Digging into boring bryozoans: new characters and new species of Immergentiidae.

Immergentia is an endolithic genus of ctenostome bryozoans and the sole member of the Immergentiidae. Etchings of their typical spindled-shaped and sometimes enantiomorphic borehole aperture in calcium carbonate substrates are accomplished by chemical dissolution. The tentacle crown of the bryozoan is essentially the only body part that extends beyond the shell surface when protruded. Previously, species were mainly described using external colony and zooidal characteristics or whole mounts, with partial histological sections conducted on a single species in 1947. Modern approaches, however, are hitherto missing. We examined the soft body morphology of Immergentia from different locations with confocal laser scanning microscopy and the production of 3D reconstructions. In addition, zooidal characteristics such as tentacle number, size, tubulets, and interzooidal distances were used to distinguish and describe species. The combination of conventional and modern methods revealed the presence of a cardiac constrictor and intercalary kenozooids that can interpose between the cystid appendages, something not previously reported in immergentiids, thus necessitating an amendment of the family diagnosis. The polypide typically has eight to ten tentacles, and the anus is positioned in the low or mid-lophophoral area. In addition, sequence data, including the mitogenome and the nuclear ribosomal genes (18S and 28S) of four species from five locations, are presented for the first time. Based on molecular and morphological data, a novel intertidal immergentiid from France, Immergentia stephanieae sp. nov., and a subtidal species from New Zealand, I. pohowskii sp. nov., are described. This work supplements the rather sparse existing knowledge on Immergentiidae and proposes additional characteristics to complement existing descriptions in order to enhance future species identification. Supplementary Information: The online version contains supplementary material available at 10.1007/s13127-024-00645-y.

Boring systematics: A genome skimmed phylogeny of ctenostome bryozoans and their endolithic family Penetrantiidae with the description of one new species.

Ctenostomes are a group of gymnolaemate bryozoans with an uncalcified chitinous body wall having few external, skeletal characters. Hence, species identification is challenging and their systematics remain poorly understood, even more so when they exhibit an endolithic (boring) lifestyle. Currently, there are four Recent families of endolithic bryozoans that live inside mineralized substrates like mollusk shells. In particular, Penetrantiidae Silén, 1946 has received considerable attention and its systematic affinity to either cheilostomes or ctenostomes has been debated. Species delimitation of penetrantiids remains difficult, owing to a high degree of colonial and zooidal plasticity. Consequently, an additional molecular approach is essential to unravel the systematics of penetrantiids, their phylogenetic placement and their species diversity. We therefore sequenced the mitochondrial (mt) genomes and two nuclear markers of 27 ctenostome species including nine penetrantiids. Our phylogeny supports the Penetrantiidae as a monophyletic group placed as sister taxon to the remaining ctenostomes alongside paludicellids, arachnidioids and terebriporids. The boring family Terebriporidae d'Orbigny, 1847 were previously considered to be among vesicularioids, but our results suggest an arachnidioid affinity instead. Ctenostome paraphyly is supported by our data, as the cheilostomes nest within them. A Multiporata clade is also well supported, including the former victorelloid genus Sundanella. Altogether, this study provides new insights into ctenostome systematics, assists with species delimitation and contributes to our understanding of the bryozoan tree of life.

Boring bryozoans: an investigation into the endolithic bryozoan family Penetrantiidae.

An endolithic lifestyle in mineralized substrates has evolved multiple times in various phyla including Bryozoa. The family Penetrantiidae includes one genus with ten extant and two fossil species. They predominantly colonize the shells of molluscs and establish colonies by chemical dissolution of calcium carbonate. Based on several morphological characters, they were described to be either cheilostome or ctenostome bryozoans. For more than 40 years, neither the characters of species identity and systematics nor the problem of their phylogeny was approached. Consequently, the aim of this study is to reevaluate species identities and the systematic position of the genus Penetrantia by analyzing at least six different species from eight regions with the aid of modern methods such as confocal laser scanning microscopy and 3D-reconstruction techniques. This study demonstrates that the musculature associated with the operculum and brood chamber shows significant differences from the cheilostome counterparts and seems to have evolved independently. Together with the presence of other ctenostome-like features such as true polymorphic stolons and uncalcified body wall, this finding supports a ctenostome affinity. Operculum morphology reveals many new species-specific characters, which, together with information about gonozooid morphology, tentacle number, and zooid size ranges, will enhance species identification. It also revealed a probable new species in Japan as well as potential cryptic species in France and New Zealand. In addition, this study increases the known distribution range of the family and its substrate diversity. Altogether, the new information collated here provides the basis for future work on a neglected taxon. Supplementary Information: The online version contains supplementary material available at 10.1007/s13127-023-00612-z.

Multiple evolutionary transitions of reproductive strategies in a phylum of aquatic colonial invertebrates.

Parental care is considered crucial for the enhanced survival of offspring and evolutionary success of many metazoan groups. Most bryozoans incubate their young in brood chambers or intracoelomically. Based on the drastic morphological differences in incubation chambers across members of the order Cheilostomatida (class Gymnolaemata), multiple origins of incubation were predicted in this group. This hypothesis was tested by constructing a molecular phylogeny based on mitogenome data and nuclear rRNA genes 18S and 28S with the most complete sampling of taxa with various incubation devices to date. Ancestral character estimation suggested that distinct types of brood chambers evolved at least 10 times in Cheilostomatida. In Eucratea loricata and Aetea spp. brooding evolved unambiguously from a zygote-spawning ancestral state, as it probably did in Tendra zostericola, Neocheilostomata, and 'Carbasea' indivisa. In two further instances, brooders with different incubation chamber types, skeletal and non-skeletal, formed clades (Scruparia spp., Leiosalpinx australis) and (Catenicula corbulifera (Steginoporella spp. (Labioporella spp., Thalamoporella californica))), each also probably evolved from a zygote-spawning ancestral state. The modular nature of bryozoans probably contributed to the evolution of such a diverse array of embryonic incubation chambers, which included complex constructions made of polymorphic heterozooids, and maternal zooidal invaginations and outgrowths.

Black and male children have an increased risk of palmoplantar psoriasis compared to White children.

A retrospective chart review of 332 pediatric psoriasis patients seen at a single academic institution from 2012 to 2022 was conducted to examine the risk factors associated with palmoplantar psoriasis (PP), a painful and treatment-resistant subtype of plaque psoriasis affecting hands and feet. Black patients have a 6.386-fold increase in the odds of having PP compared to White patients and males have a 2.241-fold increase in the odds of having PP. Black and Hispanic/Latino patients displayed a higher prevalence of nail and palm/sole involvement (p < .0001), whereas White patients exhibited more scalp involvement (p = .04). This study reveals the importance of considering the diagnosis of PP in Black male patients based on its demographic prevalence, which may in turn impact clinical care for these patients.

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.

Paleozoic origins of cheilostome bryozoans and their parental care inferred by a new genome-skimmed phylogeny.

Phylogenetic relationships and the timing of evolutionary events are essential for understanding evolution on longer time scales. Cheilostome bryozoans are a group of ubiquitous, species-rich, marine colonial organisms with an excellent fossil record but lack phylogenetic relationships inferred from molecular data. We present genome-skimmed data for 395 cheilostomes and combine these with 315 published sequences to infer relationships and the timing of key events among c. 500 cheilostome species. We find that named cheilostome genera and species are phylogenetically coherent, rendering fossil or contemporary specimens readily delimited using only skeletal morphology. Our phylogeny shows that parental care in the form of brooding evolved several times independently but was never lost in cheilostomes. Our fossil calibration, robust to varied assumptions, indicates that the cheilostome lineage and parental care therein could have Paleozoic origins, much older than the first known fossil record of cheilostomes in the Late Jurassic.

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

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

Hornera currieae n. sp. (Cyclostomatida: Horneridae): a new bathyal cyclostome bryozoan with reproductively induced skeletal plasticity.

Here we describe a new hornerid, Hornera currieae n. sp. (Bryozoa: Cyclostomatida) from bathyal depths across the New Zealand region. Colonies are irregular, finely branched fans attaining ~40 mm or more in height. Key characters include: (1) thick, semi-hyaline porcellanous skeleton; (2) loss or reduction of nervi (longitudinal striae) away from growing tips; (3) sparse, threadlike cancelli; and (4) small (6187 m), widely spaced autozooidal apertures. Diagnostic hornerid traits possessed by H. currieae n. sp. include vertical ancestrular tube, periancestrular budding of daughter zooids, and skeletal ultrastructure dominated by hexagonal semi-nacre grading to pseudofoliated fabric. The abfrontal incubation chamber develops from a cryptic tube arising from the frontally positioned aperture of the fertile zooid. We used SEM, micro-CT and electron backscatter diffractometry (EBSD) to investigate the ultrastructure and internal architecture of H. currieae n. sp. EBSD reveals that crystalline c-axes of laminated crystallites are perpendicular to skeletal walls. Threadlike cancelli, which traverse secondary calcification, connect autozooidal chambers to the colony-wide hypostegal cavity. Micro-CT reveals that abfrontal cancelli usually bend proximally towards the base, but turn distally towards reproductively active regions of the colony in synchrony with gonozooid development. The zone of affected cancelli extends for 47 branch internodes below the gonozooid. We assessed whether skeletal ultrastructure was similarly affected, but neither cancellus direction, nor gonozooid proximity, were predictive of the crystallite imbrication direction. We hypothesise that (1) hornerid cancelli are active conduits for colonial metabolite transport and (2) that changes in gradients of metabolites and/or reproductive morphogens within the hypostegal cavity affect cancellus morphogenesis. Potentially, H. currieae n. sp. skeletons may preserve a record of intra-colony metabolite translocation dynamics over time.

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.