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.

Kelp species are ecologically-important habitat-formers in coastal marine ecosystems, where they alter environmental conditions and promote local biodiversity by providing complex biogenic habitat for an array of associated organisms. While it is widely accepted that kelps harbour significant biodiversity, our current understanding of spatiotemporal variability in kelp-associated assemblages and the key environmental drivers of variability patterns remains limited. Here we examined the influence of ocean temperature and wave exposure on the structure of faunal assemblages associated with the holdfasts of Laminaria hyperborea, the dominant habitat-forming kelp in the northeast Atlantic. We sampled holdfasts from 12 kelp-dominated open-coast sites nested within four regions across the UK, spanning ~9° in latitude and ~2.7° C in mean sea surface temperature. Overall, holdfast assemblages were highly diverse, with 261 taxa representing 11 phyla recorded across the study. We examined patterns of spatial variability for sessile and mobile taxa separately, and documented high variability between regions, between sites within regions, and between replicate holdfasts for both assemblage types. Mobile assemblage structure was more strongly linked to temperature variability than sessile assemblage structure, which was principally structured by site-level variability in factors such as wave exposure. Patterns in the structure of both biogenic habitat and associated assemblages did not vary predictably along a latitudinal gradient in temperature, indicating that other processes acting across multiple spatial and temporal scales are important drivers of assemblage structure. Overall, kelp holdfasts in the UK supported high levels of diversity, that were similar to other kelp-dominated systems globally and comparable to those recorded for other vegetated marine habitats (i.e. seagrass beds), which are perhaps more widely recognised for their high biodiversity value.

Polyembryony–the production of multiple cloned embryos from a single fertilised egg–is a seemingly paradoxical combination of reproductive modes that nevertheless persists in diverse taxa. We document features of polyembryony in the Cyclostomata (Bryozoa)–an ancient order of modular colonial marine invertebrates–that suggest a substantial reduction in the paradoxical nature of this enigmatic reproductive mode. Firstly, we provide molecular evidence for polyembryony in three exemplar species, supporting the widely cited inference that polyembryony characterises the entire order. Secondly, genotyping demonstrates protracted release of cloned offspring from the primary embryo in a given gonozooid (chamber for embryonic incubation), thus exposing the same genotype to changing environmental conditions over time. Finally, we confirm that each gonozooid produces a distinct genotype, with each primary embryo being the result of a separate fertilisation event. We hypothesise that the sustained release of one or a few genotypes against varying environmental conditions achieves levels of risk-spreading similar to those in organisms that release multiple, unique genotypes at a single time. We argue that polyembryony, specifically with the production of a large number of progeny per fertilisation event, has been favoured in the Cyclostomata over long geological periods.

Foundation species elevate local biodiversity and underpin critical ecological processes and functions. Kelp species are distributed along ~25% of the world’s coastlines, where they serve as foundation species in intertidal and subtidal habitats. As well as ameliorating environmental conditions and producing organic matter, they provide biogenic habitat for a vast array of associated organisms. Here, we investigated patterns of diversity and structure in assemblages associated with the stipe of the kelp Laminaria hyperborea in the NE Atlantic. Stipes were sampled at 4 study regions (with 3 sites nested within each region) in the UK, spanning ~9° of latitude. Stipe-associated communities were highly diverse (134 species) and abundant (16-4532 ind. stipe-1), with no obvious sequential shift in diversity or overall trends in abundance/biomass of assemblages with latitude. However, we observed high degrees of variability between sites from the same region and individuals within sites, indicating that processes working across smaller spatial scales were more important than those at regional scales. While we observed high between-site variability in assemblage structure, regional differences were also evident. Most notably, sites within our southernmost region (southern England) were largely devoid of amphipods that dominated all other regions. This study highlights the important role of L. hyperborea in elevating biodiversity at local to regional scales through a facilitative interaction. Moreover, given that L. hyperborea forests may be increasingly impacted by ocean warming, changes in coastal water quality and proposed exploitation, our study serves as an important benchmark against which to detect future changes.