Trait-Fitness Associations via Fecundity and Competition in a Two-Million-Year-Long Fossil Record.

AbstractThe evolution of phenotypic traits is usually studied on generational timescales or across species on million-year timescales. We bridge this conceptual gap by using high-density sampling of a species lineage, Microporella agonistes (Bryozoa, Cheilostomatida), over 2 million years of its evolutionary history, to ask whether trait-fitness associations are consistent with evolutionary trait models often applied to phenotypic time series. We use average fecundity and competitive outcome as two different fitness components, where competitive outcome is a proxy for partial survival. Examining three quantitative traits in multivariate analyses, we present evidence that some traits experienced substantial selective pressures, in part controlled by past environments. A complex interplay of resource competition with an altering set of competitors and past temperatures has contributed to the changing patterns of phenotypes within the focal species. A comparison with congeneric species living in the same regional community suggests that size traits are more temporally variable and less constrained than shape traits. Our analyses also show that while controls on phenotypes are complex and varied in time, ecological and evolutionary processes that unfold on shorter timescales are not inconsistent with macroevolutionary patterns observed on longer timescales.

Boring life: early colony formation and growth in the endolithic bryozoan genus Penetrantia Silén, 1946.

As in most colonial and sessile marine invertebrates, bryozoan life history is characterized by asexual propagation of zooids for colonial growth and by sexual production of larvae for dispersal. However, comprehensive life histories, particularly in cryptic species such as endolithic (boring) bryozoans, remain poorly understood. The ctenostome family Penetrantiidae is widespread from temperate to tropical waters and often found in molluscan shells, offering an opportunity to study the boring lifestyle and its potential impact on bioerosion through growth and settlement experiments. Our research focused on Penetrantia clionoides from Guam in the Pacific Ocean, Penetrantia japonica from Japan, and a Penetrantia species from France in the Atlantic Ocean. We found distinct life histories and reproductive patterns potentially influenced by environmental factors such as temperature and food availability. The tropical P. clionoides displayed higher rates of larval production and growth compared to its temperate counterpart. For instance, the mean stolon extension was 335.2 µm/week in P. clionoides versus 232.1 µm/week in Penetrantia sp. Autozooid development took 13 days in P. clionoides and 31 days in Penetrantia sp. Anatomical features like apertural rims aided in species identification and in understanding larval settlement preferences, suggesting a tendency for philopatric settlement behavior. The bioerosional impact of penetrantiids remains little understood, but we generated first projections of bioerosion rates and a protocol for keeping Penetrantia under laboratory conditions, laying a foundation for further research in this field.

Effect of a generalist mesopredator on modular and unitary sessile prey associated with a foundation species.

Unitary and modular sessile organisms both dominate in marine benthic communities, commonly preyed upon by the same generalist predators. The differences between unitary and modular defensive strategies may underlie the ways generalist predators control community structure, but this has never been empirically examined. We hypothesize that the individual size of an omnivorous mesopredatory shrimp affects the relative vulnerability of unitary and modular prey and hence translates into community structure. In a short-term laboratory microcosm experiment, we assessed the effect of the shrimp individual size on an epibiotic assemblage of red algae blades initially dominated by three species of modular bryozoans and a unitary serpulid tubeworm. We found that the individual size of a shrimp determines its effect on the prey community composition. Large shrimp stronger than small shrimp increased the proportion of unitary tubeworms among the epibionts surviving predation. While large shrimp reduced the proportions of all the three dominant bryozoan species, small shrimp, in contrast, mostly increased the proportion of a bryozoan species with the smallest modules and largest colonies. This bryozoan, like the tubeworms, demonstrated a higher survival rate with larger individual (colony) size. Yet, against large shrimp this bryozoan was outperformed by the largest tubeworms almost immune to predation. Partial predation by small shrimp modestly improved survival of the largest bryozoan colonies. Thus, relative vulnerability of unitary and modular prey is determined by the predator individual size. Our findings clarify the complex way the size structures of generalist consumers and their prey shape communities by affecting the species-specific relative performance of modular and unitary organisms. The demography of a foundation species and the competitive hierarchy can have additional effects by altering the balance of predation and competition.

Alcyonidium kuklinskii sp. nov., a new species of Antarctic ctenostome bryozoan with a key to all Antarctic species of the genus.

Recent surveys of Antarctic waters in the Terra Nova Bay (Ross Sea) revealed numerous bryozoan species including ctenostome bryozoans. Whereas cheilostome bryozoans are well-studied in these latitudes, ctenostomes remain highly neglected. Large ctenostomes are easily recognized by their lack of calcified skeletons, but this lack also renders them difficult and tedious to identify. As a result, histology and reconstructions of internal soft tissues are required to classify this group of bryozoans. Thanks to the availability of new specimens from Terra Nova Bay, a detailed analysis of growth form, gut morphology and tentacle number of two colonies, initially ascribed to the ctenostome bryozoan genus Alcyonidum Lamouroux, 1813, turned out to be a new species, Alcyonidium kuklinskii sp. nov., which we described in this study. These specimens were also barcoded (COI) and sequences compared to available ones. Together with the new species described here, a total of ten species of Alcyonidium is now known for the Southern Ocean, accounting for one eighth of the entire genus diversity. All Southern Ocean species appear to be endemic. In order to speed the identification of the Antarctic Alcyonidium species, we provide an identification key and a distribution map of all type species. In brief, colony morphology, zooidal size and, in particular tentacle number represent the most suitable characters for identifying species within this genus. Supplementary Information: The online version contains supplementary material available at 10.1007/s13127-023-00629-4.

Deep-sea ctenostome bryozoans: revision of the family Pachyzoidae, with description of a new genus and three new species from Zealandia.

Pachyzoidae is a little-known family of deep-sea ctenostome Bryozoa that until now was monospecific for Pachyzoon atlanticum. Originally described from the Atlantic Ocean, the genus was also found off southeastern New Caledonia in deep waters of the geological continent of Zealandia. Pachyzoon atlanticum forms globular to flat round colonies, living on soft, muddy to sandy bottoms with a few rhizoidal cystid appendages extending from the basal, substrate-oriented side. In this study, we investigate additional pachyzoids, collected between 1965 and 2015 from over 40 sites around New Zealand, by means of detailed morphological and histological investigations. In total, several hundred colonies were encountered in the NIWA Invertebrate Collection, comprising two new species of the genus Pachyzoon, P. grischenkoi sp. nov. and P. pulvinaris sp. nov., and the new genus and species Jeanloupia zealandica gen. et sp. nov.. The genus Jeanloupia is characterized by small disc-shaped colonies with highly elongated peristomes and a quadrangular aperture, distinct from the round apertures of the genus Pachyzoon. Pachyzoid species differ in colony structure and shape, apertural papillae and polypide features such as tentacle number or digestive-tract details. Cystid appendages are non-kenozooidal, but may originate from laterally flanking kenozooids. Based on published images, alleged P. atlanticum from New Caledonia is re-interpreted as P. pulvinaris n. sp.. Morphological characters support alcyonidioidean relationships, as previously suggested. First observations on pachyzoid reproduction show macrolecithal oocytes and brooding of embryos, which seems to be the general pattern for this family. The occurrence of three new Zealandian species in a comparatively small geographical area far from the Atlantic indicates a high possibility of more species to discovered.

Genomic and transcriptomic survey of bryozoan Hox and ParaHox genes with emphasis on phylactolaemate bryozoans.

BACKGROUND: Bryozoans are mostly sessile aquatic colonial invertebrates belonging to the clade Lophotrochozoa, which unites many protostome bilaterian phyla such as molluscs, annelids and brachiopods. While Hox and ParaHox genes have been extensively studied in various lophotrochozoan lineages, investigations on Hox and ParaHox gene complements in bryozoans are scarce. RESULTS: Herein, we present the most comprehensive survey of Hox and ParaHox gene complements in bryozoans using four genomes and 35 transcriptomes representing all bryozoan clades: Cheilostomata, Ctenostomata, Cyclostomata and Phylactolaemata. Using similarity searches, phylogenetic analyses and detailed manual curation, we have identified five Hox genes in bryozoans (pb, Dfd, Lox5, Lox4 and Post2) and one ParaHox gene (Cdx). Interestingly, we observed lineage-specific duplication of certain Hox and ParaHox genes (Dfd, Lox5 and Cdx) in some bryozoan lineages. CONCLUSIONS: The bryozoan Hox cluster does not retain the ancestral lophotrochozoan condition but appears relatively simple (includes only five genes) and broken into two genomic regions, characterized by the loss and duplication of serval genes. Importantly, bryozoans share the lack of two Hox genes (Post1 and Scr) with their proposed sister-taxon, Phoronida, which suggests that those genes were missing in the most common ancestor of bryozoans and phoronids.

Anti-Inflammatory Effects of Bioactive Compounds from Seaweeds, Bryozoans, Jellyfish, Shellfish and Peanut Worms.

Inflammation is a defense mechanism of the body in response to harmful stimuli such as pathogens, damaged cells, toxic compounds or radiation. However, chronic inflammation plays an important role in the pathogenesis of a variety of diseases. Multiple anti-inflammatory drugs are currently available for the treatment of inflammation, but all exhibit less efficacy. This drives the search for new anti-inflammatory compounds focusing on natural resources. Marine organisms produce a broad spectrum of bioactive compounds with anti-inflammatory activities. Several are considered as lead compounds for development into drugs. Anti-inflammatory compounds have been extracted from algae, corals, seaweeds and other marine organisms. We previously reviewed anti-inflammatory compounds, as well as crude extracts isolated from echinoderms such as sea cucumbers, sea urchins and starfish. In the present review, we evaluate the anti-inflammatory effects of compounds from other marine organisms, including macroalgae (seaweeds), marine angiosperms (seagrasses), medusozoa (jellyfish), bryozoans (moss animals), mollusks (shellfish) and peanut worms. We also present a review of the molecular mechanisms of the anti-inflammatory activity of these compounds. Our objective in this review is to provide an overview of the current state of research on anti-inflammatory compounds from marine sources and the prospects for their translation into novel anti-inflammatory drugs.

Bioprospecting for polyesterase activity relevant for PET degradation in marine Enterobacterales isolates.

Plastics have quickly become an integral part of modern life. Due to excessive production and improper waste disposal, they are recognized as contaminants present in practically all habitat types. Although there are several polymers, polyethylene terephthalate (PET) is of particular concern due to its abundance in the environment. There is a need for a solution that is both cost-effective and ecologically friendly to address this pollutant. The use of microbial depolymerizing enzymes could offer a biological avenue for plastic degradation, though the full potential of these enzymes is yet to be uncovered. The purpose of this study was to use (1) plate-based screening methods to investigate the plastic degradation potential of marine bacteria from the order Enterobacterales collected from various organismal and environmental sources, and (2) perform genome-based analysis to identify polyesterases potentially related to PET degradation. 126 bacterial isolates were obtained from the strain collection of RD3, Research Unit Marine Symbioses-GEOMAR-and sequentially tested for esterase and polyesterase activity, in combination here referred to as PETase-like activity. The results show that members of the microbial families Alteromonadaceae, Shewanellaceae, and Vibrionaceae, derived from marine sponges and bryozoans, are the most promising candidates within the order Enterobacterales. Furthermore, 389 putative hydrolases from the α/ß superfamily were identified in 23 analyzed genomes, of which 22 were sequenced for this study. Several candidates showed similarities with known PETases, indicating underlying enzymatic potential within the order Enterobacterales for PET degradation.

Bryoherms from the lower Sarmatian (upper Serravallian, Middle Miocene) of the Central Paratethys.

Bryozoan-serpulid-algal-thrombolite bioherms of up to 50 cm size are described from the Sarmatian (upper Middle Miocene) of the Central Paratethys. They occur on top of lower Sarmatian carbonate sediments of high-energy conditions and the individual bioherms settle on crests of ripples. The buildups are overlain and partly truncated by cross-bedded oolites of late Sarmatian age. Buildup growth starts with a Cryptosula/Hydroides (bryozoan/serpulid) pioneer community, followed by nodular Schizoporella (bryozoan) colonies overgrown by coralline algae/microbial mats and a thrombolite with calcareous algal filaments. All these constituents form a framestone fabric which is overall dominated by bryozoans labeling them as bryoherms. Inside the bioherms ecological successions of higher frequencies occur which are interpreted to reflect short-time environmental fluctuations such as nutrient availability, oxygenation (possible anoxia), salinity (possible brackish water), temperature and water level. The internal succession in individual bioherms is related to long-term environmental changes including general shallowing, increasing nutrient supply and decreasing water circulation and oxygenation. The described bioherms are most similar to modern bryostromatolites of the Coorong lagoon in S Australia and also similar to structures in the Netherlands. The widespread occurrence of bryoherms/bryostromatolites in the Central Paratethys suggests a phase of considerable eutrophication during the early Sarmatian.

Phylogenomics reveals deep relationships and diversification within phylactolaemate bryozoans.

Bryozoans are mostly sessile colonial invertebrates that inhabit all kinds of aquatic ecosystems. Extant bryozoan species fall into two clades with one of them, Phylactolaemata, being the only exclusively freshwater clade. Phylogenetic relationships within the class Phylactolaemata have long been controversial owing to their limited distinguishable characteristics that reflect evolutionary relationships. Here, we present the first phylogenomic analysis of Phylactolaemata using transcriptomic data combined with dense taxon sampling of six families to better resolve the interrelationships and to estimate divergence time. Using maximum-likelihood and Bayesian inference approaches, we recovered a robust phylogeny for Phylactolaemata in which the interfamilial relationships are fully resolved. We show Stephanellidae is the sister taxon of all other phylactolaemates and confirm that Lophopodidae represents the second offshoot within the phylactolaemate tree. Plumatella fruticosa clearly falls outside Plumatellidae as previous investigations have suggested, and instead clusters with Pectinatellidae and Cristatellidae as the sister taxon of Fredericellidae. Our results demonstrate that cryptic speciation is very likely in F. sultana and in two species of Plumatella (P. repens and P. casmiana). Divergence time estimates show that Phylactolaemata appeared at the end of the Ediacaran and started to diverge in the Silurian, although confidence intervals were large for most nodes. The radiation of most extant phylactolaemate families occurred mainly in the Palaeogene and Neogene highlighting post-extinction diversification.

Collecting and Culturing Bryozoans for Regenerative Studies.

Among marine invertebrates, bryozoans are small, not well known, and complex to identify. Nevertheless, they offer unique opportunities for whole-body generation research, because of their colonial, modular mode of growth. Here, we describe detailed methods for collection of bryozoans from a range of environments, sample preparation and identification, culture and feeding, spawning and breeding, marking colonies for growth studies, and histological preparation.

Alien hotspot: Benthic marine species introduced in the Brazilian semiarid coast.

In this study, we provide a baseline assessment of introduced marine species along the extensive (~600 km) Brazilian semiarid coast. We reported 27 introduced and 26 cryptogenic species. Moreover, the main vectors of introduction were ballast water, shipping lines, oil and gas activities, biofouling, and rafting on plastic debris. The taxa were Ascidiacea (17 species) and Bryozoa (17), followed by Crustacea (6), Mollusca (6), Cnidaria (3), Echinodermata (3), and Porifera (1). Among these invertebrates, some species are recognized as drivers of impacts such as the invasive corals (Tubastraea tagusensis and Tubastraea coccinea), the bivalves Isognomom bicolor and Perna viridis, the crab Charybdis hellerii, the brittle star Ophiothela mirabilis, and, finally, the bryozoan Membraniporopsis tubigera. These species threaten the biodiversity of unique ecosystems such as intertidal sandstone reefs, shallow-water coral reefs, and mesophotic ecosystems. Moreover, the up-to-date results highlight that this region is a hotspot of bioinvasion in the tropical South Atlantic.

Amazing Discoveries of Benthic Fauna from the Abyssal Zone of Lake Baikal.

Lake Baikal is a natural laboratory for the study of species diversity and evolution, as a unique freshwater ecosystem meeting the all of the main criteria of the World Heritage Convention. However, despite many years of research, the true biodiversity of the lake is clearly insufficiently studied, especially that of deep-water benthic sessile organisms. For the first time, plastic waste was raised from depths of 110 to 190 m of Lake Baikal. The aim of this study was to examine the biological community inhabiting the plastic substrate using morphological and molecular genetic analysis. Fragments of plastic packaging materials were densely populated: bryozoans, leeches and their cocoons, capsules of gastropod eggs, and turbellaria cocoons were found. All the data obtained as a result of an analysis of the nucleotide sequences of the standard bar-coding fragment of the mitochondrial genome turned out to be unique. Our results demonstrate the prospects for conducting comprehensive studies of artificial substrates to determine the true biodiversity of benthos in the abyssal zone of Lake Baikal.

From the individual to the colony: Marine invertebrates as models to understand levels of biological organization.

The developmental and evolutionary principles of coloniality in marine animals remain largely unexplored. Although many common traits have evolved independently in different groups of colonial animals, questions about their significance for colonial life histories remain unanswered. In 2018 (Nov. 25 - Dec. 8), the inaugural course on the Evolution of Coloniality and Modularity took place at the Center for Marine Biology of the University of São Paulo (CEBIMAR-USP), Brazil. During the intensive two-week graduate-level course, we addressed some of the historical ideas about animal coloniality by focal studies in bryozoans, tunicates, cnidarians, and sponges. We discussed many historical hypotheses and ways to test these using both extant and paleontological data, and we carried direct observations of animal colonies in the different phyla to address questions about coloniality. We covered topics related to multi-level selection theory and studied colonial traits, including modular miniaturization, polymorphism, brooding, and allorecognition. Course participants carried out short research projects using local species of animals to address questions on allorecognition and regeneration in ascidians and sponges, fusion and chimerism in anthoathecate hydrozoans, and evolution of polymorphism in bryozoans. Although many questions remain unanswered, this course served as a foundation to continue to develop a developmental and evolutionary synthesis of clonal and modular development in colonial marine organisms.

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.

Ultrastructure of rhizoids in the marine bryozoan Dendrobeania fruticosa (Gymnolaemata: Cheilostomata).

Bryozoans form colonies of iterated modules, termed zooids, and display varying degrees of polymorphism. Polymorphic colonies comprise autozooids (or feeding zooids) and heteromorphic zooids, among which the most common types are avicularia and kenozooids. Kenozooids differ in shape, size, and presumed function. Among this diversity, there are rhizoids, which serve to attach colonies to the substrate or to lift them above it. To date, only general data on anatomy of kenozooids at light microscopy level are available. Here, we present the first description of the ultrastructure of the holdfast-like rhizoids of the cheilostome bryozoan Dendrobeania fruticosa. The rhizoid wall is composed of a single-layered epidermis, which produces the ectocyst. The voluminous cavity is acoelomate: it has no special cellular lining, nor any signs of an extracellular matrix toward the epidermis. It is traversed by delicate branching funicular strands that originate from the pore plate. The only cells in contact with the epidermis are the cells of the funicular system and the storage cells. The pore plate between the rhizoid and autozooid includes a variable number of communication pores. Each pore is plugged with a rosette complex, which includes a cincture cell and four special cells extending through the pore. The limiting cells are absent, and the special cells are in direct contact with the funicular strands. Cell contacts between special cells are absent; moreover, there are spaces between their proximal lobes filled with a heterogeneous matrix similar to that in the lumen of the funicular strands. Such matrix is also found outside of the extracellular matrix surrounding the special cells. These findings allow us to suggest that nutrient transport most likely occurs between, rather than through, the special cells. However, further studies are needed to understand how the rosette complex functions.

A molecular phylogeny of historical and contemporary specimens of an under-studied micro-invertebrate group.

Resolution of relationships at lower taxonomic levels is crucial for answering many evolutionary questions, and as such, sufficiently varied species representation is vital. This latter goal is not always achievable with relatively fresh samples. To alleviate the difficulties in procuring rarer taxa, we have seen increasing utilization of historical specimens in building molecular phylogenies using high throughput sequencing. This effort, however, has mainly focused on large-bodied or well-studied groups, with small-bodied and under-studied taxa under-prioritized. Here, we utilize both historical and contemporary specimens, to increase the resolution of phylogenetic relationships among a group of under-studied and small-bodied metazoans, namely, cheilostome bryozoans. In this study, we pioneer the sequencing of air-dried cheilostomes, utilizing a recently developed library preparation method for low DNA input. We evaluate a de novo mitogenome assembly and two iterative methods, using the sequenced target specimen as a reference for mapping, for our sequences. In doing so, we present mitochondrial and ribosomal RNA sequences of 43 cheilostomes representing 37 species, including 14 from historical samples ranging from 50 to 149 years old. The inferred phylogenetic relationships of these samples, analyzed together with publicly available sequence data, are shown in a statistically well-supported 65 taxa and 17 genes cheilostome tree, which is also the most broadly sampled and largest to date. The robust phylogenetic placement of historical samples whose contemporary conspecifics and/or congenerics have been sequenced verifies the appropriateness of our workflow and gives confidence in the phylogenetic placement of those historical samples for which there are no close relatives sequenced. The success of our workflow is highlighted by the circularization of a total of 27 mitogenomes, seven from historical cheilostome samples. Our study highlights the potential of utilizing DNA from micro-invertebrate specimens stored in natural history collections for resolving phylogenetic relationships among species.

Evolutionary conflict and coloniality in animals.

Despite considerable interest in the effects of evolutionary conflict in colonies of social insects, relatively little attention has been paid to this issue in clonal animals with modular construction, such as colonial ascidians, bryozoans, and cnidarians. These colonial animals are structural individuals, subdivided into repeated morphological modules, which can individually acquire, process, and share resources. While size-related selection favors colony formation, evolutionary conflicts remain a potent obstacle to such cooperation. These conflicts can occur at several levels and must be mediated for cooperation to emerge. Module-level conflicts potentially result in coalitions of genetically similar modules failing to share resources or monopolizing reproduction. Mediation occurs by a number of mechanisms including: (a) a single-module bottleneck at the initiation of colony formation, (b) allorecognition that limits colony fusion to close kin, (c) development of new modules from connective tissue, (d) synchronization of module budding, (e) programmed module death, (f) terminal differentiation of reproductive modules, and (g) architectural constraints. Effective mediation of module-level conflicts, however, may in some cases contribute to cell-level conflicts. Animal colonies typically have multipotent stem cells, and genetically variant stem cells can potentially monopolize gamete formation. Limiting colony fusion to close kin may not eliminate such conflict. Finally, in at least some taxa an association between photosymbiosis and coloniality is found. Allocation of photosynthate can lead to host-symbiont conflicts that can be mediated by housing symbionts intracellularly and using chemiosmotic mechanisms to detect defectors. Colonial animals thus serve as a living laboratory of evolutionary conflict and its mediation.

Broad but restricted detection of malacosporeans in a Neotropical cradle of diversification.

This study undertook the first investigation of malacosporean infections in Neotropical fish. We used polymerase chain reaction detection with a primer set generally targeting known malacosporeans to assay for infection in the kidney of 146 fish in 21 species belonging to 12 families collected from two areas in the Amazon Basin. Infections were found in 13 fish variously belonging to seven species in six families and included the first identification of a malacosporean infection in cartilaginous fish (a freshwater stingray). Based on ssrDNA, all infections represented a single Buddenbrockia species (Buddenbrockia sp. E) that demonstrates an exceptionally broad range of fish species infected, and countered our expectations of high Neotropical malacosporean diversity. Infections were characterized at varying and often high prevalences in fish species but sample sizes were small. Ascertaining whether highly divergent malacosporeans have not been detected by current primers, and more comprehensive sampling may reveal whether malacosporeans are truly as species poor in the Amazon Basin as present data suggest. Our results prompt speculations about evolutionary scenarios including introduction via marine incursions and patterns of host use over time.

Using encrusting bryozoan adhesion to evaluate the efficacy of fouling-release marine coatings.

Biofouling communities are spatiotemporally diverse, underscoring the need to assess fouling-release (FR) coating performance against common biofouling taxa at multiple field sites. Adhesion strength assessments of FR coatings incorporate few taxa into standardized protocols. This study tested the feasibility of incorporating existing ASTM barnacle protocols on tubeworms and encrusting bryozoans (EB). Additionally, trends in adhesion strength among these taxa were compared at two field sites. EB adhesion at both field sites showed consistent results and adhesion strength followed the same trend: tubeworms > barnacles >EB. Testing EB adhesion was feasible and enhanced assessments of FR coatings by increasing the diversity of assessed taxa.