Jimmy Jair Cabra García Revisão e análise filogenética do gênero Glenognatha Simon, 1887 (Araneae, Tetragnathidae) Revision and phylogenetic analysis of the spider genus Glenognatha Simon, 1887 (Araneae, Tetragnathidae) São Paulo 2013 Jimmy Jair Cabra García Revisão e análise filogenética do gênero Glenognatha Simon, 1887 (Araneae, Tetragnathidae) Revision and phylogenetic analysis of the spider genus Glenognatha Simon, 1887 (Araneae, Tetragnathidae) Dissertação apresentada ao Instituto de Biociências da Universidade de São Paulo, para a obtenção de Título de Mestre em Ciências Biológicas, na Área de Zoologia. Orientador(a): Antonio D. Brescovit São Paulo 2013 ABSTRACT A taxonomic revision and phylogenetic analysis of the spider genus Glenognatha Simon, 1887 is presented. The analysis is based on a data set including 24 Glenognatha species plus eight outgroup representatives of three additional tetragnathine genera and one metaine, scored for 82 morphological characters. Eight unambiguous synapomorphies support the monophyly of Glenognatha, all free of homoplasy. Some internal clades within the genus are well-supported and its relationships are discussed. The genus Glenognatha has a broad distribution occupying the Neartic, Neotropic, Afrotropic, Indo-Malaya and Oceania ecozones. As revised here, Glenognatha comprises 27 species, four of them only know from males. New morphological data are provided for the description of thirteen previously described species. Eleven species are newly described: G. sp. nov. 1, G. sp. nov. 3, G. sp. nov. 4 and G. sp. nov. 7 from southeast Brazil, G. sp. nov. 6, G. sp. nov. 9 and G. sp. nov. 10 from the Amazonian region, G. sp. nov. 2, G. sp. nov. 5 and G. sp. nov. 8 from northern Andes and G. sp. nov. 11 from central Mexico and Southern United States. Females of G. minuta Banks, 1898, G. gaujoni Simon, 1895 and G. gloriae (Petrunkevitch, 1930) and males of G. globosa (Petrunkevitch, 1925) and G. hirsutissima (Berland, 1935) are described for the first time. Three new combinations are proposed in congruence with the phylogenetic results: G. argyrostilba (O. P.-Cambridge, 1876), G. dentata (Zhu & Wen, 1978) and G. tangi (Zhu, Song & Zhang, 2003), all previously included in Dyschiriognatha. The following taxa are newly synonymized: Dyschiriognatha montana Simon, 1897, Glenognatha mira Bryant, 1945 and Glenognatha maelfaiti Baert, 1987 with Glenognatha argyrostilba (O. P.-Cambridge, 1876) and Glenognatha centralis Chamberlin, 1925 with Glenognatha minuta Banks, 1898. A key to species of Glenognatha and distribution maps are provided. RESUMO A revisão taxonômica e análise cladística do gênero Glenognatha Simon, 1887 é apresentada. A análise é baseada em uma matriz que inclui 24 espécies de Glenognatha, oito representantes do grupo externo, incluindo três gêneros da subfamília Tetragnathinae e um de Metainae, e 82 caracteres morfológicos. Oito sinapomorfias não ambíguas sustentam a monofilia de Glenognatha, todas não homoplásticas. Alguns clados internos dentro do gênero são bem suportados e suas relações são discutidas. O gênero Glenognatha apresenta uma ampla distribuição ocupando as regiões Neártica, Neotropical, Afrotropical, Indo-Malaya, Australasia e Oceania. Vinte e sete espécies de Glenognatha são reconhecidas, quatro delas somente conhecidas por machos. Novos dados morfológicos são fornecidos para a descrição de treze espécies previamente conhecidas. Onze espécies novas são descritas: G. sp. nov. 1, G. sp. nov. 3, G. sp. nov. 4 e G. sp. nov. 7 do sudeste do Brasil, G. sp. nov. 6, G. sp. nov. 9 e G. sp. nov. 10 da região Amazônica, G. sp. nov. 2, G. sp. nov. 5 e G. sp. nov. 8 do norte da cordilheira dos Andes e G. sp. nov. 11 da região central do México e sul dos Estados Unidos. As fêmeas de G. minuta Banks, 1898, G. gaujoni Simon, 1895 e G. gloriae (Petrunkevitch, 1930) e os machos de G. globosa (Petrunkevitch, 1925) e G. hirsutissima (Berland, 1935) são descritos por primeira vez. São propostas três novas combinações em congruência com os resultados da análise cladística: Glenognatha argyrostilba (O. P.- Cambridge, 1876), Glenognatha dentata (Zhu & Wen, 1978) e Glenognatha tangi (Zhu, Song & Zhang, 2003), todas previamente incluídas no gênero Dyschiriognatha. Quatro sinonímias são propostas: Dyschiriognatha montana Simon, 1897, Glenognatha mira Bryant, 1945 e Glenognatha maelfaiti Baert, 1987 com Glenognatha argyrostilba (O. P.-Cambridge, 1876) e Glenognatha centralis Chamberlin, 1925 com Glenognatha minuta Banks, 1898. É apresentada uma chave para a identificação e mapas de distribuição para as espécies. INTRODUCTION Tetragnathidae The orb-weaving spider family Tetragnathidae currently includes 47 genera and 947 species (Platnick, 2013). This family is particularly diverse in the Neotropical region which has 47% of the described genera. Most members of the family build horizontal orbicular webs on vegetation adjacent to rivers or streams (Levi, 2005), yet the group presents great morphological and ethological diversity (Levi, 1980; Eberhard, 1982; Hormiga et al., 1995; Álvarez-Padilla et al., 2009). Although tetragnathids have received increasing attention from systematic biologists in the last ten years, there is still great gaps of knowledge concerning the real richness of the family; probably only half or fewer of the species of Tetragnathidae have been described, as is the case in many other spider families (Alvarez-Padilla & Hormiga, 2011). Recent phylogenetic analyzes (Álvarez-Padilla, 2007; Dimitrov & Hormiga, 2009, Álvarez-Padilla et al., 2009; Alvarez-Padilla & Hormiga, 2011; Dimitrov & Hormiga, 2011), support the monophyly of Tetragnathidae by the following synapomorphies: copulatory duct length less than half of the spermathecae length, femora IV mesal surface without basal macrosetae, male palpal patella with one macrosetae, male palpal tibia length approximately two times the widest point of the tibia, piriform gland spigot bases separated from the spigot shaft leaving a torus, tegulum oval to spherical and cymbial tarsal organ larger than the bases of surrounding macrosetae. Based mainly on genitalia characters tetragnathids can be divided into four subfamilies: Tetragnathinae, Leucauginae, Metainae and Nanometinae (Álvarez-Padilla et al., 2009; Álvarez-Padilla & Hormiga, 2011). The taxonomic composition of these subfamilies (except Metainae) has been rather stable in phylogenetic analysis regardless of data partitions and analytical approaches (Álvarez-Padilla & Hormiga, 2011). The subfamily Tetragnathinae which includes the genera Antillognatha Bryant, 1945; Cyrtognatha Keyserling, 1881; Doryonychus Simon, 1900; Dyschiriognatha Simon, 1893; Glenognatha Simon, 1887; Hispanognatha Bryant, 1945; Tetragnatha Latreille, 1804 and Pachygnatha Sundevall, 1823, is particularly interesting among tetragnathids due to the evolution of a secondary haplogyne condition, in which fertilization ducts have been lost and there has been an overall simplification of female genitalia (Dimitrov et al., 2007). Taxonomic revisions of tetragnathid genera with species distributed in the Neotropical region are scarce. In the case of Tetragnathinae, the only current revision of a group distributed in the Neotropics, was made by Dimitrov & Hormiga (2009) for the genus Cyrtognatha. These authors established eight new combinations, described 11 new species and proposed Agriognatha O. P. Cambridge, 1896 as a junior synonym of Cyrtognatha. In addition, the monophyly of Cyrtognatha was tested and its position within Tetragnathinae discussed. The results obtained in this study highlight the importance of conducting systematic studies within Tetragnathinae. Taxonomic history of the genus Glenognatha The genus Glenognatha was erected by Simon (1887) with the type species G. emertoni Simon 1887, based on a male specimen collected in Arizona. Simon (1887) recognized the affinity of this species with species of the genus Pachygnatha, but stated that the well-developed tracheal spiracle, analogous with Anyphaena Sundevall, 1833, distinguished this new species from Pachygnatha. Before the erection of Glenognatha, Keyserling (1883) described the species Pachygnatha australis from males and females collected in Pacasmayo, Peru, which were subsequently transferred to Glenognatha by Simon (1894). After these pioneers works, in the late nineteenth century and the first half of the twentieth century, 15 Glenognatha species were proposed. Seven of these species were described in Islands; five in French Polynesia: G. nigromaculata (Berland, 1933); G. hirsutissima (Berland, 1935); G. argenteoguttata (Berland, 1935); G. chamberlini (Berland, 1942) and G. phalangiops (Berland, 1942), and two in Caribbean islands: G. gloriae (Petrunkevitch, 1930) (Puerto Rico) and G. mira Bryant, 1945 (Hispaniola). The eight remaining species were described in Argentina (G. lacteovittata (Mello-Leitão, 1944)), Ecuador (G. gaujoni Simon, 1895), United States of America (G. foxi (McCook, 1894)), Guyana (G. caporiaccoi Platnick, 1993), Mexico (G. minuta Banks, 1898 and G. spherella Chamberlin & Ivie, 1936) and Panama (G. centralis Chamberlin, 1925 and G. globosa (Petrunkevitch, 1925)). Twelve of the previously named species are known only from the type locality. Moreover, several species were initially described in other genera such as Dyschiriognatha (G. nigromaculata), Hivaoa Berland, 1935 (G. nigromaculata, G. hirsutissima, G. argenteoguttata, G. chamberlini and G. phalangiops), Diplocephalus Bertha, 1883 (G. foxi and G. gloriae), Mimognatha