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Environmental data used in the RLQ analyses.

Environmental data used in the RLQ analyses.

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Article
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Understanding community assembly is a key goal in community ecology. Environmental filtering influences community assembly by excluding ill‐adapted species, resulting in communities with similar functional traits. An RLQ (a four‐way ordination) analysis incorporating spatial data was run on a dataset of 642 species of cheilostomes (Bryozoa) from 77...

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Context 1
... NMDS was used identify groups of closely related variables prior to examination of variable inflation factors (VIFs; R package car; Fox and Weisberg 2019). The environmental variables selected for RLQ following NMDS and VIF are given in Table 1. Finally, a Moran's I test was performed to determine if environmental variables were spatially autocorrelated (R package ade4; Dufour 2007). ...
Context 2
... NMDS was used identify groups of closely related variables prior to examination of variable inflation factors (VIFs; R package car; Fox and Weisberg 2019). The environmental variables selected for RLQ following NMDS and VIF are given in Table 1. Finally, a Moran's I test was performed to determine if environmental variables were spatially autocorrelated (R package ade4; Dufour 2007). ...
Context 3
... NMDS was used identify groups of closely related variables prior to examination of variable inflation factors (VIFs; R package car; Fox and Weisberg 2019). The environmental variables selected for RLQ following NMDS and VIF are given in Table 1. Finally, a Moran's I test was performed to determine if environmental variables were spatially autocorrelated (R package ade4; Dufour 2007). ...

Citations

... Studies show significant trait-environment relationships in marine species (Fernandez, Collins, Gittenberger, et al., 2020;Jansen et al., 2018;McClain et al., 2020;Mindel et al., 2016). A study of trait-environment relationships in bryozoans found that colony shape varies with depth and substrate and that polymorphism varies with depth and oxygen levels (Schack et al., 2020). Likewise, life-history traits of marine fish, such as body length and growth, respond strongly to the environment (Beukhof et al., 2019). ...
... Although some studies have described significant traitenvironment relationships in marine species (Beukhof et al., 2019;Fernandez, Collins, Gittenberger, et al., 2020;Jansen et al., 2018;McClain et al., 2020;Mindel et al., 2016;Schack et al., 2020), few have also addressed the contribution of evolutionary history to trait biogeography in the marine environment (McLean et al., 2021). By jointly analysing spatial, environmental, trait and phylogenetic data of the species, we aim to test trait-environment relationships in hydroids across large spatial and environmental gradients and to evaluate associations between traits, environmental variables, space and phylogeny. ...
Article
The aim was to test trait–environment relationships in hydroids across large spatial and environmental gradients and to evaluate associations between traits, environmental variables, space and phylogeny. Atlantic Ocean and adjacent polar seas. Present day. Hydrozoa. Trait–environment relationships and their spatial and phylogenetic contexts were assessed in hydroids by using a combination of a fourth‐corner test and extended‐RLQ approach, in a multivariate ordination method that uses five matrices: (1) species across sites, (2) environmental data across sites, (3) traits across species, (4) latitude and longitude for each site, and (5) phylogenetic distance among species. We based our analyses on 3680 records of 431 species distributed at 1440 sites, 16 species traits, nine environmental variables and a phylogenetic tree of the studied species. Hydroid traits are significantly correlated with environmental variables and geographical space, and hydroid phylogeny is correlated with the environment and geographical space. More evidently, we observed an increased presence of larger species (taller, more branched, with greater base diameter and polysiphonic) at sites richer in nitrate and silicate, with higher dissolved oxygen, lower temperatures, lower salinities and lower current velocities. The observation of phylogenetically related species with similar traits and distributions corroborates that historical processes interact with the environment in determining community assembly and explaining patterns of distribution of species traits. Several environmental variables combined affect the distribution of hydroid traits, which are also influenced by spatial and historical factors. In the first analysis of its kind for hydrozoans, we have provided an overview of how hydroids are distributed across environments according to their traits and revealed the spatial and phylogenetic components of these trait–environment relationships.
... colonial or erect) differed between the two regions. Planktotrophic (feeding) larvae are capable of re-settling to avoid non-optimal substrate [71], colonial forms are associated with high overgrowth abilities [72], and erect forms can provide better feeding performance [73]. Therefore, trait divergence observed later in development likely resulted from competitive dynamics and a mix of common and distinct trait mechanisms that define competitive abilities at each region. ...
Article
Assembly processes are highly dynamic with biotic filters operating more intensely at local scales, yet the strength of biotic interactions can vary across time and latitude. Predation, for example, can be stronger at lower latitudes, while competition can intensify at later stages of assembly due to resource limitation. Since biotic filters act upon functional traits of organisms, we explored trait-mediated community assembly in diverse marine assemblages from four regions along the Pacific coast of North and Central America. Using predator exclusion experiments and two assembly stages, we tested the hypotheses that non-random trait patterns would emerge during late assembly at all regions due to competition and at lower latitude regions regardless of assembly stage due to predation. As expected, trait divergence occurred in late assembly but only at higher latitude regions, while in tropical Panama, relaxed predation caused trait divergence during late assembly. Moreover, colonizing trait strategies were common during early assembly while competitive strategies were favoured during late assembly at higher latitude regions. Predation-resistant traits were only favoured in Panama during both assembly stages. Our large-scale manipulative study demonstrates that different biotic interactions across time and latitude can have important consequences for trait assembly.
... However, they show profound abundance in the shelf waters up to 500m of water depths (Cook, 1981;Gordon, 1986Gordon, , 1999Wood et al., 2012;Smith, 2014). Each colony (the zoarium) is composed of many individual tubular or box-shaped modular units (zooids); each being enclosed in an exoskeletal case (the zooecium) which opens through an opening (orifice) on the outside, and it is sometimes covered by a lid like structure (operculum) (Cheetham and Cook, 1983;Taylor, 1990;Jablonski et al., 1997;Gordon et al., 2007;Naufal et al., 2018;Schack et al., 2019;2020). All the species known from this Phylum belong to three classes -Class Gymnolaemata and Class Stenolaemata comprising the majority of both uncalcified and well calcified marine forms whereas the minor Class Phylactolaemata includes uncalcified freshwater forms (Smith, 2014, Venkatraman et al., 2016. ...
... Their abundances in the marine realm are largely controlled by the availability of favourable substrate in addition to the changes in the existing physico-chemical parameters. Several studies conducted across the globe have shown that growth forms of bryozoan colonies are related to depths, hydrodynamic energy conditions, acidification of marine environment, substrate availability and rate of sedimentation Lagaaij and Gautier, 1965;Schopf, 1969;Harmelin, 1988;Smith, 1995;Taylor et al., 2004;Moissette et al., 2007;Rodolfo-Metalpa et al., 2010;Swezey et al., 2017;Schack et al., 2020). They are also found to be sensitive to global rise in temperature, bottom level fishing and anthropogenic pollutants. ...
... Nellia tenella is indicative of warm tropical waters and prefer euryhaline salinity conditions. These species are also capable of surviving high salinity and show great tolerance to changes in environment (Sonar et al., 2010) including less food availability (Schack et al., 2020). Erect, biserial colonies of Licornia sp. also point to its profound shallow water preference while attaching to a substratum which sometimes can be artificial such as marine debris (Vieira et al., 2013). ...
Article
The ecological record of bryozoans from the continental shelf-slope region of the southwestern Bay of Bengal is very sparse, prompting this investigation. Forty-five sea-floor sediment samples were collected along eight transects during the cruise on R/V Sagar Paschimi from the offshore region between Chennai and Cuddalore in the southwestern Bay of Bengal. The distribution of bryozoans was assessed based on the fragments of bryozoan colonies from these samples, which were collected at depths ranging from 6–308m. The goal was to determine the relationship between environmental parameters measured from the study area and the abundances of various bryozoan species. Results revealed the presence of 29 species of recent bryozoans in 24 genera pertaining to eight morphotypes. It is evident from the present investigation that the major control on the spatial distribution of bryozoan colonies is the type of substrate available, as sample sites dominated by coarse sediments showed high species richness though other factors such as ocean dynamics and rate of sedimentation also seem to regulate their abundance.
... To assess whether some plant species have a higher chance of being transported by particular waterfowl species, we used RLQ (Dolédec et al., 1996) and fourth-corner analyses (Dray et al., 2014). RLQ has traditionally been recommended to evaluate univariate relationships between environmental variables and species functional traits (Almeida et al., 2018;Schack et al., 2020;Silva J. L. A. et al., 2021). We have utilized the method to look instead at the relationships between waterfowl species traits (substituting environmental variables) and plant species functional traits. ...
Article
Full-text available
Endozoochory by waterfowl is important for a broad range of angiosperms, most of which lack a fleshy fruit. This dispersal function contributes to the formation and maintenance of plant communities and may allow range shifts for plant species under global change. However, our current understanding of what seed or plant traits are important for this dispersal mechanism, and how they relate to variation in waterbird traits, is extremely limited. We addressed this question using a unique dataset identifying the plant species whose seeds are ingested by 31 different waterfowl species in Europe. We used RLQ and fourth-corner analyses to explore relationships between (1) bird morphological and foraging strategy traits, and (2) plant traits related to seed morphology, environmental preferences, and growth form. We then used Generalized Additive Models to identify relationships between plant/seed traits and the number of waterfowl species that disperse them. Although many waterfowl feed intentionally on seeds, available seed trait data provided little explanation for patterns compared to plant traits such as Ellenberg indicators of habitat preference and life form. Geese were associated with terrestrial plants, ingesting seeds as they graze on land. Diving ducks were associated with strictly aquatic plants, ingesting seeds as they feed at greater depths. Dabbling ducks ingest seeds from plants with high light and temperature requirements, especially shoreline and ruderal species growing in or around the dynamic and shallow microhabitats favored by these birds. Overall, the number of waterfowl vector species (up to 13 per plant species) increases for plants with greater soil moisture requirements and salinity tolerance, reflecting the inclination of most waterfowl species to feed in coastal wetlands. Our findings underline the importance of waterfowl dispersal for plants that are not strictly aquatic, as well as for plants associated with high salinities. Furthermore, our results reveal a soil moisture gradient that drives seed-bird interactions, in line with differences between waterfowl groups in their microhabitat preferences along the land-water continuum. This study provides an important advance in our understanding of the interactions that define plant dispersal in wetlands and their surroundings, and of what plants might be affected by ongoing changes in the distributions of waterfowl species.
... Last, but not least, New Zealand is one of the better-studied marine regions taxonomically and ecologically for Bryozoa (e.g. Gordon, 1984;1986;Gordon et al., 2009;Schack et al., 2020), a phylum that is somewhat neglected in many other parts of the world. Bryozoan research has been continuously conducted in New Zealand since 1841 ) and a governmental agency, the National Institute of Water and Atmospheric Research (NIWA), is both the data manager and custodian for fisheries and invertebrate research data, hence assuring knowledge curation. ...
Article
Full-text available
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.
... Last, but not least, New Zealand is one of the better-studied marine regions taxonomically and ecologically for Bryozoa (e.g. Gordon, 1984Gordon, , 1986Gordon, , 1989Gordon et al., 2009;Schack et al., 2020), a phylum that is somewhat neglected in many other parts of the world. Bryozoan research has been continuously conducted in New Zealand since 1841 ) and a governmental agency, the National Institute of Water and Atmospheric Research (NIWA), is both the data manager and custodian for fisheries and invertebrate research data, hence assuring knowledge curation. ...
Preprint
Full-text available
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, 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 asked if its presence has any bearing on the diversification rates of cheilostomes.
Article
Bryozoans, simple invertebrates living on the sea floor, are emerging as a model system for understanding ecological and evolutionary processes on macroevolutionary scales.
Article
Cheilostomata is the most diverse and ecologically dominant order of bryozoans living today. We apply a Bayesian framework to estimate macroevolutionary rates of cheilostomes since the Late Jurassic across four datasets: (I) manually curated genus ranges; (II) published text-mined genus ranges; (III) non-revised Paleobiology Database (PBDB) records; (IV) revised and augmented PBDB records. All datasets revealed increased origination rates in the Albian, and a twin K–Pg and Danian extinction rate peak. High origination rates in the Late Selandian–Ypresian in Dataset I indicate the onset of an ascophoran-grade radiation. Lineage-through-time plots confirm the macroevolutionary lag preceding the radiation of cheilostomes in the middle Cretaceous, and their renewed diversification in the late Paleocene and Eocene. A multivariate birth–death model indicates that origination rates are shaped by diversity-dependent dynamics coupled with a positive correlation with sea surface temperature, while extinction rates negatively correlate with sea level. Text-mined data provide broadly similar rate dynamics as manually curated data, although discrepancies could be attributed to the omission of key literature in Dataset II, and the inclusion of new published and unpublished data, and revised ranges in Dataset I. Revision and augmentation of PBDB occurrences were necessary to generate rate profiles akin to those of Datasets I and II and highlight the risks of using unedited occurrence data. Our results support the widely held assumption that diversification dynamics are controlled by both biotic and abiotic factors, and pave the way for integrating fossils with molecular phylogenies to study these processes in more detail.
Thesis
Full-text available
Cheilostomatous bryozoans are biomineralisers, as such, vulnerable to acidification and increasing temperature currently affecting the world’s oceans. Cheilostomes have excelled in zooid polymorphism and they also perform a high array of growth forms, geometries and mineralogies. Flustriforms, and related taxa, are considerably speciose, ubiquitous, and highly abundant in the benthic realm. An extensive literature review coupled with the analysis of spatial distribution data was performed to address growth as a fundamental biological process of multidimensional expressions. Growth can manifest in zooidal production, somatic growth (both zooids and colonial) and area. However, a more comparable, albeit requiring additional analyses is the measurement of calcification (carbonate production). When temporal dimension is considered, growth is expressed as rates. At wider temporal scales, growth can be expressed via carbon immobilisation (secondary production) and sequestration, whereas, at wider spatial scales, growth can be measured by the expanding distribution ranges. Higher temperatures are expected to accelerate metabolism, hence growth rates, but will affect calcification. Broadly, near-future predicted scenarios will likely increase growth in flustrines in most dimensions addressed in this thesis. Evaluating these trends in a polar-temperate gradient will provide important insights of the underlying mechanisms and possible ecological trade-offs in cheilostomatous bryozoans.