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New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae)
(2016)
We describe eight new species of the genus Pholcus, and document their microhabitats. Four species are assigned to the previously described Pholcus ethagala group: P. tanahrata Huber sp. nov., P. uludong Huber sp. nov., and P. bukittimah Huber sp. nov. from the Malay Peninsula, and P. barisan Huber sp. nov. from Sumatra. These species are all litter-dwellers that build domed sheet webs on the undersides of large dead leaves on the ground. The other four species are assigned to newly created species groups: the P. tambunan group with two species from northern Borneo: P. tambunan Huber sp. nov. and P. bario Huber sp. nov.; and the P. domingo group with two species from the Philippines, Mindanao: P. domingo Huber sp. nov. and P. matutum Huber sp. nov. These latter four species are leafdwellers that build barely visible silk platforms tightly attached to the undersides of live leaves. The main rationale for this paper is to provide part of the taxonomic and natural history background for upcoming phylogenetic and evolutionary (microhabitat shifts) analyses.
The southern South American genus Guaranita includes tiny spiders (body length ~1 mm) that lead reclusive lives under ground-objects and run rapidly when disturbed. As a result, they have been poorly collected and studied. Here we report on a recent collection of Guaranita spiders from Argentina, describing one new species (G. auadae Huber sp. nov.) and the previously unknown female of G. dobby Torres et al., 2016. In addition, we provide CO1 barcodes for all (now five) known species, first SEM data, and first chromosome data for the genus. The diploid number of Guaranita goloboffi Huber, 2000 (2n♂ = 11) is among the lowest in araneomorph spiders with monocentric chromosome structure.
This paper summarizes current knowledge about Central African pholcids. Central Africa is here defined as the area between 10°N and 7°S and between 6°E and 18°E, including mainly the Lower Guinean subregion of the Guineo-Congolian center of endemism. This includes all of Gabon, Equatorial Guinea, São Tomé and Príncipe, most of Cameroon and Congo Republic, and parts of the neighboring countries. An annotated list of the 14 genera and 79 species recorded from this area is given, together with distribution maps and an identification key to genera. Seven species are newly described: Anansus kamwai sp. nov., Leptopholcus gabonicus sp. nov., Ninetis faro sp. nov., Pholcus punu sp. nov., P. rawiriae sp. nov., Spermophora abibae sp. nov., and S. awalai sp. nov. Additional new records are given for 16 previously described species, including 17 new country records. Distribution and diversity patterns are compared with data on West and East Africa. While West Africa contains a similar set of genera it is significantly less diverse than Central Africa. East Africa is taxonomically more distinct. It has similar levels of diversity as Central Africa, but appears to be less undersampled.
Daddy-long-leg giants: revision of the spider genus Artema Walckenaer, 1837 (Araneae, Pholcidae)
(2017)
This is the first revision of Artema Walckenaer, 1837, a genus consisting of large and phylogenetically interesting species. Even though Artema is not species-rich (now eight nominal species), it has suffered from poor descriptions and synonymies. Our main goal was to gather all available material and to clarify species limits. Four species are easily distinguished from other congeners: Artema atlanta Walckenaer, 1837, the type species; A. kochi Kulczyński, 1901 (revalidated); A. bunkpurugu Huber & Kwapong, 2013; and A. nephilit sp. nov. All other species are problematic for varying reasons: species limits are unclear between A. doriae Thorell, 1881 and A. transcaspica Spassky, 1934; A. magna Roewer, 1960 and A. ziaretana (Roewer, 1960) are problematic because they are based on female and juvenile types respectively and little new material is available. The material available to us suggests the existence of a few further species; however, they are not formally described, either because of small sample sizes (Artema sp. a and A. sp. b are represented by only one specimen each) or because of unclear species limits (between Artema sp. c, A. transcaspica and A. doriae).This study is the first serious step towards understanding the genus. Intensive collecting effort is needed in order to fully clarify species limits.
The North American-Caribbean genera Pholcophora Banks, 1896 and Tolteca Huber, 2000 are representatives of Ninetinae, a group of small, cryptic, and thus poorly known pholcid spiders. We present the first comprehensive revisions of the two genera, including extensive SEM data and descriptions of seven new species from Mexico (Pholcophora mazatlan Huber sp. nov., P. papanoa Huber sp. nov., P. tehuacan Huber sp. nov., Tolteca huahua Huber sp. nov., T. manzanillo Huber sp. nov., T. oaxaca Huber sp. nov., and T. sinnombre Huber sp. nov.). We add new CO1 sequences of nine species to previously published molecular data and use these for a preliminary analysis of relationships. We recover a North American-Caribbean clade including ‘true’ (mainland) Pholcophora, Tolteca (Mexico), and a Caribbean clade consisting of the genus Papiamenta Huber, 2000 (Curaçao) and Caribbean ‘Pholcophora’. First karyotype data for Tolteca (2n♂ = 13, X1X2Y and 15, X1X2Y, respectively) reveal a strong reduction of the number of chromosome pairs within the North American-Caribbean clade, and considerable karyotype differentiation among congeners. This agrees with considerable CO1 divergence among species of Tolteca but contrasts with very inconspicuous morphological divergence. Environmental niche analyses show that the widespread P. americana Banks, 1896 (western USA, SW Canada) occupies a very different niche than its Mexican congeners and other close relatives. Caribbean taxa also have a low niche overlap with ‘true’ Pholcophora and Tolteca, supporting the idea that Caribbean ‘Pholcophora’ are taxonomically misplaced.
The previously poorly known Southeast Asian spider genus Aetana Huber, 2005 is revised. Fifteen species are newly described, and the first SEM data and a first phylogenetic analysis of the genus are presented. Four species groups are well supported, one restricted to Borneo, two restricted to the Philippines, and one ranging from the Philippines to Fiji. The cladistic analysis and field observations suggest that the ancestor of Aetana built its web close to the ground, in confined spaces among and under rocks and logs. In at least two cases, evolutionary shifts of microhabitat resulted in species being adapted to life in higher forest strata, with correlated morphological and behavioral changes (lighter coloration; longer abdomen; additional sheet in web or more strongly domed web). The following species are newly described: A. abadae Huber, sp. nov., A. baganihan Huber, sp. nov., A. banahaw Huber, sp. nov., A. kiukoki Huber, sp. nov., A. libjo Huber, sp. nov., A. loboc Huber, sp. nov., A. lozadae Huber, sp. nov., A. manansalai Huber, sp. nov., A. ocampoi Huber, sp. nov., A. paragua Huber, sp. nov. and A. pasambai Huber, sp. nov. from the Philippines; A. gaya Huber, sp. nov., A. indah Huber, sp. nov., A. lambir Huber, sp. nov. and A. poring Huber, sp. nov. from northern Borneo. The female of A. kinabalu Huber, 2005 is newly described. A potential case of female genital dimorphism is documented in A. ocampoi Huber, sp. nov.
We describe several new species of the previously monotypic Panjange lanthana species group from the Philippines and document their extraordinary morphology. Some species show strong male genital asymmetry, a phenomenon that seems to be exceedingly rare in spiders. Males of most species have eye stalks, and in two species these eye stalks are among the longest ever recorded in spiders. Some species show a tendency for male genital (pedipalp) elongation, and one species has the longest and thinnest palps ever recorded in Pholcidae. A cladistic analysis is performed including all described and several undescribed species of Panjange (except for one “problem species”), supporting the lanthana group and its close relationship with members of the possibly paraphyletic cavicola group. The following eight new species are described: Panjange malagos Huber sp. nov.; Pa. casaroro Huber sp. nov.; Pa. camiguin Huber sp. nov.; Pa. hamiguitan Huber sp. nov.; Pa. isarog Huber sp. nov.; Pa. dinagat Huber sp. nov.; Pa. marilog Huber sp. nov.; Pa. bukidnon Huber sp. nov.
We revise the Southeast Asian Pholcus bicornutus group in which males are characterized by a unique pair of horns on their ocular area, each of which carries at its tip a brush of hairs. In two species, the two hair brushes are ‘glued’ or ‘waxed’ together by an unidentified substance into a very consistently curved and pointed single median tip. In the other five species known, the hairs are unglued. We present a first revision of ocular modifications in Pholcidae and identify twenty supposedly independent origins. Most cases are in Pholcinae, and all but one case are limited to the male, suggesting sexual selection as the main driving force in the evolution of ocular modifications in Pholcidae. Previously, the Pholcus bicornutus group consisted of four species limited to the Philippines. We describe four new species, including three species from the Philippines (P. olangapo Huber, sp. nov.; P. kawit Huber, sp. nov.; P. baguio Huber, sp. nov.) and the first representative from outside the Philippines (P. mulu Huber, sp. nov. from Sarawak, NE Borneo) and provide new records and SEM data for three previously described species.
We revise the Panjange nigrifrons group in Borneo and document an unexpected diversity in western Sarawak forests. Five species occur within 80 km from Kuching, each species being known from its type locality only. Further species occur east until Niah, but the genus seems to be absent from Sabah. We contrast this with another pholcid genus (Aetana Huber, 2005), which is diverse in Sabah and westward until Niah, but does not seem to occur in central and western Sarawak. Five species are newly described: Panjange kapit Huber, sp. nov., Panjange kubah Huber, sp. nov., Panjange niah Huber, sp. nov., Panjange pueh Huber, sp. nov., Panjange seowi Huber, sp. nov.; Panjange tahai (Huber, 2011) comb. nov. is transferred from Pholcus.
Two small, ground and litter-dwelling pholcid species from northern Borneo are described as representatives of a new genus, Hantu gen. nov.: H. kapit gen. et sp. nov. and H. niah gen. et sp. nov. Previous cladistic analyses suggested a closer relationship with the genera Savarna Huber, 2005 and Khorata Huber, 2005 (mainland Southeast Asia) than with the geographically closer genus Aetana Huber, 2005 (Borneo and Philippines to Fiji). Since the two species do not share any of the synapomorphies of Khorata and Savarna while having several synapomorphies on their own (ventral apophysis on male palpal coxa; male palpal trochanter apophysis with small teeth or scales; spines on male femora 1; high density of vertical hairs on male femora; presence of scape on epigynum), they are here proposed as representing a new genus.