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Chelicerates are a diverse group of arthropods, represented by such forms as predatory spiders and scorpions, parasitic ticks, humic detritivores, and marine sea spiders (pycnogonids) and horseshoe crabs. Conflicting phylogenetic relationships have been proposed for chelicerates based on both morphological and molecular data, the latter usually not recovering arachnids as a clade and instead finding horseshoe crabs nested inside terrestrial Arachnida. Here, using genomic-scale datasets and analyses optimised for countering systematic error, we find strong support for monophyletic Acari (ticks and mites), which when considered as a single group represent the most biodiverse chelicerate lineage. In addition, our analysis recovers marine forms (sea spiders and horseshoe crabs) as the successive sister groups of a monophyletic lineage of terrestrial arachnids, suggesting a single colonisation of land within Chelicerata and the absence of wholly secondarily marine arachnid orders. Morphological and molecular data have led to conflicting phylogenetic hypotheses for the Chelicerata. Here, the authors reconstruct the phylogeny of the Chelicerata using genomic-scale datasets, finding evidence for a monophyletic Acari and a single terrestrialisation of Arachnida.

Despite significant advances in invertebrate phylogenomics over the past decade, the higher-level phylogeny of Pycnogonida (sea spiders) remains elusive. Due to the inaccessibility of some small-bodied lineages, few phylogenetic studies have sampled all sea spider families. Previous efforts based on a handful of genes have yielded unstable tree topologies. Here, we inferred the relationships of 89 sea spider species using targeted capture of the mitochondrial genome, 56 conserved exons, 101 ultraconserved elements, and 3 nuclear ribosomal genes. We inferred molecular divergence times by integrating morphological data for fossil species to calibrate 15 nodes in the arthropod tree of life. This integration of data classes resolved the basal topology of sea spiders with high support. The enigmatic family Austrodecidae was resolved as the sister group to the remaining Pycnogonida and the small-bodied family Rhynchothoracidae as the sister group of the robust-bodied family Pycnogonidae. Molecular divergence time estimation recovered a basal divergence of crown group sea spiders in the Ordovician. Comparison of diversification dynamics with other marine invertebrate taxa that originated in the Paleozoic suggests that sea spiders and some crustacean groups exhibit resilience to mass extinction episodes, relative to mollusk and echinoderm lineages.

Regenerative abilities and their evolution in the different animal lineages have fascinated generations of biologists. While some taxa are capable of restoring entire individuals from small body fragments, others can regrow only specific structures or lack structural regeneration completely. In contrast to many other protostomes, including the segmented annelids, molting animals (Ecdysozoa) are commonly considered incapable of primary body axis regeneration, which has been hypothesized to be linked to the evolution of their protective cuticular exoskeleton. This holds also for the extraordinarily diverse, segmented arthropods. Contradicting this long-standing paradigm, we here show that immatures of the sea spider Pycnogonum litorale reestablish the posterior body pole after transverse amputation and can regrow almost complete segments and the terminal body region, including the hindgut, anus, and musculature. Depending on the amputation level, normal phenotypes or hypomeric six-legged forms develop. Remarkably, also the hypomeric animals regain reproductive functionality by ectopic formation of gonoducts and gonopores. The discovery of such complex regenerative patterns in an extant arthropod challenges the hitherto widely assumed evolutionary loss of axial regeneration during ecdysozoan evolution. Rather, the branching of sea spiders at the base of Chelicerata and their likely ancestral anamorphic development suggests that the arthropod stem species may have featured similar regenerative capabilities. Accordingly, our results provide an incentive for renewed comparative regeneration studies across ecdysozoans, with the aim to resolve whether this trait was potentially even inherited from the protostome ancestor.

To investigate whether the biogeographical regions proposed by J. W. Hedgpeth and widely adopted by other authors hold true, are an oversimplification or with further data might show a unified Antarctic province. Southern Hemisphere. The distributions of 1318 species of bivalves, 4656 species of gastropods, 1465 species of cheilostome and 167 species of cyclostome bryozoans were analysed for 29 regions in the Southern Hemisphere, including South American, South African, Tasmanian, New Zealand, sub-Antarctic and Antarctic regions. We present data on species richness, rates of endemism, patterns of radiation, faunal similarities and multivariate biogeographical analyses. The most striking pattern to emerge from our data set of species counts per region was a strong east-west hemispheric asymmetry, with high species numbers in New Zealand, Tasmania and South Africa and low numbers in South America. In contrast, no difference was found in richness between the east and west parts of the Southern Ocean. We compared findings in our model taxa with published data on ascidians, cephalopods and pycnogonids. Further evidence of strong faunal links between the Antarctic and South America is reported in this study, although we found little evidence for a biogeographical relationship between the Antarctic or South America and New Zealand/Tasmania. Strong evidence exists for a long-term influence of the Antarctic Circumpolar Current upon the distribution of Southern Ocean benthos. This is demonstrated by the reduced prevalence of South American species in the Antarctic and sub-Antarctic with increasing distance from South America in the direction of the current. Three of our four study taxa (bivalves, cheilostomes and cyclostomes) show the Southern Ocean as a 'single functional unit' with no evidence for a biogeographical split between east and west. Unlike the biogeographical schemes previously proposed, we show that biogeographical regions in the Southern Ocean differ depending upon the class of animals being considered. Despite this we suggest that some general rules are viable, including species endemism rates of around 50%, a single Antarctic province and a definite distinction between the sub-Antarctic islands influenced by South America and those of New Zealand.

Neofunctionalization of duplicated gene copies is thought to be an important process underlying the origin of evolutionary novelty and provides an elegant mechanism for the origin of new phenotypic traits. One putative case where a new gene copy has been linked to a novel morphological trait is the origin of the arachnid patella, a taxonomically restricted leg segment. In spiders, the origin of this segment has been linked to the origin of the paralog dachshund-2, suggesting that a new gene facilitated the expression of a new trait. However, various arachnid groups that possess patellae do not have a copy of dachshund-2, disfavoring the direct link between gene origin and trait origin. We investigated the developmental genetic basis for patellar patterning in the harvestman Phalangium opilio, which lacks dachshund-2. Here, we show that the harvestman patella is established by a novel expression domain of the transcription factor extradenticle. Leveraging this definition of patellar identity, we surveyed targeted groups across chelicerate phylogeny to assess when this trait evolved. We show that a patellar homolog is present in Pycnogonida (sea spiders) and various arachnid orders, suggesting a single origin of the patella in the ancestor of Chelicerata. A potential loss of the patella is observed in Ixodida. Our results suggest that the modification of an ancient gene, rather than the neofunctionalization of a new gene copy, underlies the origin of the patella. Broadly, this work underscores the value of comparative data and broad taxonomic sampling when testing hypotheses in evolutionary developmental biology.

A large bloom of Salpa spp. in the northeastern Pacific during the spring of 2012 resulted in a major deposition of tunics and fecal pellets on the seafloor at ~ 4000 m depth (Sta. M) over a period of 6 months. Continuous monitoring of this food pulse was recorded using autonomous instruments: sequencing sediment traps, a timelapse camera on the seafloor, and a bottom-transiting vehicle measuring sediment community oxygen consumption (SCOC). These deep-sea measurements were complemented by sampling of salps in the epipelagic zone by California Cooperative Ocean Fisheries Investigations. The particulate organic carbon (POC) flux increased sharply beginning in early March, reaching a peak of 38 mg C m−2 d−1 in mid-April at 3400 m depth. Salp detritus started appearing in images of the seafloor taken in March and covered a daily maximum of 98% of the seafloor from late June to early July. Concurrently, the SCOC rose with increased salp deposition, reaching a high of 31 mg C m−2 d−1 in late June. A dominant megafauna species, Peniagone sp. A, increased 7-fold in density beginning 7 weeks after the peak in salp deposition. Estimated food supply from salp detritus was 97–327% of the SCOC demand integrated over the 6-month period starting in March 2012. Such large episodic pulses of food sustain abyssal communities over extended periods of time.

Glacio-marine fjords occur widely at high latitudes and have been extensively studied in the Arctic, where heavy meltwater inputs and sedimentation yield low benthic faunal abundance and biodiversity in inner-middle fjords. Fjord benthic ecosystems remain poorly studied in the subpolar Antarctic, including those in extensive fjords along the West Antarctic Peninsula (WAP). Here we test ecosystem predictions from Arctic fjords on three subpolar, glacio-marine fjords along the WAP. With seafloor photographic surveys we evaluate benthic megafaunal abundance, community structure, and species diversity, as well as the abundance of demersal nekton and macroalgal detritus, in soft-sediment basins of Andvord, Flandres and Barilari Bays at depths of 436-725 m. We then contrast these fjord sites with three open shelf stations of similar depths. Contrary to Arctic predictions, WAP fjord basins exhibited 3 to 38-fold greater benthic megafaunal abundance than the open shelf, and local species diversity and trophic complexity remained high from outer to inner fjord basins. Furthermore, WAP fjords contained distinct species composition, substantially contributing to beta and gamma diversity at 400-700 m depths along the WAP. The abundance of demersal nekton and macroalgal detritus was also substantially higher in WAP fjords compared to the open shelf. We conclude that WAP fjords are important hotspots of benthic abundance and biodiversity as a consequence of weak meltwater influences, low sedimentation disturbance, and high, varied food inputs. We postulate that WAP fjords differ markedly from their Arctic counterparts because they are in earlier stages of climate warming, and that rapid warming along the WAP will increase meltwater and sediment inputs, deleteriously impacting these biodiversity hotspots. Because WAP fjords also provide important habitat and foraging areas for Antarctic krill and baleen whales, there is an urgent need to develop better understanding of the structure, dynamics and climate-sensitivity of WAP subpolar fjord ecosystems.

Pycnogonid specimens of the genus Pallenopsis collected during the Antarctic Polarstern XXIII/8 cruise (23 November 2006 to 30 January 2007) were studied. Seven species are recognized in this collection and two of them are proposed as new species: Pallenopsis tubercula n. sp., P. elephantensis n. sp., P. gracilis Cano-Sánchez and López-González, 2019Cano-SánchezE., López-GonzálezP.J.2019. Two new species and new findings in the genus Pallenopsis (Pycnogonida: Pallenopsidae) with an updated identification key to Antarctic and Sub-Antarctic species. Zootaxa4585: 517-530. 10.11646/zootaxa.4585.3.7, P. hodgsoni Gordon, 1938, P. macronyx Bouvier, 1911, P. spicata Hodgson, 1914 and P. aff. vanhoeffeni Hodgson, 1915. Pallenopsis gracilis is reported for the first time since its original description. The new species are described, illustrated and compared with their closest congeners, and their morphological variability is discussed. A cluster analysis (Euclidean distance) including our Pallenopsis specimens and the holotype of P. gracilis is carried out, based on 24 morphological characters. Finally, an updated key to the Southern Ocean species is presented.

A new sea spider (Arthropoda: Pycnogonida) from the Lower Devonian Hunsrueck Slate (Germany) is described as Pentapantopus vogteli gen. et sp. nov. This is the fifth pycnogonid species known from this exceptional fossil Lagerstaette. The most conspicuous character of the new species is the presence of five pairs of walking legs. This character, in concert with a reduced abdomen, indicates a phylogenetic position of P. vogteli among the crown group pycnogonids. P. vogteli extends the knowledge of fossil pycnogonid body plans and underlines the significance of the Hunsrueck Slate, as this locality shows the highest diversity of sea spiders for the entire fossil record so far.

Neuroectodermal signalling centres induce and pattern many novel vertebrate brain structures but are absent, or divergent, in invertebrate chordates. This has led to the idea that signalling-centre genetic programs were first assembled in stem vertebrates and potentially drove morphological innovations of the brain. However, this scenario presumes that extant cephalochordates accurately represent ancestral chordate characters, which has not been tested using close chordate outgroups. Here we report that genetic programs homologous to three vertebrate signalling centres — the anterior neural ridge, zona limitans intrathalamica and isthmic organizer — are present in the hemichordate Saccoglossus kowalevskii . Fgf8/17/18 (a single gene homologous to vertebrate Fgf8 , Fgf17 and Fgf18 ), sfrp1/5 , hh and wnt1 are expressed in vertebrate-like arrangements in hemichordate ectoderm, and homologous genetic mechanisms regulate ectodermal patterning in both animals. We propose that these genetic programs were components of an unexpectedly complex, ancient genetic regulatory scaffold for deuterostome body patterning that degenerated in amphioxus and ascidians, but was retained to pattern divergent structures in hemichordates and vertebrates. Genetic programs homologous to three vertebrate signalling centres are present in the hemichordate Saccoglossus kowalevskii and may be components of a complex, ancient genetic regulatory scaffold for deuterostome body patterning that degenerated in amphioxus and ascidians, but was retained to pattern divergent structures in hemichordates and vertebrates. Evolutionary roots of the vertebrate brain The vertebrate brain is a complex structure, and how it evolved from a simpler nervous system remains obscure. The invertebrates most closely related to vertebrates, such as sea squirts and lancelets, have very much simpler brains, and it has been widely assumed that the vertebrate brain has a uniquely vertebrate evolutionary history. But work by Christopher Lowe and colleagues now shows that the genetic program that specifies the anterior end of acorn worms — very distant relatives of vertebrates, akin to echinoderms such as starfish — is very like that of vertebrates. This means that the program for specifying the vertebrate brain started out as a more generalized routine for the development of the front end of the animal. Confusion arose because the pathways involved have been lost or highly modified in lancelets and sea squirts.