Convergent adaptation of true crabs (Decapoda: Brachyura) to a gradient of terrestrial environments.

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 tips spanning 88 of 109 brachyuran 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 seven and up to 17 times convergently, and returned to the sea from non-marine environments at least twice. 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 extensive new 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.

Review and guide to the isopods (Crustacea, Isopoda) of littoral and sublittoral marine habitats in the Southern California Bight.

The isopod crustaceans reported from or expected to occur in littoral and sublittoral marine habitats of the Southern California Bight (SCB) in the northeastern Pacific Ocean are reviewed. A total of 190 species, representing 105 genera in 42 families and six suborders are covered. Approximately 84% of these isopods represent described species with the remaining 16% comprising well-documented "provisional" but undescribed species. Cymothoida and Asellota are the most diverse of the six suborders, accounting for ca. 36% and 29% of the species, respectively. Valvifera and Sphaeromatidea are the next most speciose suborders with between 13-15% of the species each, while the suborder Limnorioidea represents fewer than 2% of the SCB isopod fauna. Finally, the mostly terrestrial suborder Oniscidea accounts for ca. 5% of the species treated herein, each which occurs at or above the high tide mark in intertidal habitats. A key to the suborders and superfamilies is presented followed by nine keys to the SCB species within each of the resultant groups. Figures are provided for most species. Bathymetric range, geographic distribution, type locality, habitat, body size, and a comprehensive list of references are included for most species.

A manager's guide to using eDNA metabarcoding in marine ecosystems.

Environmental DNA (eDNA) metabarcoding is a powerful tool that can enhance marine ecosystem/biodiversity monitoring programs. Here we outline five important steps managers and researchers should consider when developing eDNA monitoring program: (1) select genes and primers to target taxa; (2) assemble or develop comprehensive barcode reference databases; (3) apply rigorous site occupancy based decontamination pipelines; (4) conduct pilot studies to define spatial and temporal variance of eDNA; and (5) archive samples, extracts, and raw sequence data. We demonstrate the importance of each of these considerations using a case study of eDNA metabarcoding in the Ports of Los Angeles and Long Beach. eDNA metabarcoding approaches detected 94.1% (16/17) of species observed in paired trawl surveys while identifying an additional 55 native fishes, providing more comprehensive biodiversity inventories. Rigorous benchmarking of eDNA metabarcoding results improved ecological interpretation and confidence in species detections while providing archived genetic resources for future analyses. Well designed and validated eDNA metabarcoding approaches are ideally suited for biomonitoring applications that rely on the detection of species, including mapping invasive species fronts and endangered species habitats as well as tracking range shifts in response to climate change. Incorporating these considerations will enhance the utility and efficacy of eDNA metabarcoding for routine biomonitoring applications.

Redescription of Gnorimosphaeroma oregonense (Dana, 1853) (Crustacea, Isopoda, Sphaeromatidae), designation of neotype, and 16S-rDNA molecular phylogeny of the north-eastern Pacific species.

Gnorimosphaeroma oregonense (Dana, 1852) is revised, a male neotype is designated, photographed, and illustrated; the species occurs from Vancouver British Columbia to the central California coast. 16S-rDNA sequences (~650 bp) for all available ethanol preserved species of Gnorimosphaeroma were used to hypothesize their relationships. Our analyses revealed a sister taxon relationship between the fully marine G. oregonense and the brackish to freshwater species, G. noblei. The oyster associated and introduced G. rayi is sister to a previously not recognized or identified, but genetically distinct, Gnorimosphaeroma sp. collected at two sites in San Francisco Bay. Gnorimosphaeroma sp. is probably also a western Pacific species based on its genetic relationship to G. rayi. Photographic comparisons are offered for G. oregonense (marine), G. noblei (freshwater), G. rayi (introduced), G. sp. (presumably introduced), and G. insulare (San Nicolas Island). Records of the holdings at the Los Angeles County Museum of Natural History are summarized. Without material available north of Vancouver through Alaska, the range of G. oregonense could not be genetically verified. This review includes a diagnosis and description of the genus Gnorimosphaeroma Menzies, 1954.

Landscape analyses using eDNA metabarcoding and Earth observation predict community biodiversity in California.

Ecosystems globally are under threat from ongoing anthropogenic environmental change. Effective conservation management requires more thorough biodiversity surveys that can reveal system-level patterns and that can be applied rapidly across space and time. Using modern ecological models and community science, we integrate environmental DNA and Earth observations to produce a time snapshot of regional biodiversity patterns and provide multi-scalar community-level characterization. We collected 278 samples in spring 2017 from coastal, shrub, and lowland forest sites in California, a complex ecosystem and biodiversity hotspot. We recovered 16,118 taxonomic entries from eDNA analyses and compiled associated traditional observations and environmental data to assess how well they predicted alpha, beta, and zeta diversity. We found that local habitat classification was diagnostic of community composition and distinct communities and organisms in different kingdoms are predicted by different environmental variables. Nonetheless, gradient forest models of 915 families recovered by eDNA analysis and using BIOCLIM variables, Sentinel-2 satellite data, human impact, and topographical features as predictors, explained 35% of the variance in community turnover. Elevation, sand percentage, and photosynthetic activities (NDVI32) were the top predictors. In addition to this signal of environmental filtering, we found a positive relationship between environmentally predicted families and their numbers of biotic interactions, suggesting environmental change could have a disproportionate effect on community networks. Together, these analyses show that coupling eDNA with environmental predictors including remote sensing data has capacity to test proposed Essential Biodiversity Variables and create new landscape biodiversity baselines that span the tree of life.

Worldwide Engagement for Digitizing Biocollections (WeDigBio): The Biocollections Community's Citizen-Science Space on the Calendar.

The digitization of biocollections is a critical task with direct implications for the global community who use the data for research and education. Recent innovations to involve citizen scientists in digitization increase awareness of the value of biodiversity specimens; advance science, technology, engineering, and math literacy; and build sustainability for digitization. In support of these activities, we launched the first global citizen-science event focused on the digitization of biodiversity specimens: Worldwide Engagement for Digitizing Biocollections (WeDigBio). During the inaugural 2015 event, 21 sites hosted events where citizen scientists transcribed specimen labels via online platforms (DigiVol, Les Herbonautes, Notes from Nature, the Smithsonian Institution's Transcription Center, and Symbiota). Many citizen scientists also contributed off-site. In total, thousands of citizen scientists around the world completed over 50,000 transcription tasks. Here, we present the process of organizing an international citizen-science event, an analysis of the event's effectiveness, and future directions-content now foundational to the growing WeDigBio event.

Redescription of Dynoides elegans (Boone, 1923) (Crustacea, Isopoda, Sphaeromatidae) from the north-eastern Pacific.

Dynoides elegans (Boone, 1923) from southern California is reviewed, redescribed, and figured. The original species description did not include figures, making it difficult to attribute individuals to the species. Dynoides saldanai Carvacho and Haasmann, 1984 and Dynoides crenulatus Carvacho & Haasman, 1984 from the Pacific Coast of Mexico and Dynoides brevicornis Kussakin & Malyutina, 1987, from Furugelm Island, Peter the Great Gulf in the Sea of Japan, appear morphologically more similar to each other than to western Pacific species. A large pleonal process is present in about half of the Dynoides species, but is absent in this north-eastern Pacific clade and the north-western Pacific Dynoides brevicornis and Dynoides brevispina. Dynoides dentisinus Shen, 1929 possess a large pleonal spine. It is known from China, Japan, and Korea and is introduced in San Francisco Bay; it can be easily distinguished from Dynoides elegans by the presence of a pleonal process in the former. A key to the Pacific West Coast Dynoides is provided.

Collecting and preserving marine and freshwater isopoda (crustacea: peracarida).

BACKGROUND: Isopoda are the most diverse Crustacea. In order to encourage the study of isopod crustaceans and their use in biodiversity studies, systematics, ecology, physiology and more, one needs to know who the isopods are and where to find them. NEW INFORMATION: This is a short "how to" guide focusing on the free-living marine and freshwater isopods: where they live and how to collect and preserve them. The tools and techniques described here are simple, but invaluable in accessing the natural history of these remarkable creatures.

Status of Exosphaeromaamplicauda (Stimpson, 1857), E.aphrodita (Boone, 1923) and description of three new species (Crustacea, Isopoda, Sphaeromatidae) from the north-eastern Pacific.

Exosphaeromaamplicauda (Stimpson, 1857) from the west coast of North America is reviewed and redescribed and revealed to be a group of closely related species. A neotype is designated and the species redescribed based on the neotype and topotypic specimens. Exosphaeromaamplicauda is known only from the coast of California, at Marin, Sonoma and San Mateo Counties. Exosphaeromaaphrodita (Boone, 1923), type locality La Jolla, California and previously considered nomen dubium is taken out of synonymy and re-validated. A further three species: Exosphaeromapaydenae sp. n., Exosphaeromarussellhansoni sp. n., and Exosphaeromapentcheffi sp. n. are described herein. Sphaeromaoctonctum Richardson, 1899 is placed into junior synonymy with Exosphaeromaamplicauda. A key to the Pacific West Coast Exosphaeroma is provided.

Genetic utility of natural history museum specimens: endangered fairy shrimp (Branchiopoda, Anostraca).

We examined the potential utility of museum specimens as a source for genetic analysis of fairy shrimp. Because of loss of their vernal pool habitat, some fairy shrimp (including Branchinectasandiegonensis and Branchinectalynchi) are listed as threatened or endangered in Southern California by the United States Fish and Wildlife Service. Management of those species requires extensive population genetics studies and the resolution of important genetic complexity (e.g. possible hybridization between endangered and non-endangered species). Regulations mandating deposition of specimens of listed species have resulted in thousands of specimens accessioned into the Natural History Museum of Los Angeles County that have been preserved in a variety of solutions. We subsampled those specimens, as well as other Anostraca with known collection and preservation histories, to test their potential for genetic analysis by attempting DNA extraction and amplification for mt16SrDNA. Fixation and preservation in not denatured ethanol had a far greater sequencing success rate than other (and unknown) fixatives and preservatives. To maximize scientific value we recommend field preservation in 95% not denatured ethanol (or, if pure ethanol is unavailable, high-proof drinking spirits, e.g. Everclear™, or 151 proof white rum), followed by storage in 95% not denatured ethanol.

Phylogenetic relationships of the family Sphaeromatidae Latreille, 1825 (Crustacea: Peracarida: Isopoda) within Sphaeromatidea based on 18S-rDNA molecular data.

Based on 18S-rDNA sequences of 97 isopods including 18 Sphaeromatidea, we show Sphaeromatidae, Valvifera, Serolidae, and Ancinidae is a well supported clade. The within clade relationships of these taxa are not as definitively demonstrated because taxon sampling for some groups is still limited. In our analyses the Sphaeromatidae are shown to be unequivocally monophyletic. This is contrary to the morphology-based analysis by A. Brandt and G. Poore in 2003, which included only five Sphaeromatidae and found the family to be paraphyletic. The Ancinidae are also upheld, and the Valvifera is the sister taxon to Serolidae. Surprisingly Plakarthrium (Plakarthiidae) is nested within the Sphaeromatidae in most analyses. We point out short-comings in our sampling and suggest areas which would benefit from better sampling. We also review the long and convoluted nomenclatural history of the Sphaeromatidea, Sphaeromatoidea, and
Sphaeromatidae.

Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences.

The remarkable antiquity, diversity and ecological significance of arthropods have inspired numerous attempts to resolve their deep phylogenetic history, but the results of two decades of intensive molecular phylogenetics have been mixed. The discovery that terrestrial insects (Hexapoda) are more closely related to aquatic Crustacea than to the terrestrial centipedes and millipedes (Myriapoda) was an early, if exceptional, success. More typically, analyses based on limited samples of taxa and genes have generated results that are inconsistent, weakly supported and highly sensitive to analytical conditions. Here we present strongly supported results from likelihood, Bayesian and parsimony analyses of over 41 kilobases of aligned DNA sequence from 62 single-copy nuclear protein-coding genes from 75 arthropod species. These species represent every major arthropod lineage, plus five species of tardigrades and onychophorans as outgroups. Our results strongly support Pancrustacea (Hexapoda plus Crustacea) but also strongly favour the traditional morphology-based Mandibulata (Myriapoda plus Pancrustacea) over the molecule-based Paradoxopoda (Myriapoda plus Chelicerata). In addition to Hexapoda, Pancrustacea includes three major extant lineages of 'crustaceans', each spanning a significant range of morphological disparity. These are Oligostraca (ostracods, mystacocarids, branchiurans and pentastomids), Vericrustacea (malacostracans, thecostracans, copepods and branchiopods) and Xenocarida (cephalocarids and remipedes). Finally, within Pancrustacea we identify Xenocarida as the long-sought sister group to the Hexapoda, a result confirming that 'crustaceans' are not monophyletic. These results provide a statistically well-supported phylogenetic framework for the largest animal phylum and represent a step towards ending the often-heated, century-long debate on arthropod relationships.

Phylogenetic relationships within the coral crab genus Carpilius (Brachyura, Xanthoidea, Carpiliidae) and of the Carpiliidae to other xanthoid crab families based on molecular sequence data.

The coral crab genus Carpilius currently includes three widely distributed species that inhabit tropical coral reefs and adjacent waters. The relationship of Carpilius to other xanthoid crabs is unknown. Previously, carcinologists considered Carpilius to be allied with crabs of the family Xanthidae (e.g., Euryozius, Liagore, and Liomera), however, recent workers have considered it to be a monotypic genus within its own family, Carpiliidae. Mitochondrial 12S- and 16S-rDNA gene fragments confirm the monophyly and distinct status of the family Carpiliidae. Within the genus Carpilius, the Caribbean species C. corallinus is basal to the two Pacific species C. maculatus and C. convexus. The Pacific species are sister taxa, despite the greater morphological resemblance of C. corallinus to the Pacific C. convexus. The relationship of the Carpiliidae (Carpilius) to other xanthoid crabs is investigated, and results of a preliminary analysis of higher xanthoid relationships did not resolve the relationships of Carpiliidae, "Xanthidae," Menippidae, Trapeziidae, and Ocypodidae to one another. A Menippidae and Carpilius relationship could not be rejected, although a Liomera, Liagore, and Carpilius relationship was.