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Cladistic Analysis of the Afrotropical Hersiliidae (Arachnida, Araneae) with the First Records of Murricia and the Description of a New Genus from Madagascar

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Cladistic Analysis of the Afrotropical Hersiliidae (Arachnida, Araneae) with the First Records of Murricia and the Description of a New Genus from Madagascar 0940-08_JAZ 2008-4_09 09-04-2008 13:34 Pagina 111 Cladistic analysis of the Afrotropical Hersiliidae (Arachnida, Araneae) with the first records of Murricia and the description of a new genus from Madagascar S.H. FOORD ABSTRACT FOORD, S.H. 2008. Cladistic analysis of the family Hersiliidae (Arachnida, Araneae) of the Afrotropical Region, with the description of a new genus from Madagascar. J. Afrotrop. Zool. 4: 111-142. This study infers a phylogenetic hypothesis for the family Hersiliidae from the Afrotropical Region. A new genus is pro- posed, Prima gen. n., to include P. ansiae sp. n. from Madagascar. The first record of the genus Murricia Simon from the Afrotropical Region is presented, with the description of a new species, Murricia uva sp. n. from Central Africa. The species level phylogenetic analysis comprises 43 taxa, of which 27 are Afrotropical hersiliids with both sexes known, 13 species non-Afrotropical and three outgroup species; 66 characters were scored. The analysis resulted in 56 most parsimonious trees. The preferred tree supports the monophyly of Tyrotama Foord & Dippenaar-Schoeman, 2005, and retrieves it as the sister-group of Hersiliola Thorell, 1870. These two genera form the most basal lineage of the family Hersiliidae. Tama Simon, 1882, is the sister-group of all the remaining hersiliids, again divided in two large lineages. The first of these clades comprises all arboreal hersiliids without biarticulations on the metatarsi I, II and III. Neotama is basal in this clade and the new genus Prima part of its sister-group (Neotama (Tamopsis (Prima (Ypypuera (Iviraiva + Yabisi))))). The second clade consists of two sister clades viz. Hersilia, whose monophyly is confirmed, and the genus Murricia.The family Hersiliidae now includes six genera in the Afrotropical Region, viz. Hersilia, Hersiliola, Murricia, Neotama, Tyro- tama Foord & Dippenaar-Schoeman, 2005, and Prima n.gen. A key to the Afrotropical genera is provided. S.H. FOORD, Department of Biological Sciences, University of Venda, Private Bag X5050, Thohoyandou, 0950 South Africa ([email protected]) Keywords: phylogeny, hersiliid, Hersilia, Neotama, Tyrotama, Hersiliola INTRODUCTION genus Tamopsis Baehr & Baehr, 1987. In this work, 29 species, of which 93 % were new to science, Hersiliidae is a small spider family with 141 species were either described or redescribed. Subsequent and 11 genera excluding the results from this study publications doubled the number of species known (Platnick 2007, Rheims & Brescovit 2004). The fam- for the Australian Region to 55 in total (Baehr & ily is characterised by conspicuously long posterior Baehr 1988b, 1989, 1992, 1993b, 1995). In their lateral spinnerets, elongated legs and is limited to the revision of the Oriental Region, Baehr & Baehr tropical and subtropical regions of the world. All her- (1993a) described 27 new species out of a total of siliids are arboreal except for the representatives of 33 taxa and proposed a tentative phylogeny for the Hersiliola Thorell, 1870, Tama Simon, 1882 and Tyro- family. Rheims & Brescovit (2004) described six new tama Foord & Dippenaar-Schoeman, 2005. These species, redescribed five species, and proposed three genera are all ground active, inhabiting arid regions new genera, Yabisi Rheims & Brecovit, 2004, Iviraiva of Africa, the Mediterranean and the western parts Rheims & Brecovit, 2004, and Ypypuera Rheims & of the Oriental Region (Smithers 1945, Levy 2003). Brecovit, 2004, for the Neotropical and Nearctic The family has recently been the focus of extensive Regions. Once again all the Neotropical and Nearc- revisions in the Australian (Baehr & Baehr 1987), tic representatives of Tama were transferred to the Oriental (Baehr & Baehr 1993a; Chen 2007), latter three genera and to Neotama Baehr & Baehr, Neotropical and Nearctic Regions (Rheims & 1993. Brescovit 2004), and the Afrotropical Region (Foord Members of the family Hersiliidae are found & Dippenaar-Schoeman 2005a, 2005b; Foord & throughout the Afrotropical Region and include both Dippenaar-Schoeman 2006). Prior to these revisions, arboreal (Foord & Dippenaar-Schoeman 2005b, Tama was the most widely distributed genus in the 2006) and ground active genera (Foord & Dippenaar- family. However, Baehr & Baehr’s 1987 revision of Schoeman 2005a). Descriptive work was done Australian hersiliids resulted in the transfer of most between 1827 and 1976. Smithers (1945) revised the species previously attributed to Tama to the new three genera in southern Africa and Benoit (1967) 111 0940-08_JAZ 2008-4_09 09-04-2008 13:34 Pagina 112 S.H. FOORD revised the genus Hersilia Audouin, 1826, of the uncertain. Recent work on fossil spiders suggests that Afrotropical Region excluding Madagascar. None of the family is an ancient taxon, and predicts the pres- these studies attempted to determine phylogenetic ence of Hersiliidae in the early Cretaceous (Penney relationships of the hersiliid taxa. Before this study 2002). Fossil evidence also points to a much wider the Afrotropical Region included records of 43 distribution in Europe during the Oligocene and species in four genera, Hersilia (31 species), Hersili- Miocene (Penney 1999; 2006). Lehtinen (1967) con- ola Thorell, 1870 (three species), Neotama (one sidered the Hersiliidae to be related to the Oecobi- species) and Tyrotama (eight species). All Afrotrop- idae based on similarity of genital structures, modi- ical species in the genus Tama have been transferred fication of the carapace, eye pattern and spinnerets. to the new genus Tyrotama (Foord & Dippenaar- Coddington & Levi (1991), in a cladistic analysis of Schoeman 2005a). Only Tama edwardsi (Lucas, 1846) all spider families, grouped the hersiliids with the remains in Tama and is restricted to the Mediter- oecobiids based on the similarity of the elongated ranean and the Palearctic Regions. posterior lateral spinnerets and the distinct prey cap- ture behaviour of the two families. While facing away, Both Tyrotama and Hersiliola occur in the warmer the spider rapidly encircles the prey and covers and parts of the Afrotropical Region, where they are found enswathes it with silk from the long posterior lateral under stones (Smithers 1945). Hersiliola construct spinnerets. Although Coddington & Levi (1991) a pholcid-like web and Tyrotama a circular retreat of placed the hersiliids in the Eresoidea (Hersiliidae, closely woven webbing plastered with small pebbles, Oecobidae and Eresidae), basal within the Entelege- chips and vegetable debris. Anchor threads attached nae, representative species of Hersiliidae were not to the substratum warn the spider of approaching included in their compilation of data. prey (Smithers 1945; Dippenaar-Schoeman et al. Baehr & Baehr (1993a) did not consider these rela- 1999; Jocqué & Dippenaar-Schoeman 2006). They tionships well supported and based their phyloge- move at great speed, overpower their prey and drag it netic analysis on morphoclines. Baehr & Baehr’s back to their retreat where they feed (Lawrence 1964). (1993a) attempt at inferring phylogenetic relation- Their egg sacs are attached to the underside of rocks ships for hersiliids, included representatives of the and covered with stone chips (Dippenaar-Schoeman family from Australia, the Oriental Region and New & Jocqué 1997). The absence of ground-dwelling Guinea. They subsequently also proposed a phy- representatives from the Oriental, Neotropical and logeny for Hersilia from Australia and New Guinea Australian regions is peculiar. Penney (2006) sug- (Baehr 1998). Their hersiliid phylogeny has Hersiliola gested that Yabisi guaba Rheims & Brescovit, 2004, basal, based on the following plesiomorphic charac- might be a ground wanderer. However, this is refuted ters: short legs, shorter posterior lateral spinnerets, by the capture of specimens in the Mangrove Islands a convex opisthosoma and no flexible zone on leg off the Florida Keys (Simberloff 1971). metatarsi. Tama is defined by longer legs, long pos- Hersilia is the most widely distributed genus in the terior lateral spinnerets as well as the presence of a family. Platnick (2007) catalogues the Afrotropical, flexible zone on the metatarsi of legs I, II and IV, Oriental and Australian Regions, and one species that whereas Hersilia has a biarticulate metatarsus, very penetrates into the Palaeartic. All Afrotropical species long I, II, and IV legs and armed chelicerae. Rheims of this genus live on tree trunks. They are well cam- and Brescovit (2004) suggested the use of Uroctea ouflaged with their flattened bodies pressed against Dufour, 1820, as an outgroup based on the proposal the bark, and when disturbed they move at great by Lehtinen (1967) that Hersiliidae is a sister group speed. Although specimens are very cryptic, the of ecribellate Oecobiids belonging to the genus reflection of sunlight on the guide threads left on Uroctea and followed the principles set forth by Nixon trunks is a good indicator of their presence (Dippe- & Carpenter (1993) for outgroup selection. In spite naar-Schoeman & Jocqué 1997). Females cover their of the differences in approach, their proposed phy- eggs with an oval-shaped layer of silk camouflaged logeny corresponds with that of Baehr & Baehr with bits of bark. They stand guard over the eggs in (1993a) except for placing Neotama plus Hersilia as an upside-down position above the silk patch until the sister-group to Tamopsis. the young hatch (pers. obs.). Baehr & Baehr (1993) suggested that the hersiliids Neotama is represented in the Afrotropical Region
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