Under the hood: Phylogenomics of hooded tick spiders (Arachnida, Ricinulei) uncovers discordance between morphology and molecules.

Ricinulei or hooded tick-spiders are a cryptic and ancient group of arachnids. The order consists of around 100 highly endemic extant species restricted to the Afrotropics and the Neotropics along with 22 fossil species. Their antiquity and low vagility make them an excellent group with which to interrogate biogeographic questions. To date, only four molecular analyses have been conducted on the group and they failed to resolve the relationships of the main lineages and even recovering the non-monophyly of the three genera. These studies were limited to a few Sanger loci or phylogenomic analyses with at most seven ingroup samples. To increase phylogenetic resolution in this little-understood and poorly studied group, we present the most comprehensive phylogenomic study of Ricinulei to date leveraging the Arachnida ultra-conserved element probe set. With a data set of 473 loci across 96 ingroup samples, analyses resolved a monophyletic Neotropical clade consisting of four main lineages. Two of them correspond to the current genera Cryptocellus and Pseudocellus while topology testing revealed one lineage to likely be a phylogenetic reconstruction artefact. The fourth lineage, restricted to Northwestern, Andean South America, is consistent with the Cryptocellus magnus group, likely corresponding to the historical genus Heteroricinoides. Since we did not sample the type species for this old genus, we do not formally re-erect Heteroricinoides but our data suggest the need for a thorough morphological re-examination of Neotropical Ricinulei.

Volcanism and palaeoclimate change drive diversification of the world's largest whip spider (Amblypygi).

The tropics contain many of the most biodiverse regions on Earth but the processes responsible for generating this diversity remain poorly understood. This study investigated the drivers of diversification in arthropods with stenotopic ecological requirements and limited dispersal capability using as a model the monotypic whip spider (Amblypygi) genus Acanthophrynus, widespread in the tropical deciduous forests of Mexico. We hypothesized that for these organisms, the tropical deciduous forests serve as a conduit for dispersal, with their disappearance imposing barriers. Given that these forests are located in a region of complex geological history and that they fluctuated in extent during the Pliocene-Pleistocene glacial/interglacial cycles we combine molecular divergence dating, palaeoclimatic niche modelling and ancestral area reconstruction to test if and when habitat fragmentation promoted diversification in Acanthophrynus. Concomitant with the expected role of landscape change, we demonstrate that orogeny of the Trans-Mexican Volcanic Belt, in the Late Miocene or Early Pliocene (6.95-5.21 million years ago), drove the earliest divergence of Acanthophrynus by vicariance. Similarly, as expected, the later onset of glaciations strongly impacted diversification. Whereas a more stable climate in the southern part of the distribution enabled further diversification, a marked loss of suitable habitat during the glaciations only allowed dispersal and diversification in the north to occur later, resulting in a lower overall diversity in this region. Barriers and diversification patterns identified in Acanthophrynus are reflected in the phylogeography of codistributed vertebrates and arthropods, emphasizing the profound impact of Trans-Mexican Volcanic Belt orogeny and glacial/interglacial cycles as drivers of diversification in the Mexican Neotropics.

Microhabitat change drives diversification in pholcid spiders.

BACKGROUND: Microhabitat changes are thought to be among the main drivers of diversification. However, this conclusion is mostly based on studies on vertebrates. Here, we investigate the influence of microhabitat on diversification rates in pholcid spiders (Araneae, Pholcidae). Diversification analyses were conducted in the framework of the largest molecular phylogeny of pholcid spiders to date based on three nuclear and three mitochondrial loci from 600 species representing more than 85% of the currently described pholcid genera. RESULTS: Assessments of ancestral microhabitat revealed frequent evolutionary change. In particular, within the largest subfamily Pholcinae, numerous changes from near-ground habitats towards leaves and back were found. In general, taxa occupying leaves and large sheltered spaces had higher diversification rates than ground-dwelling taxa. Shifts in speciation rate were found in leaf- and space-dwelling taxa. CONCLUSIONS: Our analyses result in one of the most comprehensive phylogenies available for a major spider family and provide a framework for any subsequent studies of pholcid spider biology. Diversification analyses strongly suggest that microhabitat is an important factor influencing diversification patterns in pholcid spiders.

A new troglobitic species of the spider genus Tengella Dahl (Araneae, Tengellidae) from Chiapas, Mexico.

A new species of troglobitic spider of the genus Tengella Dahl is described from Mexico: Tengella kalebi new species, from a limestone cave in Chiapas, Mexico. The species is described on the basis of adult male and females. This is the fifth species described for the genus Tengella, the third species described from Mexico, the first species that has notable troglomorphic modifications, and the first ecribellate Tengella species.

Taxonomic revision of the spider genus Ixchela Huber, 2000 (Araneae: Pholcidae), with description of ten new species from Mexico and Central America.

 A taxonomic revision of the genus Ixchela Huber, 2000 is presented. The previously described five species are redescribed with new taxonomic information and new distribution records. The males of Ixchela placida (Gertsch, 1971) and Ixchela simoni (O. Pickard-Cambridge, 1898) are described for the first time. Ten new species are described, nine from Mexico and one from Honduras: Ixchela mixe new species, Ixchela huberi new species, Ixchela juarezi new species, Ixchela grix new species, Ixchela taxco new species, Ixchela franckei new species, Ixchela tzotzil new species, Ixchela santibanezi new species, Ixchela huasteca new species, and Ixchela viquezi new species. Identification keys for males and females are presented.

Two new species of ricinuleids of the genus Pseudocellus (Arachnida: Ricinulei: Ricinoididae) from southern Mexico.

Two new species of ricinuleids of the genus Pseudocellus are described from Mexico: Pseudocellus cruzlopezi sp. nov. from Oaxaca, and Pseudocellus monjarazi sp. nov. from Chiapas. Both species are described from adult males and females. The first species is epigean and edaphomorphic, whereas the second is cavernicolous and troglomorphic. The number of known species of the genus Pseudocellus increases to 25, and Mexican species to 16, indicating that Mexico has the highest diversity of ricinuleids in the world. An updated identification key to adult males of the 16 described species found in Mexico and southern USA is provided.

To be or not to be Integrative taxonomy and species delimitation in the daddy long-legs spiders of the genus Physocyclus (Araneae, Pholcidae) using DNA barcoding and morphology.

Integrative taxonomy is crucial for discovery, recognition, and species delimitation, especially in underestimated species complex or cryptic species, by incorporating different sources of evidence to construct rigorous species hypotheses. The spider genus Physocyclus Simon, 1893 (Pholcidae, Arteminae) is composed of 37 species, mainly from North America. In this study, traditional morphology was compared with three DNA barcoding markers regarding their utility in species delimitation within the genus: 1) Cytochrome c Oxidase subunit 1 (CO1), 2) Internal Transcribed Spacer 2 (ITS2), and 3) Ribosomal large subunit (28S). The molecular species delimitation analyses were carried out using four methods under the corrected p-distances Neighbor-Joining (NJ) criteria: 1) Automatic Barcode Gap Discovery (ABGD), 2) Assemble Species by Automatic Partitioning (ASAP), 3) General Mixed Yule Coalescent model (GMYC), and 4) Bayesian Poisson Tree Processes (bPTP). The analyses incorporated 75 terminals from 22 putative species of Physocyclus. The average intraspecific genetic distance (p-distance) was found to be < 2%, whereas the average interspecific genetic distance was 20.6%. The ABGD, ASAP, and GMYC methods were the most congruent, delimiting 26 or 27 species, while the bPTP method delimited 33 species. The use of traditional morphology for species delimitation was congruent with most molecular methods, with the male palp, male chelicerae, and female genitalia shown to be robust characters that support species-level identification. The barcoding with CO1 and 28S had better resolution for species delimitation in comparison with ITS2. The concatenated matrix and traditional morphology were found to be more robust and informative for species delimitation within Physocyclus.

COI mtDNA barcoding and morphology for species delimitation in the spider genus Ixchela Huber (Araneae: Pholcidae), with the description of two new species from Mexico.

The spider genus Ixchela Huber 2000, is comprised of 22 species distributed from north-eastern Mexico to Central America, including the two new species described herein from Mexico: Ixchela panchovillai sp. nov. and Ixchela zapatai sp. nov., both from the state of Oaxaca and described for both sexes. DNA barcoding utilizing mitochondrial cytochrome c oxidase subunit 1 (COI) and morphology were used for species delimitation. Molecular analyses and species delimitation included four methods: 1) corrected p-distances under neighbor-joining (NJ), 2) general mixed yule coalescent model (GMYC), 3) automatic barcode gap discovery (ABGD), and 4) Poisson tree processes (bPTP). All molecular methods and morphology were consistent in delimiting and recognizing the two new species describing herein. The average inter-specific genetic distance (p-distance) within the genus Ixchela is 12%. Ixchela panchovillai sp. nov. is closely related in the NJ analysis with I. placida, with an average p-distance of 7.9%, whereas I. zapatai sp. nov. is closely related to I. taxco, with an average p-distance of 8.4% between both species. Additionally, identification keys for males and females of the genus Ixchela are presented.

Under an integrative taxonomic approach: the description of a new species of the genus Loxosceles (Araneae, Sicariidae) from Mexico City.

A new species of the spider genus Loxosceles Heineken & Lowe, 1832, Loxosceles tenochtitlan Valdez-Mondragón & Navarro-Rodríguez, sp. nov., is described based on adult male and female specimens from the states of Mexico City, Estado de Mexico and Tlaxcala. Integrative taxonomy including traditional morphology, geometric and lineal morphology, and molecules (DNA barcodes of cytochrome c oxidase subunit 1 (CO1) and internal transcribed spacer 2 (ITS2)), were used as evidence to delimit the new species. Four methods were used for molecular analyses and species delimitation: 1) corrected p-distances under neighbor joining (NJ), 2) automatic barcode gap discovery (ABGD), 3) general mixed yule coalescent model (GMYC), and 4) poisson tree processes (bPTP). All molecular methods, traditional, geometric and lineal morphology were consistent in delimiting and recognizing the new species. Loxosceles tenochtitlan sp. nov. is closely related to L. misteca based on molecular data. Although both species are morphologically similar, the average p-distance from CO1 data was 13.8% and 4.2% for ITS2 data. The molecular species delimitation methods recovered well-supported monophyletic clusters for samples of L. tenochtitlan sp. nov. from Mexico City + Tlaxcala and for samples of L. misteca from Guerrero. Loxosceles tenochtitlan sp. nov. is considered a unique species for three reasons: (1) it can be distinguished by morphological characters (genitalic and somatic); (2) the four different molecular species delimitation methods were congruent to separate both species; and (3) there is variation in leg I length of males between both species, with the males of L. misteca having longer legs than males of L. tenochtitlan sp. nov., also morphometrically, the shape of tibiae of the palp between males of both species is different.

A new species of Loxosceles Heineken & Lowe (Araneae, Sicariidae), with updated distribution records and biogeographical comments for the species from Mexico, including a new record of Loxoscelesrufescens (Dufour).

A new species of the spider genus Loxosceles Heineken & Lowe, 1832, Loxoscelesmalintzi sp. n., is described from the states of Puebla, Morelos and Guerrero, in the central region of Mexico. The description is based on adult males and females with morphological and ultra-morphological images. Updated distribution maps are provided for the 39 species recorded from the Mexican territory (including the new species). The states with the greatest diversity are Baja California Sur, Baja California and Sonora, with five species each. A total of 441 records for the 39 species, based on arachnological collections, data bases and literature, were used to update the distribution maps. Loxoscelesboneti Gertsch, 1958 is the species with the highest number of records in Mexico, with a total of 58 records from different localities. The states with the most records so far are Guerrero, with 55 records, Morelos, with 35 records, and Baja California Sur, with 30 records. Loxoscelesrufescens (Dufour, 1820), an introduced species, is recorded for the second time in Mexico, from the state of Chihuahua, being the first well-documented record for the country. Mexico has the greatest diversity of species of Loxosceles worldwide, with 39 (two introduced species) of the 134 described species. Additionally, biogeographical comments for the species from Mexico are provided.

On the poorly known haplogynae spiders of the genus Ochyrocera Simon (Araneae, Ochyroceratidae) from Mexico: description of two new species with an updated identification key for Mexican species.

Two new species of the spider genus Ochyrocera Simon 1891 are described from Mexico. Ochyrocera jarocha new species was collected under rotten trunks and hollow trunks in a tropical rainforest, in San Martin Volcano, Veracruz, Mexico. Ochyrocera pojoj new species was collected in a mixed forest, under rotten trunks, in La Trinitaria, Chiapas, Mexico, which represents the third species described from the state of Chiapas. With the description of the two new species herein, six species of Ochyrocera are recorded from Mexico. An updated taxonomic identification key and a distribution map to the Mexican species are provided.

A remarkable new species of the magnus species-group of Cryptocellus (Arachnida, Ricinulei) from Ecuador, with observations on the taxonomy of the New World genera.

A new ricinuleid species, Cryptocellus chimaera sp. nov., is described based on a male specimen from Northwest Ecuador (Esmeraldas, Reserva Ecológica Mache-Chindul, Estación Biológica Bilsa). This species is unique among all Cryptocellus in having very large longitudinal carapacial translucent areas together with a markedly incrassate femur of leg II. Representing only the second species of the order described from Ecuador, C. chimaera sp. nov. is assigned to the magnus species-group of Cryptocellus Westwood, 1874. Cryptocellus chimaera sp. nov. is remarkable, for its morphology resembles that of Cryptocellus magnus Ewing, 1929, especially with regard to the male copulatory apparatus, although both resemble Pseudocellus Platnick, 1980, due to the presence of diffuse longitudinal carapacial translucent areas. Along with the new species description, a comparative diagnosis and supplementary images are provided for C. magnus. Based on direct observations of some species belonging to the five species-groups of Cryptocellus, we discuss on the occurrence of different morphologies of carapacial translucent areas within the genus. We deem it important to continue the survey of morphological characters, especially within Cryptocellus, in order to increase our understanding of the species-groups and to unravel their relationships.

On the trapdoor spiders of Mexico: description of the first new species of the spider genus Aptostichus from Mexico and the description of the female of Eucteniza zapatista (Araneae, Mygalomorphae, Euctenizidae).

A new species of the spider genus Aptostichus Simon, 1891 is described from a cave in Huautla de Jiménez, Oaxaca, Mexico: Aptostichus sabinaesp. n. This species represents the first new species described from Mexico and the southernmost record in North America for the genus so far. Aptostichus sabinaesp. n. represents the forty-first species described for the genus, which has the highest species diversity in the family Euctenizidae. Eucteniza zapatista is redescribed based on five new males and the first known female from the Parque Nacional La Malinche (PNLM), Tlaxcala Mexico. Eucteniza zapatista is the fourth species of the genus where a female is known, and one of fourteen species described for the genus to date.

A reanalysis of the morphological phylogeny of the spider genus Physocyclus Simon (Araneae: Pholcidae) with the description of a new species and description of the female of Physocyclus paredesi Valdez-Mondragón from Mexico.

A new species of spider from Michoacán, Physocyclus peribanensis sp. nov. is described. This description is based on a male holotype and one female paratype. Also, the first description of the female of Physocyclus paredesi Valdez-Mondragón from Oaxaca, Mexico is provided, as well as the redescription of the male. This paper provides a cladistic reanalysis of the spider genus Physocyclus Simon, corroborating the monophyly of the genus with morphological data. The phylogenetic reanalysis was done with 54 morphological characters (44 binary and 10 multistate) using equal and implied weighting approach. The equal weighting analysis found two most parsimonious trees, whereas the analysis with implied weighting found just one most parsimonious trees with the concavity values (K= 5-10). The genus Physocyclus is composed by two clades or species groups: the globosus and the dugesi groups. Physocyclus peribanensis sp. nov. belongs to the dugesi group composed of 21 species, and P. paredesi to the globosus group composed of 11 species. With the new species described here, the number of known species of the genus Physocyclus increases to 32 species. The globosus group has a biogeographical distribution pattern in the Mesoamerican and Mexican Mountain biotic components, whereas the dugesi group has a biogeographical distribution in the Mesoamerican and Continental Nearctic biotic components.

First record of the mygalomorph spider family Paratropididae (Arachnida, Araneae) in North America with the description of a new species of Paratropis Simon from Mexico, and with new ultramorphological data for the family.

A new species of the genus Paratropis is described from North America: Paratropis tuxtlensis sp. n., from a tropical rainforest in Veracruz, Mexico. This is the fifth Paratropis and the tenth paratropidid species described and the first North American record of this Neotropical family. The species is described based on adult males and females, and juveniles. The juveniles show ontogenetic variation in the number of cuspules on the labium and endites, and in the number and position of leg trichobothria. This is the second Paratropis species, and the third paratropidid known from both sexes. The scanning electron photographs (SEM) reveal new morphological data and contribute to the knowledge of the family.