Increasing risk of invasions by organisms on marine debris in the Southeast coast of India.

Increasing amount of anthropogenic litter in the marine environment has provided an enormous number of substrates for a wide range of marine organisms, thus serving as a potential vector for the transport of fouling organisms. Here, we examined the fouling organisms on different types of stranded litter (plastic, glass, rubber, foam sponge, cloth, metal and wood) on eight beaches along the southeast coast of India. In total, 17 encrusting species belonging to seven phyla (Arthropoda, Bryozoa, Mollusca, Annelida, Cnidaria, Chlorophyta and Foraminifera) were identified on 367 items, with one invasive species, the mussel Mytella strigata, detected. The most common species associated with marine litter were the cosmopolitan bryozoans Jellyella tuberculata (%O = 31.64 %) and J. eburnea (28.61 %), the barnacle species Lepas anserifera (29.97 %), Amphibalanus amphitrite (22.34 %) and Amphibalanus sp. (14.16 %), and the oyster species Saccostrea cucullata (13.62 %) and Magallana bilineata (5.44 %). We also reported the first records on stranded litter of four species: the gastropod species Pirenella cingulata and Umbonium vestiarium, the foraminiferan Ammonia beccarii, and the oyster M. bilineata. This study is thus the first documentation of marine litter as a vector for species dispersal in India, where the production and consumption of plastic rank among the highest in the world. We also highlight the increasing risk of invasions by non-indigenous organisms attached to debris along the southeast coast of India. Comprehensive monitoring efforts are thus needed to elucidate the type of vectors responsible for the arrival of invasive species in this region. Raising awareness and promoting education are vital components in fostering sustainable solutions to combat plastic pollution in the country and globally.

Changing allometric relationships among fossil and Recent populations in two colonial species.

Allometry is vital for understanding the mechanisms underlying phenotypic evolution. Despite a large body of literature on allometry, studies based on fossil time series are limited for solitary organisms and nonexistent for colonial organisms. Allometric relationships have been found to be relatively constant across Recent populations of the same species, separated by space, but variable among fossil populations separated by thousands of years. How stable are allometric relationships at the module level for colonial organisms? We address this question using two extant species of the cheilostome bryozoan Microporella with fossil records spanning the Pleistocene of New Zealand. We investigate size covariation between feeding modules and three traits with separate functions (reproductive, resource uptake, and defense). We found that within-population (static) allometry can change on timescales of at least 0.1 million years. These within-population relationships do not consistently predict overintraspecific evolutionary allometry, which in turn does not predict those estimated at the genus level. Different functional traits are constrained to different extents by module size with defensive traits being the least constrained and most evolvable, compared with reproductive and resource uptake traits. Our study highlights the potential of colonial organisms in understanding the constraints and drivers of long-term phenotypic change.

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.

Marine biofouling organisms on beached, buoyant and benthic plastic debris in the Catalan Sea.

Plastic debris provides long-lasting substrates for benthic organisms, thus acting as a potential vector for their dispersion. Its interaction with these colonizers is, however, still poorly known. This study examines fouling communities on beached, buoyant and benthic plastic debris in the Catalan Sea (NW Mediterranean), and characterizes the plastic type. We found 14 specimens belonging to two phyla (Annelida and Foraminifera) on microplastics, and more than 400 specimens belonging to 26 species in 10 phyla (Annelida, Arthropoda, Brachiopoda, Bryozoa, Chordata, Cnidaria, Echinodermata, Mollusca, Porifera and Sipuncula) on macroplastics. With 15 species, bryozoans are the most diverse group on plastics. We also report 17 egg cases of the catshark Scyliorhinus sp., and highlight the implications for their dispersal. Our results suggest that plastic polymers may be relevant for distinct fouling communities, likely due to their chemical structure and/or surface properties. Our study provides evidence that biofouling may play a role in the sinking of plastic debris, as the most abundant fouled plastics had lower densities than seawater, and all bryozoan species were characteristic of shallower depths than those sampled. More studies at low taxonomic level are needed in order to detect new species introduction and potential invasive species associated with plastic debris.

When fossil clades 'compete': local dominance, global diversification dynamics and causation.

Examining the supposition that local-scale competition drives macroevolutionary patterns has become a familiar goal in fossil biodiversity studies. However, it is an elusive goal, hampered by inadequate confirmation of ecological equivalence and interactive processes between clades, patchy sampling, few comparative analyses of local species assemblages over long geological intervals, and a dearth of appropriate statistical tools. We address these concerns by reevaluating one of the classic examples of clade displacement in the fossil record, in which cheilostome bryozoans surpass the once dominant cyclostomes. Here, we analyse a newly expanded and vetted compilation of 40 190 fossil species occurrences to estimate cheilostome and cyclostome patterns of species proportions within assemblages, global genus richness and genus origination and extinction rates while accounting for sampling. Comparison of time-series models using linear stochastic differential equations suggests that inter-clade genus origination and extinction rates are causally linked to each other in a complex feedback relationship rather than by simple correlations or unidirectional relationships, and that these rates are not causally linked to changing within-assemblage proportions of cheilostome versus cyclostome species.

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.

Trait-fitness associations do not predict within-species phenotypic evolution over 2 million years.

Long-term patterns of phenotypic change are the cumulative results of tens of thousands to millions of years of evolution. Yet, empirical and theoretical studies of phenotypic selection are largely based on contemporary populations. The challenges in studying phenotypic evolution, in particular trait-fitness associations in the deep past, are barriers to linking micro- and macroevolution. Here, we capitalize on the unique opportunity offered by a marine colonial organism commonly preserved in the fossil record to investigate trait-fitness associations over 2 Myr. We use the density of female polymorphs in colonies of Antartothoa tongima as a proxy for fecundity, a fitness component, and investigate multivariate signals of trait-fitness associations in six time intervals on the backdrop of Pleistocene climatic shifts. We detect negative trait-fitness associations for feeding polymorph (autozooid) sizes, positive associations for autozooid shape but no particular relationship between fecundity and brood chamber size. In addition, we demonstrate that long-term trait patterns are explained by palaeoclimate (as approximated by ∂18O), and to a lesser extent by ecological interactions (i.e. overgrowth competition and substrate crowding). Our analyses show that macroevolutionary outcomes of trait evolution are not a simple scaling-up from the trait-fitness associations.

Revisiting a Landmark Study System: No Evidence for a Punctuated Mode of Evolution in Metrarabdotos.

Is speciation generally a "special time" in morphological evolution, or are lineage-splitting events just "more of the same" where the end product happens to be two separate lineages? Data on evolutionary dynamics during anagenetic and cladogenetic events among closely related lineages within a clade are rare, but the fossil record of the bryozoan genus Metrarabdotos is considered a textbook example of a clade where speciation causes rapid evolutionary change against a backdrop of morphological stasis within lineages. Here, we point to some methodological and measurement theoretical issues in the original work on Metrarabdotos. We then reanalyze a subset of the original data that can be meaningfully investigated using quantitative statistical approaches similar to those used in the original studies. We consistently fail to find variation in the evolutionary process during within-lineage evolution compared with cladogenetic events: the rates of evolution, the strength of selection, and the directions traveled in multivariate morphospace are not different when comparing evolution within lineages and at speciation events in Metrarabdotos, and genetic drift cannot be excluded as a sufficient explanation for the morphological differentiation within lineages and during speciation. Although widely considered the best example of a punctuated mode of evolution, morphological divergence and speciation are not linked in Metrarabdotos.

A genome-skimmed phylogeny of a widespread bryozoan family, Adeonidae.

BACKGROUND: Understanding the phylogenetic relationships among species is one of the main goals of systematic biology. Simultaneously, credible phylogenetic hypotheses are often the first requirement for unveiling the evolutionary history of traits and for modelling macroevolutionary processes. However, many non-model taxa have not yet been sequenced to an extent such that statistically well-supported molecular phylogenies can be constructed for these purposes. Here, we use a genome-skimming approach to extract sequence information for 15 mitochondrial and 2 ribosomal operon genes from the cheilostome bryozoan family, Adeonidae, Busk, 1884, whose current systematics is based purely on morphological traits. The members of the Adeonidae are, like all cheilostome bryozoans, benthic, colonial, marine organisms. Adeonids are also geographically widely-distributed, often locally common, and are sometimes important habitat-builders. RESULTS: We successfully genome-skimmed 35 adeonid colonies representing 6 genera (Adeona, Adeonellopsis, Bracebridgia, Adeonella, Laminopora and Cucullipora). We also contributed 16 new, circularised mitochondrial genomes to the eight previously published for cheilostome bryozoans. Using the aforementioned mitochondrial and ribosomal genes, we inferred the relationships among these 35 samples. Contrary to some previous suggestions, the Adeonidae is a robustly supported monophyletic clade. However, the genera Adeonella and Laminopora are in need of revision: Adeonella is polyphyletic and Laminopora paraphyletically forms a clade with some Adeonella species. Additionally, we assign a sequence clustering identity using cox1 barcoding region of 99% at the species and 83% at the genus level. CONCLUSIONS: We provide sequence data, obtained via genome-skimming, that greatly increases the resolution of the phylogenetic relationships within the adeonids. We present a highly-supported topology based on 17 genes and substantially increase availability of circularised cheilostome mitochondrial genomes, and highlight how we can extend our pipeline to other bryozoans.

Early Pleistocene and Holocene bryozoans from Indonesia.

This paper describes 40 bryozoan species, comprising one cyclostome and 39 cheilostomes (8 anascan- and 31 ascophoran-grade), obtained from early Pleistocene and Holocene samples from two localities in Indonesia. Five of the cheilostomes are described as new species: Acanthodesia variegata n. sp., Pleurocodonellina javanensis n. sp., Calyptotheca sidneyi n. sp., Characodoma wesselinghi n. sp., and Turbicellepora yasuharai n. sp. All of the remaining species are extant and characterized by a tropical to subtropical Indo-West Pacific distribution.

Differences in extinction rates drove modern biogeographic patterns of tropical marine biodiversity.

Marine biodiversity in the Coral Triangle is several times higher than anywhere else, but why this is true is unknown because of poor historical data. To address this, we compared the first available record of fossil cheilostome bryozoans from Indonesia with the previously sampled excellent record from the Caribbean. These two regions differ several-fold in species richness today, but cheilostome diversity was strikingly similar until the end of the Miocene 5.3 million years ago so that the modern disparity must have developed more recently. However, the Miocene faunas were ecologically very different, with a greater proportion of erect and free-living species in the Caribbean compared to the less well-known Coral Triangle. Our results support the hypothesis that modern differences in diversity arose primarily from differential extinction of Caribbean erect and free-living species concomitant with oceanographic changes due to the uplift of the Isthmus of Panama, rather than exceptional rates of diversification in the Indo-Pacific.

Relative size predicts competitive outcome through 2 million years.

Competition is an important biotic interaction that influences survival and reproduction. While competition on ecological timescales has received great attention, little is known about competition on evolutionary timescales. Do competitive abilities change over hundreds of thousands to millions of years? Can we predict competitive outcomes using phenotypic traits? How much do traits that confer competitive advantage and competitive outcomes change? Here we show, using communities of encrusting marine bryozoans spanning more than 2 million years, that size is a significant determinant of overgrowth outcomes: colonies with larger zooids tend to overgrow colonies with smaller zooids. We also detected temporally coordinated changes in average zooid sizes, suggesting that different species responded to a common external driver. Although species-specific average zooid sizes change over evolutionary timescales, species-specific competitive abilities seem relatively stable, suggesting that traits other than zooid size also control overgrowth outcomes and/or that evolutionary constraints are involved.

Interspecific interactions through 2 million years: are competitive outcomes predictable?

Ecological interactions affect the survival and reproduction of individuals. However, ecological interactions are notoriously difficult to measure in extinct populations, hindering our understanding of how the outcomes of interactions such as competition vary in time and influence long-term evolutionary changes. Here, the outcomes of spatial competition in a temporally continuous community over evolutionary timescales are presented for the first time. Our research domain is encrusting cheilostome bryozoans from the Wanganui Basin of New Zealand over a ca 2 Myr time period (Pleistocene to Recent). We find that a subset of species can be identified as consistent winners, and others as consistent losers, in the sense that they win or lose interspecific competitive encounters statistically more often than the null hypothesis of 50%. Most species do not improve or worsen in their competitive abilities through the 2 Myr period, but a minority of species are winners in some intervals and losers in others. We found that conspecifics tend to cluster spatially and interact more often than expected under a null hypothesis: most of these are stand-off interactions where the two colonies involved stopped growing at edges of encounter. Counterintuitively, competitive ability has no bearing on ecological dominance.