Neglected no longer: Phylogenomic resolution of higher-level relationships in Solifugae.

Advanced sequencing technologies have expedited resolution of higher-level arthropod relationships. Yet, dark branches persist, principally among groups occurring in cryptic habitats. Among chelicerates, Solifugae ("camel spiders") is the last order lacking a higher-level phylogeny and have thus been historically characterized as "neglected [arachnid] cousins". Though renowned for aggression, remarkable running speed, and xeric adaptation, inferring solifuge relationships has been hindered by inaccessibility of diagnostic morphological characters, whereas molecular investigations have been limited to one of 12 recognized families. Our phylogenomic dataset via capture of ultraconserved elements sampling all extant families recovered a well-resolved phylogeny, with two distinct groups of New World taxa nested within a broader Paleotropical radiation. Divergence times using fossil calibrations inferred that Solifugae radiated by the Permian, and most families diverged prior to the Paleogene-Cretaceous extinction, likely driven by continental breakup. We establish Boreosolifugae new suborder uniting five Laurasian families, and Australosolifugae new suborder uniting seven Gondwanan families using morphological and biogeographic signal.

Congruence between ultraconserved element-based matrices and phylotranscriptomic datasets in the scorpion Tree of Life.

Scorpions are ancient and historically renowned for their potent venom. Traditionally, the systematics of this group of arthropods was supported by morphological characters, until recent phylogenomic analyses (using RNAseq data) revealed most of the higher-level taxa to be non-monophyletic. While these phylogenomic hypotheses are stable for almost all lineages, some nodes have been hard to resolve due to minimal taxonomic sampling (e.g. family Chactidae). In the same line, it has been shown that some nodes in the Arachnid Tree of Life show disagreement between hypotheses generated using transcritptomes and other genomic sources such as the ultraconserved elements (UCEs). Here, we compared the phylogenetic signal of transcriptomes vs. UCEs by retrieving UCEs from new and previously published scorpion transcriptomes and genomes, and reconstructed phylogenies using both datasets independently. We reexamined the monophyly and phylogenetic placement of Chactidae, sampling an additional chactid species using both datasets. Our results showed that both sets of genome-scale datasets recovered highly similar topologies, with Chactidae rendered paraphyletic owing to the placement of Nullibrotheas allenii. As a first step toward redressing the systematics of Chactidae, we establish the family Anuroctonidae (new family) to accommodate the genus Anuroctonus.

Old and cold: Diverse phylogenomic datasets support an ancient transantarctic dispersive route on the scorpion family Bothriuridae in temperate Gondwana.

In this contribution we try to unveil the diversification process of Bothriuridae in temperate Gondwana through dated phylogenomic analyses using UCE and transcriptomics, and including in the analyses species of genera Urophonius and Cercophonius, the most closely related genera of Bothriuridae from South America and Australia respectively. Additionally we explored the hypothesis that the winter activity period of some species of Urophonius, as well as the cold environmental preferences of this genus, could be related to the climatic conditions of the time frame and area in which it evolved. Genus Urophonius was recovered as sister group to Cercophonius using amino acids and UCE. The time frame obtained for the split between South American and Australian bothriurids is 94 Ma., which suggests a dispersal event through temperate Gondwana, before the final breakup of the land bridge of South America-Antarctica-Australia ca. 35 Ma. The split between summer and winter species of Urophonius, taking place at 64 Ma, is considered representative to the turnover time from the summer activity period to the winter activity period in some species of the genus. This time frame is compatible with a period of global warming of the late Cretaceous greenhouse episode that could have triggered this change.

Description of Bothriurus mistral n. sp., the highest-dwelling Bothriurus from the western Andes (Scorpiones, Bothriuridae), using multiple morphometric approaches.

We describe Bothriurus mistral n. sp. (Scorpiones, Bothriuridae) from the Chilean north-central Andes of the Coquimbo Region. This is the highest elevational discovery for Bothriurus in the western slopes of the Andes. This species was collected in the Estero Derecho Private Protected Area and Natural Sanctuary as part of the First National Biodiversity Inventory of Chile of the Integrated System for Monitoring and Evaluation of Native Forest Ecosystems (SIMEF). Bothriurus mistral n. sp. is closely related to Bothriurus coriaceus Pocock, 1893, from the lowlands of central Chile. This integrative research includes a combination of traditional morphometrics and geometric morphometric analyses to support the taxonomic delimitation of the species.

A Fine-Scale Hotspot at the Edge: Epigean Arthropods from the Atacama Coast (Paposo-Taltal, Antofagasta Region, Chile).

The Atacama Desert at its margins harbors a unique biodiversity that is still very poorly known, especially in coastal fog oases spanning from Perú towards the Atacama coast. An outstanding species-rich fog oasis is the latitudinal fringe Paposo-Taltal, that is considered an iconic site of the Lomas formation. This contribution is the first to reveal the knowledge on arthropods of this emblematic coastal section. We used pitfall traps to study the taxonomic composition, richness, and abundance of terrestrial arthropods in 17 sample sites along a 100 km section of the coast between 24.5 and 25.5 southern latitude, in a variety of characteristic habitats. From a total of 9154 individuals, we were able to identify 173 arthropod species grouped into 118 genera and 57 families. The most diverse group were insects, with 146 species grouped in 97 genera and 43 families, while arachnids were represented by 27 species grouped into 21 genera and 14 families. Current conservation challenges on a global scale are driving the creation and evaluation of potential conservation sites in regions with few protected areas, such as the margins of the Atacama Desert. Better taxonomic, distributional, and population knowledge is urgently needed to perform concrete conservation actions in a biodiversity hotspot at a desert edge.

Low temperatures induce physiological changes in lipids, fatty acids and hydrocarbons, in two rare winter scorpions of genus Urophonius (Scorpiones, Bothriuridae).

Different organisms (mainly poikilotherms) are subject to environmental fluctuations that could affect their normal physiological functioning (e.g., by destabilization of biomembranes and rupture of biomolecules). As a result, animals regulate their body temperature and adapt to different environmental conditions through various physiological strategies. These adaptations are crucial in all organisms, although they are more relevant in those that have reached a great adaptive diversity such as scorpions. Within scorpions, the genus Urophonius presents species with winter activity, being this a peculiarity within the Order and an opportunity to study the strategies deployed by these organisms when facing different temperatures. Here, we explore three basic issues of lipid remodeling under high and low temperatures, using adults and juveniles of Urophonius achalensis and U. brachycentrus. First, as an indicator of metabolic state, we analyzed the lipidic changes in different tissues observing that low temperatures generate higher quantities of triacylglycerols and fewer amount of structural lipids and sphyngomielin. Furthermore, we studied the participation of fatty acids in adaptive homeoviscosity, showing that there are changes in the quantity of saturated and unsaturated fatty acids at low temperature (mainly 16:0, 18:0, 18:1 and 18:2). Finally, we observe that there are quantitative and qualitative variations in the cuticular hydrocarbons (with possible water barrier and chemical recognition function). These fluctuations are in some cases species-specific, metabolic-specific, tissue-specific and in others depend on the ontogenetic state.

Two new species of the sun-spider genus Gaucha from Argentina and Brazil (Solifugae, Mummuciidae).

Two new species in the South American sun-spider family Mummuciidae are herein described. Gaucha ramirezi sp. nov. is known from the Chancaní Provincial Park and Forest Reserve, Córdoba province, Argentina, and further reported for a single locality to the northeast, in Santiago del Estero province. The systematic position of this species is uncertain and it is not assigned to any species-group of Gaucha Mello-Leitão, 1924. The other species, Gaucha santana sp. nov., is only known from the Ibirapuitã Environmental Protection Area, in the southern Brazilian state of Rio Grande do Sul, and is a member of the fasciata species-group. With these descriptions, the number of known species of Gaucha is raised to eleven.

Five new scorpion species of genus Brachistosternus (Scorpiones: Bothriuridae) from the deserts of Chile and Peru, with comments about some poorly studied diagnostic characters of the genus.

Five new scorpion species of genus Brachistosternus of Chile and Peru are described. Brachistosternus gayi n. sp. is a high Andean species of north central Chile. Brachistosternus philippii n. sp. occurs near the coast of Antofagasta. Brachistosternus misti n. sp. occurs at intermediates altitudes of southern Peru. Brachistosternus contisuyu n. sp. occurs in Lomas formation in southern Peru. Brachistosternus anandrovestigia n. sp. occurs in coastal areas of southern Peru, and is the second known species of the genus without metasomal glands or androvestigia. Two diagnostic characters are discussed: the Internal Laminar Apophysis of the right hemispermatophore, and the sternum macrosetae.

Multiple approaches to understanding the taxonomic status of an enigmatic new scorpion species of the genus Tityus (Buthidae) from the biogeographic island of Paraje Tres Cerros (Argentina).

Tityus curupi n. sp., belonging to the bolivianus complex, is described from the biogeographically distinct area of Paraje Tres Cerros in north-eastern Argentina. We also present a molecular species delimitation analysis between Tityus curupi n. sp. and its sister species Tityus uruguayensis Borelli 1901 to confirm species integrity. Furthermore, a cytogenetic analysis is presented for these two species which contain different multivalent associations in meiosis, as a consequence of chromosome rearrangements, and the highest chromosome numbers in the genus.

Dated phylogenetic studies of the southernmost American buthids (Scorpiones; Buthidae).

A dated molecular phylogeny of the southernmost American species of the family Buthidae, based on two nuclear and two mitochondrial genes, is presented. Based on this study, analyzed species of the subgenus Tityus (Archaeotityus) are neither sister to the remaining species of the genus Tityus, nor are they closely related to the New World microbuthids with decreasing neobothriotaxy. Analyzed species of the subgenus Tityus do not form a monophyletic group. Based on ancestral area estimation analyses, known geoclimatic events of the region and comparisons to the diversification processes of other epigean groups from the area, a generalized hypothesis about the patterns of historical colonization processes of the family Buthidae in southern South America is presented. Furthermore, for the first time, a Paleogene-African ingression route for the colonization of America by the family Buthidae is proposed as a plausible hypothesis.

Phylogeny, species delimitation and convergence in the South American bothriurid scorpion genus Brachistosternus Pocock 1893: Integrating morphology, nuclear and mitochondrial DNA.

A phylogenetic analysis of the scorpion genus Brachistosternus Pocock, 1893 (Bothriuridae Simon, 1880) is presented, based on a dataset including 41 of the 43 described species and five outgroups, 116 morphological characters and more than 4150 base-pairs of DNA sequence from the nuclear 18S rDNA and 28S rDNA gene loci, and the mitochondrial 12S rDNA, 16S rDNA, and Cytochrome c Oxidase Subunit I gene loci. Analyses conducted using parsimony, Maximum Likelihood and Bayesian Inference were largely congruent with high support for most clades. The results confirmed the monophyly of Brachistosternus, the nominal subgenus, and subgenus Ministernus Francke, 1985, as in previous analyses based only on morphology, but differed in several other respects. Species from the plains of the Atacama Desert diverged basally whereas the high altitude Andean species radiated from a more derived ancestor, presumably as a consequence of Andean uplift and associated changes in climate. Species limits were assessed among species that contain intraspecific variation (e.g., different morphs), are difficult to separate morphologically, and/or exhibit widespread or disjunct distributions. The extent of convergence in morphological adaptation to life on sandy substrata (psammophily) and the complexity of the male genitalia, or hemispermatophores, was investigated. Psammophily evolved on at least four independent occasions. The lobe regions of the hemispermatophore increased in complexity on three independent occasions, and decreased in complexity on another three independent occasions.

Scorpion sheds 'tail' to escape: consequences and implications of autotomy in scorpions (Buthidae: Ananteris).

Autotomy, the voluntary shedding or detachment of a body part at a determined cleavage plane, is a common anti-predation defense mechanism in several animal taxa, including arthropods. Among arachnids, autotomy has been observed in harvestmen, mites, and spiders, always involving the loss of legs. Autotomy of the opisthosoma (abdomen) was recently reported in a single species of the Neotropical buthid scorpion genus Ananteris Thorell, 1891, but few details were revealed. Based on observations in the field and laboratory, examination of material in museum collections, and scanning electron microscopy, we document autotomy of the metasoma (the hind part of the opisthosoma, or 'tail') in fourteen species of Ananteris. Autotomy is more common in males than females, and has not been observed in juveniles. When the scorpion is held by the metasoma, it is voluntarily severed at the joints between metasomal segments I and II, II and III, or III and IV, allowing the scorpion to escape. After detachment, the severed metasoma moves (twitches) automatically, much like the severed tail of a lizard or the severed leg of a spider, and reacts to contact, even attempting to sting. The severed surface heals rapidly, scar tissue forming in five days. The lost metasomal segments and telson cannot be regenerated. Autotomy of the metasoma and telson results in permanent loss of the posterior part of the scorpion's digestive system (the anus is situated posteriorly on metasomal segment V) and the ability to inject venom by stinging. After autotomy, scorpions do not defecate and can only capture small prey items. However, males can survive and mate successfully for up to eight months in the laboratory. In spite of diminished predation ability after autotomy, survival allows males to reproduce. Autotomy in Ananteris therefore appears to be an effective, adaptive, anti-predation escape mechanism.

Two new scorpion species from Paposo, in the coastal desert of Taltal, Chile (Scorpiones, Bothriuridae, Brachistosternus).

Brachistosternus paposo n. sp. and Brachistosternus barrigai n. sp. are described from Paposo, in the coastal transitional desert of northern Chile. Brachistosternus paposo n. sp. is closely related to Brachistosternus roigalsinai Ojanguren-Affilastro 2002, and B. barrigai n. sp. is closely related with Brachistosternus kamanchaca Ojanguren-Affilastro, Mattoni & Prendini 2007.