For much of terrestrial biodiversity, the evolutionary pathways of adaptation from marine ancestors are poorly understood, and have usually been viewed as a binary trait. True crabs, the decapod crustacean infraorder Brachyura, comprise over 7,600 species representing a striking diversity of morphology and ecology, including repeated adaptation to non-marine habitats. Here, we reconstruct the evolutionary history of Brachyura using new and published sequences of 10 genes for 344 species spanning 88 of 104 families. Using 36 newly vetted fossil calibrations, we infer that brachyurans most likely diverged in the Triassic, with family-level splits in the late Cretaceous and early Paleogene. By contrast, the root age is underestimated with automated sampling of 328 fossil occurrences explicitly incorporated into the tree prior, suggesting such models are a poor fit under heterogeneous fossil preservation. We apply recently defined trait-by-environment associations to classify a gradient of transitions from marine to terrestrial lifestyles. We estimate that crabs left the marine environment at least five and up to 15 times convergently, and returned to the sea from non-marine environments three or four times. Although the most highly terrestrial- and many freshwater-adapted crabs are concentrated in Thoracotremata, Bayesian threshold models of ancestral state reconstruction fail to identify shifts to higher terrestrial grades due to the degree of underlying change required. Lineages throughout our tree inhabit intertidal and marginal marine environments, corroborating the inference that the early stages of terrestrial adaptation have a lower threshold to evolve. Our framework and newly compiled fossil and natural history datasets will enable future comparisons of non-marine adaptation at the morphological and molecular level. Crabs provide an important window into the early processes of adaptation to novel environments, and different degrees of evolutionary constraint that might help predict these pathways.
Lying at the southernmost point of the Lucayan Archipelago, the Turks and Caicos Islands are amongst the better studied localities for anchialine cave biodiversity. For nearly five decades, novel invertebrate fauna, comprised primarily of crustaceans, have been collected from these tidally influenced pools – but new findings are always on the horizon. Herein we present new records of crustaceans and annelids from anchialine blue holes and horizontal caves of the Turks and Caicos. These findings include two potentially new species of meiofaunal annelids and a new species of remipede collected from a shallow water cave pool. Our 2019 expedition additionally expands known faunal distributions for several taxa across the Caicos islands, and raises the biodiversity of the region to 35 species, 13 of them considered endemic. This is the first comprehensive faunal list for the anchialine systems in the Caicos Bank.
The Yucatán Peninsula, Mexico is a carbonate platform well-known for extensive karst networks of densely stratified aquifer ecosystems. This aquifer supports diverse anchialine fauna, including species of the globally distributed anchialine shrimp genus Typhlatya (Atyidae). Four species (T. campecheae, T. pearsei, T. dzilamensis and T. mitchelli) are endemic to the Peninsula, of which three are federally listed in Mexico. This first integrative evaluation (i.e., molecular, morphological, broad geographic and type locality sampling, and environmental data) of Yucatán Typhlatya reveals considerable species identity conflict in prior phylogenetic assessments, broad species ranges, syntopy within cave systems and five genetic lineages (of which two are new to science). Despite sampling from the type locality of endangered T. campecheae, specimens (and molecular data) were indistinguishable from vulnerable T. pearsei. Ancestral/divergence reconstructions support convergent evolution of a low-salinity ancestor for a post-Paleogene arc Yucatán + Cuba Typhlatya clade within the anchialine Atyidae clade. A secondary adaptation for the coastal-restricted euryhaline (2–37 psu), Typhlatya dzilamensis (unknown conservation status) was identified, while remaining species lineages were low-salinity (< 5 psu) adapted and found within the meteoric lens of inland and coastal caves. This study demonstrates the need for integrative/interdisciplinary approaches when conducting biodiversity assessments in complex and poorly studied aquifers.
Remipedia is a stygobitic group commonly associated with coastal anchialine caves. This class consists of 12 genera, ten of which are found within the Lucayan Archipelago. Herein, we describe a new species within the genus Godzillius from Conch Sound Blue Hole, North Andros Island, Bahamas. Godzillius louriei sp. nov. is the third known remipede observed from a subseafloor marine cave, and the first from the Godzilliidae. Remipedes dwell within notoriously difficult to access cave habitats and thus integrative and comprehensive systematic studies at family or genus level are often absent in the literature. In this study, all species of Godzillius are compared using morphological and molecular approaches. Specifically, the feeding appendages of G. louriei sp. nov., G. fuchsi Gonzalez, Singpiel & Schlagner, 2013 and G. robustus Schram, Yager & Emerson, 1986 were examined using scanning electron microscopy (SEM). Species of Godzillius are identified based on the spines of maxilla 1 segment 4 and by the denticles on the lacinia mobilis of the left mandible. A molecular phylogeny using the mitochondrial 16S rRNA and nuclear histone 3 genes recovered G. louriei sp. nov. within the Godzillius clade and 16S genetic distances revealed a 13–15% difference between species of Godzillius.
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