Ligia R. Benavides

Rounding up the usual suspects: a standard target-gene approach for resolving the interfamilial phylogenetic relationships of ecribellate orb-weaving spiders with a new family-rank classification (Araneae, Araneoidea)

We test the limits of the spider superfamily Araneoidea and reconstruct its interfamilial relationships using standard molecular markers. The taxon sample (363 terminals) comprises for the first time representatives of all araneoid families, including the first molecular data of the family Synaphridae. We use the resulting phylogenetic framework to study web evolution in araneoids. Araneoidea is monophyletic and sister to Nicodamoidea rank. n. Orbiculariae are not monophyletic and also include the RTA clade, Oecobiidae and Hersiliidae. Deinopoidea is paraphyletic with respect to a lineage that includes the RTA clade, Hersiliidae and Oecobiidae. The cribellate orb‐weaving family Uloboridae is monophyletic and is sister group to a lineage that includes the RTA Clade, Hersiliidae and Oecobiidae. The monophyly of most Araneoidea families is well supported, with a few exceptions. Anapidae includes holarchaeids but the family remains diphyletic even if Holarchaea is considered an anapid. The orb‐web is ancient, having evolved by the early Jurassic; a single origin of the orb with multiple “losses” is implied by our analyses. By the late Jurassic, the orb‐web had already been transformed into different architectures, but the ancestors of the RTA clade probably built orb‐webs. We also find further support for a single origin of the cribellum and multiple independent losses. The following taxonomic and nomenclatural changes are proposed: the cribellate and ecribellate nicodamids are grouped in the superfamily Nicodamoidea rank n. (Megadictynidae rank res. and Nicodamidae stat. n.). Araneoidea includes 17 families with the following changes: Araneidae is re‐circumscribed to include nephilines, Nephilinae rank res., Arkyidae rank n., Physoglenidae rank n., Synotaxidae is limited to the genus Synotaxus, Pararchaeidae is a junior synonym of Malkaridae (syn. n.), Holarchaeidae of Anapidae (syn. n.) and Sinopimoidae of Linyphiidae (syn. n.).

Forest refugia in Western and Central Africa as ‘museums’ of Mesozoic biodiversity

The refugial speciation model, or ‘species pump’, is widely accepted in the context of tropical biogeography and has been advocated as an explanation for present species distributions in tropical Western and Central Africa. In order to test this hypothesis, a phylogeny of the cryptic arachnid order Ricinulei, based on four nuclear and mitochondrial DNA markers, was inferred. This ancient clade of litter-dwelling arthropods, endemic to the primary forests of Western and Central Africa and the Neotropics, might provide insights into the mode and tempo of evolution in Africa. Twenty-six African ricinuleid specimens were sampled from eight countries spanning the distribution of Ricinulei on the continent, and analysed together with Neotropical samples plus other arachnid outgroups. The phylogenetic and molecular dating results suggest that Ricinulei diversified in association with the fragmentation of Gondwana. The early diversification of Ricinoides in Western and Central Africa around 88 (±33) Ma fits old palaeogeographical events better than recent climatic fluctuations. Unlike most recent molecular studies, these results agree with fossil evidence, suggesting that refugia may have acted as ‘museums’ conserving ancient diversity rather than as engines generating diversity during successive episodes of climatic fluctuation in Africa.

A revision of selected clades of Neotropical mite-harvestmen (Arachnida, Opiliones, Cyphophthalmi, Neogoveidae) with the description of eight new species

Abstract Among the least-studied families of mite harvestmen (Opiliones, Cyphophthalmi) is the family Neogoveidae, a group of arachnids that inhabits tropical rain forests on both sides of the Atlantic Ocean around the Equator, and an exquisite example of Gondwanan vicariance. To evaluate the diversity within Neogoveidae we studied the morphology of all currently recognized genera using a recent phylogeny of the group to provide a framework for the family in the Neotropics. That study identifies several clades that correspond to the recognized genera Neogovea, Huitaca, Metagovea, Canga, and the recently resurrected genus Brasilogovea, formerly synonymized with Neogovea. Neogovea is restricted to eastern Amazonia (Brazil, Guyana, French Guiana, Suriname), Huitaca remains endemic to Colombia, Canga is restricted to its type locality in the Serra de Carajás (Pará State, Brazil), Brasilogovea to Central Amazonas and the Tepuis region in Colombia, and Metagovea is the most widespread genus, found along the northern Andes from Perú to Venezuela, and extending to the Amazon region. All Neotropical genera are diagnosed and discussed. The formerly monotypic Huitaca is revised with the description of six additional species, all found in the high-altitude Colombian mountain areas called cordilleras, between 2,030 and 3,050 m. These include: Huitaca bitaco new species, Huitaca boyacaensis new species, Huitaca caldas new species, Huitaca depressa new species, Huitaca sharkeyi new species, and Huitaca tama new species, all from Colombia. Two additional species, Brasilogovea chiribiqueta new species, from Colombia, and Neogovea hormigai new species, from Guyana, are also described.

Molecular phylogenetic analysis of “pirate spiders” (Araneae, Mimetidae) with the description of a new African genus and the first report of maternal care in the family

We investigate the phylogeny of “pirate spiders” (Mimetidae), a family of araneophagic spiders known for their use of aggressive mimicry as a foraging strategy, but poorly understood phylogenetically. Relationships are inferred by including molecular data from six loci for 92 mimetid terminals spanning four genera, and 119 outgroups representing 12 families. Phylogenetic analyses based on parsimony, maximum‐likelihood and Bayesian approaches, as well as static and dynamic homology, robustly support monophyly of Mimetidae and a sister‐group relationship to a clade comprising Tetragnathidae + Arkyidae. Relationships among the mimetid genera are largely congruent across methods, as follows: (Gelanor (Ero (Anansi n. gen. (Australomimetus, Mimetus)))). Diversification of Mimetidae is estimated to be around 114 Ma, in the Early Cretaceous. In light of the results of our phylogenetic analyses, we erect Anansi n. gen. to include a clade of mimetids from West Africa that contains at least four species, including the newly described A. luki n. sp. We present the first report of maternal care in Mimetidae based on novel field observations.

Taxonomic revision of the Neotropical pirate spiders of the genus Gelanor Thorell, 1869 (Araneae, Mimetidae) with the description of five new species.

We revise the Neotropical spider genus Gelanor Thorell, 1869 (Mimetidae). Gelanor is distributed from northeast Mexico to southern Uruguay , from sea level to 1,600 m. We describe five new species of Gelanor and report eleven new synonymies. Gelanor is here circumscribed to include ten species: Gelanor fortuna new species, Gelanor juruti new species, Gelanor moyobamba new species, Gelanor siquirres new species, Gelanor waorani new species, Gelanor altithorax Keyserling, 1893 (= Gelanor lanei Soares, 1941 new synonymy), Gelanor consequus O. P.-Cambridge, 1902 (= Gelanor depressus Chickering, 1956 new synonymy, Gelanor gertschi Chickering, 1947 new synonymy and Gelanor heraldicus Petrunkevitch, 1925 new synonymy), Gelanor innominatus Chamberlin, 1916, Gelanor latus (Keyserling, 1881) (= Gelanor mixtus O. P.-Cambridge, 1899 new synonymy, Gelanor mabelae Chickering, 1947 new synonymy, Gelanor ornatus Schenkel, 1953 new synonymy and Gelanor proximus Mello-Leitão, 1929 new synonymy) and Gelanor zonatus (C.L. Koch, 1845) (= Gelanor distinctus O-P. Cambridge, 1899 new synonymy, Gelanor insularis Mello-Leitão, 1929 new synonymy and Gelanor obscurus Mello-Leitão, 1929 new synonymy). In addition, we describe for the first time the males of G. altithorax and G. consequus. Species descriptions are provided for all ten species in the genus, together with a compilation of available data, including type specimens, type localities and morphological diagnoses. Light and electron microscope images and updated data on known geographical distributions, are also provided. We also discuss the phylogenetic placement of Gelanor in Mimetidae.

The ‘Peripatos’ in Eurogondwana? – Lack of evidence that south-east Asian onychophorans walked through Europe

Abstract. Onychophorans, or velvet worms, are cryptic but extremely charismatic terrestrial invertebrates that have often been the subject of interesting biogeographic debate. Despite great interest, a well resolved and complete phylogeny of the group and a reliable chronogram have been elusive due to their broad geographic distribution, paucity of samples, and challenging molecular composition. Here we present a molecular phylogenetic analysis of Onychophora that includes previously unsampled and undersampled lineages and we analyse the expanded dataset using a series of nested taxon sets designed to increase the amount of information available for particular subclades. These include a dataset with outgroups, one restricted to the ingroup taxa, and three others for Peripatopsidae, Peripatidae and Neopatida (= the Neotropical Peripatidae). To explore competing biogeographic scenarios we generate a new time tree for Onychophora using the few available reliable fossils as calibration points. Comparing our results to those of Cyphophthalmi, we reconsider the hypothesis that velvet worms reached Southeast Asia via Eurogondwana, and conclude that a more likely scenario is that they reached Southeast Asia by rafting on the Sibumasu terrane. Our phylogenetic results support the reciprocal monophyly of both families as well as an early division between East and West Gondwana, also in both families, each beginning to diversify between the Permian and the Jurassic. Peripatopsidae clearly supports paraphyly of South Africa with respect to southern South America (Chile) and a sister group relationship of the Southeast Asian/New Guinean Paraperipatus to the Australian/New Zealand taxa. The latter includes a clade that divides between Western Australia and Eastern Australia and two sister clades of trans-Tasman species (one oviparous and one viviparous). This pattern clearly shows that oviparity is secondarily derived in velvet worms. Peripatidae finds a sister group relationship between the Southeast Asian Eoperipatus and the West Gondwanan clade, which divides into the African Mesoperipatus and Neopatida. The latter shows a well supported split between the Pacific Oroperipatus (although it is unclear whether they form one or two clades) and a sister clade that includes the members of the genera Peripatus, Epiperipatus, Macroperipatus and representatives of the monotypic genera Cerradopatus, Plicatoperipatus and Principapillatus. However, Peripatus, Epiperipatus and Macroperipatus are not monophyletic, and all the species from the monotypic genera are related to geographically close species. The same goes for the type species of Macroperipatus (from Trinidad, and sister group to other Trinidad and Tobago species of Epiperipatus) and Epiperipatus (from French Guiana, and related to other Guyana shield species of Epiperipatus and Peripatus). Geographic structure within Neopatida is largely obscured by an unresolved backbone, but many well supported instances of generic non-monophyly challenge the current taxonomic framework, which has often relied on anatomical characters that are untested phylogenetically.

The systematics and biogeography of the mite harvestman family Sironidae (Arachnida : Opiliones : Cyphophthalmi) with the description of five new species

Abstract. Sironidae, the first described family of Cyphophthalmi, is among the least understood phylogenetically. After examining recent collections across their distribution range, we provide the first comprehensive treatment of Sironidae by including molecular data from most of the known species, and all genera except for the monotypic Odontosiro Juberthie, 1961. We also revisit the male genitalic morphology for most genera by using confocal laser scanning microscopy and provide descriptions of five new species belonging to Iberosiro de Bivort & Giribet, 2004 (monotypic until now), Paramiopsalis Juberthie, 1962 and Siro Latreille, 1802. While the monophyly of Sironidae remains poorly supported using traditional Sanger-based markers, with the Mediterranean Parasiro Hansen & Sørensen, 1904 and the Japanese Suzukielus Juberthie, 1970b sometimes branching basally with respect to the other sironids, the remaining genera form a well-supported Laurentian/Laurasian clade. This group divides into a Western European/North American clade of Siro and the remaining genera, Iberosiro, Paramiopsalis and Cyphophthalmus Joseph, 1868. Iberosiro and Paramiopsalis form a well-supported clade from the NW corner of the Iberian Peninsula, while Cyphophthalmus is widespread in the Balkan region and Eastern Mediterranean. Finally, the following new taxa are described: Iberosiro rosae Giribet, Merino-Sáinz & Benavides, sp. nov., Paramiopsalis anadonae Giribet, Merino-Sáinz & Benavides, sp. nov., Paramiopsalis ramblae Benavides & Giribet, sp. nov., Siro ligiae Giribet, sp. nov., and Siro richarti Benavides & Giribet, sp. nov.

Figures 8 – 13. Metagovea Oviformis Martens, 1969 In An Illustrated Catalogue Of The South American Species Of The Cyphophthalmid Family Neogoveidae (Arthropoda, Opiliones, Cyphophthalmi) With A Report On 37 Undescribed Species

FIGURES 8 – 13. Metagovea oviformis Martens, 1969, male holotype (SFM 23959) (8 – 10), female paratype (SFM 23960) (11 – 13). 8, 11 dorsal view; 9, 12 lateral view; 10, 13 ventral view.