite, Amphibalanus improvisus, Bugula neritina, Ulva linza, and Navicula incerta) to better understand what surface properties drive attachment across multiple fouling organisms. A library of ten xerogel coatings and a glass standard provided a range of values for the selected surface properties to compare to biofouling attachment patterns. Results from the surface characterization and biological assays were analyzed separately and in combination using multivariate statistical methods. Principal coordinate analysis of the surface property characterization and the biological assays resulted in different groupings of the xerogel coatings. In particular, the biofouling organisms were able to distinguish four coatings that were not distinguishable by the surface properties of this study. The authors used canonical analysis of principal coordinates (CAP) to identify surface properties governing attachment across all five biofouling species. The CAP pointed to surface energy and surface charge as important drivers of patterns in biological attachment, but also suggested that differentiation of the surfaces was influenced to a comparable or greater extent by the dispersive component of surface energy.

Phenotypic plasticity has the potential to allow organisms to respond rapidly to global environmental change, but the range and effectiveness of these responses are poorly understood across taxa and growth strategies. Colonial organisms might be particularly resilient to environmental stressors, as organizational modularity and successive asexual generations can allow for distinctively flexible responses in the aggregate form. We performed laboratory experiments to examine the effects of increasing dissolved carbon dioxide (CO 2 ) (i.e. ocean acidification) on the colonial bryozoan Celleporella cornuta sampled from two source populations within a coastal upwelling region of the northern California coast. Bryozoan colonies were remarkably plastic under these CO 2 treatments. Colonies raised under high CO 2 grew more quickly, investing less in reproduction and producing lighter skeletons when compared with genetically identical clones raised under current surface atmosphere CO 2 values. Bryozoans held under high CO 2 conditions also changed the Mg/Ca ratio of skeletal calcite and increased the expression of organic coverings in new growth, which may serve as protection against acidified water. We also observed strong differences between source populations in reproductive investment and organic covering reaction norms, consistent with adaptive responses to persistent spatial variation in local oceanographic conditions. Our results demonstrate that phenotypic plasticity and energetic trade-offs can mediate biological responses to global environmental change, and highlight the broad range of strategies available to colonial organisms., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

Few studies to date have investigated the effects of ocean acidification on non-reef forming marine invertebrates with non-feeding larvae. Here, we exposed adults of the bryozoan Bugula neritina and their larvae to lowered pH. We monitored spawning, larval swimming, settlement, and post-settlement individual sizes at two pHs (7.9 vs. 7.6) and settlement dynamics alone over a broader pH range (8.0 down to 6.5). Our results show that spawning was not affected by adult exposure (48h at pH7.6), larvae swam 32% faster and the newly-settled individuals grew significantly larger (5%) at pH7.6 than in the control. Although larvae required more time to settle when pH was lowered, reduced pH was not lethal, even down to pH6.5. Overall, this fouling species appeared to be robust to acidification, and yet, indirect effects such as prolonging the pelagic larval duration could increase predation risk, and might negatively impact population dynamics., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Background: Phylactolaemata is commonly regarded the earliest branch within Bryozoa and thus the sister group to the other bryozoan taxa, Cyclostomata and Gymnolaemata. Therefore, the taxon is important for the reconstruction of the bryozoan morphological ground pattern. In this study the myoanatomy of Pectinatella magnifica, Cristatella mucedo and Hyalinella punctata was analysed by means of histology, f-actin staining and confocal laser-scanning microscopy in order to fill gaps in knowledge concerning the myoanatomy of Phylactolaemata., Results: The retractor muscles and muscles of the aperture, gut, body wall, tentacle sheath, lophophore constitute the most prominent muscular subsets in these species. The lophophore shows longitudinal muscle bands in the tentacles, lophophoral arm muscles, epistome musculature and hitherto undescribed muscles of the ring canal. In general the muscular system of the three species is very similar with differences mainly in the body wall, tentacle sheath and epistome. The body wall contains an orthogonal grid of musculature. The epistome exhibits either a muscular meshwork in the epistomal wall or muscle fibers traversing the epistomal cavity. The whole tentacle sheath possesses a regular mesh of muscles in Pectinatella and Cristatella, whereas circular muscles are limited to the tentacle sheath base in Hyalinella., Conclusion: This study is the first to describe muscles of the ring canal and contributes to reconstructing muscular features for the last common ancestor of all bryozoans. The data available suggest that two longitudinal muscle bands in the tentacles, as well as retractor muscles and longitudinal and circular muscles in the tentacle sheath, were present in the last common bryozoan ancestor. Comparisons among bryozoans shows that several apomorphies are present in the myoanatomy of each class- level taxon such as the epistomal musculature and musculature of the lophophoral arms in phylactolaemates, annular muscles in cyclostomes and parietal muscles in gymnolaemates.

In this study we produce a standardised dataset for benthic macrofauna and sediments through integration of data (33,198 samples) from 777 grab surveys. The resulting dataset is used to identify spatial and temporal patterns in faunal distribution around the UK, and the role of sediment composition and other explanatory variables in determining such patterns. We show how insight into natural variability afforded by the dataset can be used to improve the sustainability of activities which affect sediment composition, by identifying conditions which should remain favourable for faunal recolonisation. Other big data applications and uses of the dataset are discussed.

Bryozoans are among the most abundant and diverse members of the Antarctic benthos, however the role of bioactive metabolites in ecological interactions has been scarcely studied. To extend our knowledge about the chemical ecology of Antarctic bryozoans, crude ether extracts (EE) and butanol extracts (BE) obtained from two Antarctic common species (Cornucopina pectogemma and Nematoflustra flagellata), were tested for antibacterial and repellent activities. The extracts were screened for quorum quenching and antibacterial activities against four Antarctic bacterial strains (Bacillus aquimaris, Micrococcus sp., Oceanobacillus sp. and Paracoccus sp.). The Antarctic amphipod Cheirimedon femoratus and the sea star Odontaster validus were selected as sympatric predators to perform anti-predatory and substrate preference assays. No quorum quenching activity was detected in any of the extracts, while all EE exhibited growth inhibition towards at least one bacterium strain. Although the species were not repellent against the sea star, they caused repellence to the amphipods in both extracts, suggesting that defence activities against predation derive from both lipophilic and hydrophilic metabolites. In the substrate preference assays, one EE and one BE deriving from different specimens of the species C. pectogemma were active. This study reveals intraspecific variability of chemical defences and supports the fact that chemically mediated interactions are common in Antarctic bryozoans as means of protection against fouling and predation., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Background: The nervous system in brachiopods has seldom been studied with modern methods. An understanding of lophophore innervation in adult brachiopods is useful for comparing the innervation of the same lophophore type among different brachiopods and can also help answer questions about the monophyly of the lophophorates. Although some brachiopods are studied with modern methods, rhynchonelliform brachiopods still require investigation. The current study used transmission electron microscopy, immunocytochemistry, and confocal laser scanning microscopy to investigate the nerve system of the lophophore and tentacles in the rhynchonelliform Hemithiris psittacea., Results: Four longitudinal nerves pass along each brachium of the lophophore: the main, accessory, second accessory, and lower. The main brachial nerve extends at the base of the dorsal side of the brachial fold and gives rise to the cross nerves, passing through the extracellular matrix to the tentacles. Cross nerves skirt the accessory brachial nerve, branch, and penetrate into adjacent outer and inner tentacles, where they are referred to as the frontal tentacular nerves. The second accessory nerve passes along the base of the inner tentacles. This nerve consists of Ʊ-like parts, which repetitively skirt the frontal and lateral sides of the inner tentacle and the frontal sides of the outer tentacles. The second accessory nerve gives rise to the latero-frontal nerves of the inner and outer tentacles. The abfrontal nerves of the inner tentacles also originate from the second accessory nerve, whereas the abfrontal nerves of the outer tentacles originate from the lower brachial nerve. The lower brachial nerve extends along the outer side of the lophophore brachia and gives rise to the intertentacular nerves, which form a T-like branch and penetrate the adjacent outer tentacles where they are referred to as abfrontal nerves. The paired outer radial nerves start from the lower brachial nerve, extend into the second accessory nerve, and give rise to the lateroabfrontal tentacular nerves of the outer tentacles., Conclusions: The innervation of the lophophore in the rhynchonelliform Hemithiris psittacea differs from that in the inarticulate Lingula anatina in several ways. The accessory brachial nerve does not participate in the innervation of the tentacles in H. psittacea as it does in L. anatina. The second accessory nerve is present in H. psittacea but not in L. anatina. There are six tentacular nerves in the outer tentacles of H. psittacea but only four in all other brachiopods studied to date. The reduced contribution of the accessory brachial nerve to tentacle innervation may reflect the general pattern of reduction of the inner lophophoral nerve in both phoronids and brachiopods. Bryozoan lophophores, in contrast, have a weakened outer nerve and a strengthened inner nerve. Our results suggest that the ancestral lophophore of all lophophorates had a simple shape but many nerve elements.

Stressors associated with global change will be experienced simultaneously and may act synergistically, so attempts to estimate the capacity of marine systems to cope with global change requires a multi-stressor approach. Because recent evidence suggests that stressor effects can be context-dependent, estimates of how stressors are experienced in ecologically realistic settings will be particularly valuable. To enhance our understanding of the interplay between environmental effects and the impact of multiple stressors from both natural and anthropogenic sources, we conducted a field experiment. We explored the impact of multiple, functionally varied stressors from both natural and anthropogenic sources experienced during early life history in a common sessile marine invertebrate, Bugula neritina. Natural spatial environmental variation induced differences in conspecific densities, allowing us to test for density-driven context-dependence of stressor effects. We indeed found density-dependent effects. Under high conspecific density, individual survival increased, which offset part of the negative effects of experiencing stressors. Experiencing multiple stressors early in life history translated to a decreased survival in the field, albeit the effects were not as drastic as we expected: our results are congruent with antagonistic stressor effects. We speculate that when individual stressors are more subtle, stressor synergies become less common.

While the direct physical impact on seabed biota is well understood, no studies have defined thresholds to inform an ecosystem-based approach to managing fishing impacts. We addressed this knowledge gap using a large-scale experiment that created a controlled gradient of fishing intensity and assessed the immediate impacts and short-term recovery. We observed a mosaic of taxon-specific responses at various thresholds. The lowest threshold of significant lasting impact occurred between 1 and 3 times fished and elicited a decrease in abundance of 39 to 70% for some sessile epifaunal organisms (cnidarians, bryozoans). This contrasted with significant increases in abundance and/or biomass of scavenging species (epifaunal echinoderms, infaunal crustaceans) by two to four-fold in areas fished twice and more. In spite of these significant specific responses, the benthic community structure, biomass and abundance at the population level appeared resilient to fishing. Overall, natural temporal variation in community metrics exceeded the effects of fishing in this highly dynamic study site, suggesting that an acute level of disturbance (fished over six times) would match the level of natural variation. We discuss the implications of our findings for natural resources management with respect to context-specific human disturbance and provide guidance for best fishing practices.

Colonial hosts offer unique opportunities for exploitation by endoparasites resulting from extensive clonal propagation, but these interactions are poorly understood. The freshwater bryozoan, Fredericella sultana, and the myxozoan, Tetracapsuloides bryosalmonae, present an appropriate model system for examining such interactions. F. sultana propagates mainly asexually, through colony fragmentation and dormant propagules (statoblasts). Our study examines how T. bryosalmonae exploits the multiple transmission routes offered by the propagation of F. sultana, evaluates the effects of such transmission on its bryozoan host, and tests the hypothesis that poor host condition provokes T. bryosalmonae to bail out of a resource that may soon be unsustainable, demonstrating terminal investment. We show that infections are present in substantial proportions of colony fragments and statoblasts over space and time and that moderate infection levels promote statoblast hatching and hence effective fecundity. We also found evidence for terminal investment, with host starvation inducing the development of transmission stages. Our results contribute to a growing picture that interactions of T. bryosalmonae and F. sultana are generally characterized by parasite persistence, facilitated by multiple transmission pathways and host condition-dependent developmental cycling, and host tolerance, promoted by effective fecundity effects and an inherent capacity for renewed growth and clonal replication.

Population density affects individual performance, though its effects are often mixed. For sessile species, increases in population density typically reduce performance. Still, cases of positive density-dependence do occur in sessile systems and demand explanation. The stress gradient hypothesis (SGH) predicts that under stressful conditions, positive effects of facilitation may outweigh the negative effects of competition. While some elements of the SGH are well studied, its potential to explain intraspecific facilitation has received little attention. Further, there have been questions regarding whether the SGH holds if the stressor is a resource. Most studies of interactions between the environment and intraspecific facilitation have relied on natural environmental gradients; manipulative studies are much rarer. To test the effects of intraspecific density and resources, we manipulated resource availability over natural population densities for the marine bryozoan Watersipora subtorquata. We found negative effects of density on colony performance in low resource environments, but mainly positive density-dependence in high resource environments. By adding resources, competition effects were reduced and the positive effects of facilitation were revealed. Our results suggest that resource availability mediates the relative strength of competition and facilitation in our system. We also suggest that intraspecific facilitation is more common than may be appreciated and that environmental variation may mediate the balance between negative and positive density-dependence., (© 2017 The Authors. Journal of Animal Ecology © 2017 British Ecological Society.)

There are a number of remote archipelagos distributed between 45 and 60 °S. The biota of these islands provide useful information to describe and understand patterns in biodiversity and biogeography as well as potential impacts of climate change on marine ecosystems. They are in key locations either side of the Polar Front but also have limited influence from human activities. Here we investigate one taxon, bryozoans, on South Atlantic shelf habitats of the Falkland (FI) and the sub-Antarctic island of South Georgia (SG). We present new data on spatial distribution in these islands, as well as an analysis of the bryozoological similarities between these and neighbouring regions. A total of 85 species of cheilostome bryozoans (351 samples) were found, belonging to 33 genera, including 18 potentially new genera and 23 new species. Remarkably 65% and 41% of species were reported for the first time at FI and SG, respectively. The highest and the lowest value of species richness and species/genus ratio were found at East (EFI) and West Falkland (WFI), respectively, likely showing a tendency for stronger intrageneric competition. New data from this study were jointly analysed with data from the literature and existing databases, revealing new bathymetric ranges in 32 species. The biogeographic affinities of the bryozoans found give further evidence of the hypothesis of sequential separation of Gondwana and support the changing concept that although the Polar Front acts as a circumpolar biogeographic barrier it is not as impermeable as originally thought. Potential dispersal mechanisms are also discussed., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Marine invertebrates with skeletons made of high-magnesium calcite may be especially susceptible to ocean acidification (OA) due to the elevated solubility of this form of calcium carbonate. However, skeletal composition can vary plastically within some species, and it is largely unknown how concurrent changes in multiple oceanographic parameters will interact to affect skeletal mineralogy, growth and vulnerability to future OA. We explored these interactive effects by culturing genetic clones of the bryozoan Jellyella tuberculata (formerly Membranipora tuberculata ) under factorial combinations of dissolved carbon dioxide (CO 2 ), temperature and food concentrations. High CO 2 and cold temperature induced degeneration of zooids in colonies. However, colonies still maintained high growth efficiencies under these adverse conditions, indicating a compensatory trade-off whereby colonies degenerate more zooids under stress, redirecting energy to the growth and maintenance of new zooids. Low-food concentration and elevated temperatures also had interactive effects on skeletal mineralogy, resulting in skeletal calcite with higher concentrations of magnesium, which readily dissolved under high CO 2 For taxa that weakly regulate skeletal magnesium concentration, skeletal dissolution may be a more widespread phenomenon than is currently documented and is a growing concern as oceans continue to warm and acidify., (© 2017 The Author(s).)

Many aquatic animals grow into colonies of repeated, genetically identical, modules (zooids). Zooid interconnections enable colonies to behave as integrated functional units, while plastic responses to environmental changes may affect individual zooids. Plasticity includes the variable partitioning of resources to sexual reproduction, colony growth and maintenance. Maintenance often involves regeneration, which is also a routine part of the life history in some organisms, such as bryozoans. Here we investigate changes in regenerative capacity in the encrusting bryozoan Cryptosula pallasiana when cultured at different seawater pCO 2 levels. The proportion of active zooids showing polypide regeneration was highest at current oceanic pH (8.1), but decreased progressively as pH declined below that value, reaching a six-fold reduction at pH 7.0. The zone of budding of new zooids at the colony periphery declined in size below pH 7.7. Under elevated pCO 2 conditions, already experienced sporadically in coastal areas, skeletal corrosion was accompanied by the proportional reallocation of resources from polypide regeneration in old zooids to the budding of new zooids at the edge of the colony. Thus, future ocean acidification can affect colonial organisms by changing how they allocate resources, with potentially profound impacts on life-history patterns and ecological interactions., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Across a wide range of taxa, larger mothers produce larger offspring. Theory assumes that larger, more fecund mothers create higher local densities of siblings, and so larger mothers produce larger offspring to offset sibling competition. This assumption has been debated for over 30 yr, but direct empirical tests are surprisingly rare. Here, we test two key assumptions of classic theories that predict sibling competition drives maternal-size-offspring-size (MSOS) correlations: (1) independent effects of offspring size and sibling density on offspring performance or (2) as a product of an interaction between these two factors. To simultaneously test these alternative assumptions, we manipulate offspring size and sibling density in the marine invertebrate, Bugula neritina, and monitor offspring performance in the field. We found that, depending on the fitness metric being considered, offspring size and sibling density can either independently or interactively affect offspring performance. Yet sibling density did not affect offspring performance in the ways that classic theories assume. Given our results, it is unlikely that sibling competition drives the positive MSOS correlation observed in this species. Empirical support for these classic theories remains lacking, suggesting alternative explanations are necessary., (© 2016 by the Ecological Society of America.)

Variation in life-history traits is ubiquitous, even though genetic variation is thought to be depleted by selection. One potential mechanism for the maintenance of trait variation is spatially variable selection. We explored spatial variation in selection in the field for a colonial marine invertebrate that shows phenotypic differences across a depth gradient of only 3 m. Our analysis included life-history traits relating to module size, colony growth, and phenology. Directional selection on colony growth varied in strength across depths, while module size was under directional selection at one depth but not the other. Differences in selection may explain some of the observed phenotypic differentiation among depths for one trait but not another: instead, selection should actually erode the differences observed for this trait. Our results suggest selection is not acting alone to maintain trait variation within and across environments in this system., (© 2016 The Author(s). Evolution © 2016 The Society for the Study of Evolution.)

The bryozoan Bugula neritina has a biphasic life cycle that consists of a planktonic larval stage and a sessile juvenile/adult stage. The transition between these two stages is crucial for the development and recruitment of B. neritina. Metamorphosis in B. neritina is mediated by both the nervous system and the release of developmental signals. However, no research has been conducted to investigate the expression of neuropeptides (NP)/peptide hormones in B. neritina larvae. Here, we report a comprehensive study of the NP/peptide hormones in the marine bryozoan B. neritina based on in silico identification methods. We recovered 22 transcripts encompassing 11 NP/peptide hormone precursor transcript sequences. The transcript sequences of the 11 isolated NP precursors were validated by cDNA cloning using gene-specific primers. We also examined the expression of three peptide hormone precursor transcripts (BnFDSIG, BnILP1, BnGPB) in the coronate larvae of B. neritina, demonstrating their distinct expression patterns in the larvae. Overall, our findings serve as an important foundation for subsequent investigations of the peptidergic control of bryozoan larval behavior and settlement.

In marine systems, oxygen availability varies at small temporal and spatial scales, such that current oxygen levels may not reflect conditions of the past. Different studies have shown that marine invertebrate larvae can select settlement sites based on local oxygen levels and oxygenation history of the biofilm, but no study has examined the interaction of both. The influence of normoxic and hypoxic water and oxygenation history of biofilms on pre-settlement behavior and settlement of the bryozoan Bugula neritina was tested. Larvae used cues in a hierarchical way: the oxygen levels in the water prime larvae to respond, the response to different biofilms is contingent on oxygen levels in the water. When oxygen levels varied throughout biofilm formation, larvae responded differently depending on the history of the biofilm. It appears that B. neritina larvae integrate cues about current and historical oxygen levels to select the appropriate microhabitat and maximize their fitness.

Metabolic rate reflects the 'pace of life' in every organism. Metabolic rate is related to an organism's capacity for essential maintenance, growth and reproduction-all of which interact to affect fitness. Although thousands of measurements of metabolic rate have been made, the microevolutionary forces that shape metabolic rate remain poorly resolved. The relationship between metabolic rate and components of fitness are often inconsistent, possibly because these fitness components incompletely map to actual fitness and often negatively covary with each other. Here we measure metabolic rate across ontogeny and monitor its effects on actual fitness (lifetime reproductive output) for a marine bryozoan in the field. We also measure key components of fitness throughout the entire life history including growth rate, longevity and age at the onset of reproduction. We found that correlational selection favours individuals with higher metabolic rates in one stage and lower metabolic rates in the other-individuals with similar metabolic rates in each developmental stage displayed the lowest fitness. Furthermore, individuals with the lowest metabolic rates lived for longer and reproduced more, but they also grew more slowly and took longer to reproduce initially. That metabolic rate is related to the pace of the life history in nature has long been suggested by macroevolutionary patterns but this study reveals the microevolutionary processes that probably generated these patterns., (© 2016 The Author(s).)

Offshore structures provide habitat that could facilitate species range expansions and the introduction of non-native species into new geographic areas. Surveys of assemblages of seven offshore oil and gas platforms in the Santa Barbara Channel revealed a change in distribution of the non-native sessile invertebrate Watersipora subtorquata, a bryozoan with a planktonic larval duration (PLD) of 24 hours or less, from one platform in 2001 to four platforms in 2013. We use a three-dimensional biophysical model to assess whether larval dispersal via currents from harbors to platforms and among platforms is a plausible mechanism to explain the change in distribution of Watersipora and to predict potential spread to other platforms in the future. Hull fouling is another possible mechanism to explain the change in distribution of Watersipora. We find that larval dispersal via currents could account for the increase in distribution of Watersipora from one to four platforms and that Watersipora is unlikely to spread from these four platforms to additional platforms through larval dispersal. Our results also suggest that larvae with PLDs of 24 hours or less released from offshore platforms can attain much greater dispersal distances than larvae with PLDs of 24 hours or less released from nearshore habitat. We hypothesize that the enhanced dispersal distance of larvae released from offshore platforms is driven by a combination of the offshore hydrodynamic environment, larval behavior, and larval release above the seafloor.

The Southern Ocean archipelago, the South Orkney Islands (SOI), became the world's first entirely high seas marine protected area (MPA) in 2010. The SOI continental shelf (~44 000 km(2) ), was less than half covered by grounded ice sheet during glaciations, is biologically rich and a key area of both sea surface warming and sea-ice losses. Little was known of the carbon cycle there, but recent work showed it was a very important site of carbon immobilization (net annual carbon accumulation) by benthos, one of the few demonstrable negative feedbacks to climate change. Carbon immobilization by SOI bryozoans was higher, per species, unit area and ice-free day, than anywhere-else polar. Here, we investigate why carbon immobilization has been so high at SOI, and whether this is due to high density, longevity or high annual production in six study species of bryozoans (benthic suspension feeders). We compared benthic carbon immobilization across major regions around West Antarctica with sea-ice and primary production, from remotely sensed and directly sampled sources. Lowest carbon immobilization was at the northernmost study regions (South Georgia) and southernmost Amundsen Sea. However, data standardized for age and density showed that only SOI was anomalous (high). High immobilization at SOI was due to very high annual production of bryozoans (rather than high densities or longevity), which were 2x, 3x and 5x higher than on the Bellingshausen, South Georgia and Amundsen shelves, respectively. We found that carbon immobilization correlated to the duration (but not peak or integrated biomass) of phytoplankton blooms, both in directly sampled, local scale data and across regions using remote-sensed data. The long bloom at SOI seems to drive considerable carbon immobilization, but sea-ice losses across West Antarctica mean that significant carbon sinks and negative feedbacks to climate change could also develop in the Bellingshausen and Amundsen seas., (© 2015 John Wiley & Sons Ltd.)

Bio-invasions depend on the number and frequency of invaders arriving in new habitats. Yet, as is often the case, it is not only quantity that counts, but also quality. The process of dispersal can change disperser quality and establishment success. Invasions are a form of extra-range dispersal, so that invaders often experience changes in quality through dispersal. To study effects of dispersal on invader quality, and its interactions with quantity on invasion success, we manipulated both in a field experiment using an invasive marine invertebrate. Establishment success increased with the number of individuals arriving in a new habitat. Prolonged larval durations--our manipulation of prolonged dispersal--decreased individual quality and establishment success. Groups of invaders with prolonged larval durations contributed only a third of the offspring relative to invaders that settled immediately. We also found an interaction between the quality and quantity of invaders on individual growth: only within high-quality cohorts did individuals experience density-dependent effects on growth. Our findings highlight that dispersal not only affects the quantity of invaders arriving in a new habitat but also their quality, and both mediate establishment success.

Within species, larger offspring typically outperform smaller offspring. While the relationship between offspring size and performance is ubiquitous, the cause of this relationship remains elusive. By linking metabolic and life-history theory, we provide a general explanation for why larger offspring perform better than smaller offspring. Using high-throughput respirometry arrays, we link metabolic rate to offspring size in two species of marine bryozoan. We found that metabolism scales allometrically with offspring size in both species: while larger offspring use absolutely more energy than smaller offspring, larger offspring use proportionally less of their maternally derived energy throughout the dependent, non-feeding phase. The increased metabolic efficiency of larger offspring while dependent on maternal investment may explain offspring size effects-larger offspring reach nutritional independence (feed for themselves) with a higher proportion of energy relative to structure than smaller offspring. These findings offer a potentially universal explanation for why larger offspring tend to perform better than smaller offspring but studies on other taxa are needed., (© 2015 The Author(s).)

Ocean acidification is receiving increasing attention because of its potential to affect marine ecosystems. Rare CO2 vents offer a unique opportunity to investigate the response of benthic ecosystems to acidification. However, the benthic habitats investigated so far are mainly found at very shallow water (less than or equal to 5 m depth) and therefore are not representative of the broad range of continental shelf habitats. Here, we show that a decrease from pH 8.1 to 7.9 observed in a CO2 vent system at 40 m depth leads to a dramatic shift in highly diverse and structurally complex habitats. Forests of the kelp Laminaria rodriguezii usually found at larger depths (greater than 65 m) replace the otherwise dominant habitats (i.e. coralligenous outcrops and rhodolith beds), which are mainly characterized by calcifying organisms. Only the aragonite-calcifying algae are able to survive in acidified waters, while high-magnesium-calcite organisms are almost completely absent. Although a long-term survey of the venting area would be necessary to fully understand the effects of the variability of pH and other carbonate parameters over the structure and functioning of the investigated mesophotic habitats, our results suggest that in addition of significant changes at species level, moderate ocean acidification may entail major shifts in the distribution and dominance of key benthic ecosystems at regional scale, which could have broad ecological and socio-economic implications., (© 2015 The Author(s).)

Many animal phyla have the physiological ability to produce biomineralized skeletons with functional roles that have been shaped by natural selection for more than 500 million years. Among these are bryozoans, a moderately diverse phylum of aquatic invertebrates with a rich fossil record and importance today as bioconstructors in some shallow-water marine habitats. Biomineralizational patterns and, especially, processes are poorly understood in bryozoans but are conventionally believed to be similar to those of the related lophotrochozoan phyla Brachiopoda and Mollusca. However, bryozoan skeletons are more intricate than those of these two phyla. Calcareous skeletons have been acquired independently in two bryozoan clades - Stenolaemata in the Ordovician and Cheilostomata in the Jurassic - providing an evolutionary replicate. This review aims to highlight the importance of biomineralization in bryozoans and focuses on their skeletal ultrastructures, mineralogy and chemistry, the roles of organic components, the evolutionary history of bimineralization in bryozoans with respect to changes in seawater chemistry, and the impact of contemporary global changes, especially ocean acidification, on bryozoan skeletons. Bryozoan skeletons are constructed from three different wall types (exterior, interior and compound) differing in the presence/absence and location of organic cuticular layers. Skeletal ultrastructures can be classified into wall-parallel (i.e. laminated) and wall-perpendicular (i.e. prismatic) fabrics, the latter apparently found in only one of the two biomineralizing clades (Cheilostomata), which is also the only clade to biomineralize aragonite. A plethora of ultrastructural fabrics can be recognized and most occur in combination with other fabrics to constitute a fabric suite. The proportion of aragonitic and bimineralic bryozoans, as well as the Mg content of bryozoan skeletons, show a latitudinal increase into the warmer waters of the tropics. Responses of bryozoan mineralogy and skeletal thickness to oscillations between calcite and aragonite seas through geological time are equivocal. Field and laboratory studies of living bryozoans have shown that predicted future changes in pH (ocean acidification) combined with global warming are likely to have detrimental effects on calcification, growth rate and production of polymorphic zooids for defence and reproduction, although some species exhibit reasonable levels of resilience. Some key questions about bryozoan biomineralization that need to be addressed are identified., (© 2014 Natural History Museum. Biological Reviews © 2014 Cambridge Philosophical Society.)