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The authors declare that all data supporting the findings of this study are available within the article and its Supplementary Information files. Abstract Climate is a strong driver of global diversity and will become increasingly important as human influences drive temperature changes at unprecedented rates. Here we investigate diversification and speciation trends within a diverse group of aquatic crustaceans, the Anomura. We use a phylogenetic framework to demonstrate that speciation rate is correlated with global cooling across the entire tree, in contrast to previous studies.

Additionally, we find that marine clades continue to show evidence of increased speciation rates with cooler global temperatures, while the single freshwater clade shows the opposite trend with speciation rates positively correlated to global warming. Our findings suggest that both global cooling and warming lead to diversification and that habitat plays a role in the responses of species to climate change.

These results have important implications for our understanding of how extant biota respond to ongoing climate change and are of particular importance for conservation planning of marine ecosystems.

Changes in climate, particularly global mean temperatures, have long been identified as drivers of diversity turnover on macroevolutionary scales 1 , 2 , 3 , 4. However, to predict the probable effects of current and future climate change it is vital that we understand past processes and trends.

Of particular concern is the threat to our oceans and freshwater ecosystems 7. However, while the responses of shelf marine invertebrates, as a whole, to a variety of climatic and other factors has been investigated at a taxic level, this investigation has yielded contradictory results; some studies found that warming induced increased diversification, while others found the opposite effect 10 , 11 , By contrast, more focussed studies on marine molluscs over shorter time scales have found no relationship between diversification and environmental factors.

Rather, species either track habitat 13 or nutrient availability 14 , or they go extinct. Terrestrial vertebrates, on the other hand, appear to show a positive correlation between speciation and global temperature 15 , 16 , 17 , while marine vertebrates show either a positive correlation with warming or no correlation with climate 16 , 17 , To date, no work has been carried out on the relationship between speciation and climate of marine invertebrates within a phylogenetic framework.

Here we examine the relationship between climate and biological diversification for Anomura hermit crabs, king crabs, porcelain crabs, mole crabs and squat lobsters.

Critically, we do this in a phylogenetic framework, using the largest tree ever assembled for the group. Anomura are an enigmatic infraorder, containing species with a diverse array of ecologies and encompassing a particularly impressive disparity of forms They are also remarkable in having independently evolved carcinisation in multiple lineages Anomura are united by the general synapomorphy of a reduced anterior-most pair of pereiopods used in gill cleaning, and the absence or severe reduction of the tail.

To date, 2, extant species have been described 21 , Anomura have colonized deep marine from depths of over 5, m to littoral environments, and are also found in freshwater and semi-terrestrial habitats. Historical taxonomic controversies are now largely resolved 19 , therefore their evolution provides an excellent system in which to investigate the relationship between climate and speciation in aquatic invertebrates.

Previous studies investigating the link between climate and biological diversification have largely used a taxic approach, treating species, genera and families as independent and assessing changes in taxonomic diversity through time. Using a time-calibrated phylogenetic supertree of Anomura, which broadly matches a recent molecular phylogeny and is consistent with recent taxonomic revisions, we calculate diversification rates for the history of Anomura.

We then perform correlation analyses of palaeo-temperature proxies, and these diversification rates using a Bayesian framework. Our analyses show a negative correlation between the whole tree speciation rates and palaeo-temperature. In detail, marine taxa show the same negative correlation, but the single freshwater clade shows the opposite relationship: These results suggest that habitat plays a role in how speciation rates are affected by temperature changes, which in turn has implications for how we manage and conserve extant aquatic biota.

Results Supertree construction We constructed a phylogenetic supertree 24 , 25 using 60 source trees from 40 papers published between and The resulting tree of taxa is the largest phylogeny of Anomura published to date Fig. The overall structure of the supertree is broadly similar to that in a recent molecular phylogeny taxa of Anomura 19 , and is also consistent with recent taxonomic revisions 26 , notably a much reduced Galatheoidea Supplementary Fig.

In our tree all superfamilies are recovered as monophyletic with the exception of the Paguroidea. The resolution of Lithodoidea within Paguroidea is well-supported by both molecular and morphological data The supertree was time-calibrated using fossil age data. Three additional trees were generated using random perturbations of the fossil age data to simulate changes to node calibrations that could affect the robustness of our findings.

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This course is designed for computer science, computer engineering, electrical engineering, and mathematics students interested in understanding, designing, developing, analyzing, and validating elliptic cryptographic algorithms and protocols. The course is targeted to a diverse audience, and generally assumes no more than an undergraduate degree in computer science, electrical or computer engineering, or mathematics.

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Anomura are an enigmatic infraorder, containing species with a diverse array of ecologies and encompassing a particularly impressive disparity of forms They are also remarkable in having independently evolved carcinisation in multiple lineages Anomura are united by the general synapomorphy of a reduced anterior-most pair of pereiopods used in gill cleaning, and the absence or severe reduction of the tail. To date, 2, extant species have been described 21 , Anomura have colonized deep marine from depths of over 5, m to littoral environments, and are also found in freshwater and semi-terrestrial habitats.

Historical taxonomic controversies are now largely resolved 19 , therefore their evolution provides an excellent system in which to investigate the relationship between climate and speciation in aquatic invertebrates. Previous studies investigating the link between climate and biological diversification have largely used a taxic approach, treating species, genera and families as independent and assessing changes in taxonomic diversity through time.

Using a time-calibrated phylogenetic supertree of Anomura, which broadly matches a recent molecular phylogeny and is consistent with recent taxonomic revisions, we calculate diversification rates for the history of Anomura. We then perform correlation analyses of palaeo-temperature proxies, and these diversification rates using a Bayesian framework. Our analyses show a negative correlation between the whole tree speciation rates and palaeo-temperature.

In detail, marine taxa show the same negative correlation, but the single freshwater clade shows the opposite relationship: These results suggest that habitat plays a role in how speciation rates are affected by temperature changes, which in turn has implications for how we manage and conserve extant aquatic biota.

Results Supertree construction We constructed a phylogenetic supertree 24 , 25 using 60 source trees from 40 papers published between and The resulting tree of taxa is the largest phylogeny of Anomura published to date Fig. The overall structure of the supertree is broadly similar to that in a recent molecular phylogeny taxa of Anomura 19 , and is also consistent with recent taxonomic revisions 26 , notably a much reduced Galatheoidea Supplementary Fig.

In our tree all superfamilies are recovered as monophyletic with the exception of the Paguroidea. The resolution of Lithodoidea within Paguroidea is well-supported by both molecular and morphological data

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