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Cladistic analysis of the Afrotropical Hersiliidae (Arachnida, Araneae) with the first records of Murricia and the description of a new 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 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. is basal in this clade and the new genus Prima part of its sister-group (Neotama ( (Prima (Ypypuera (Iviraiva + ))))). The second clade consists of two sister clades viz. , 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 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 , 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)

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revised the genus Hersilia Audouin, 1826, of the uncertain. Recent work on fossil 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 (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- (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 (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 (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 . 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 by have an African origin based on the principle that one species, Neotama corticola (Lawrence 1937), plesiomorphy and place of origin should correspond. which is restricted to the south-eastern region of The presence of the plesiomorphic hook-shaped South Africa (Foord & Dippenaar-Schoeman 2005b). median apophysis found on the male palp in some Petrunkevitch (1963), Schawaller (1981) and Wun- Afrotropical Hersilia species and the occurrence of derlich (2004) have described several fossil genera the ground dwelling genera of Hersiliola in Africa whose relationships with extant Hersiliidae remain points in that direction. However, Penney (2006) has

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questioned this assertion with reference to fossil evi- (see Rheims et al. 2004). Species from North Africa dence of a much wider distribution of hersiliids (countries outside the Afrotropical Region) are only throughout the northern hemisphere and invoked the included when such species also occur in the alternative hypothesis of ousted relics (Eskov & Golo- Afrotropical Region. vatch 1986). This would suggest that the current dis- Slide preparation: The epigyne of the female and the junct ‘Gondwanan’ distribution of hersiliids is the left palp of the male (when available, otherwise the result of a formerly worldwide distribution followed right) were removed and mounted temporarily on by extinction of the northern hemisphere represen- slides in Heinze’s modified PVA mounting medium tatives (Penney 2006). (Smith-Meyer & Rodrigues 1966). References to The taxonomic status of two Oriental genera, Murri- illustrations in this publication are capitalised cia Simon, 1882, and Promurricia Baehr & Baehr, whereas references to figures from other papers are 1993 has also been doubted (Rheims & Brescovit noted in lower case. 2004). Murricia Simon is a small genus in the family Illustrations: All illustrations were made with a stere- Hersiliidae, only known from females, with three omicroscope Zeizz Stemi SV 6, using a camera species that have been recorded from India, Sri Lanka lucida. Scanning electron micrographs were taken and Singapore, respectively (Platnick 2007). The with a JEOL (JSM 840) microscope. genus is characterised by the lateral eyes that are on Abbreviations: The following abbreviations are used distinct eye tubercles, the markedly triangular in this paper: opisthosoma and the second pair of dorsal muscular ALE – anterior lateral eye; AME – anterior median pits that are transverse (Baehr & Baehr 1993). Little eye; bS – basal segment of posterior lateral ; is known about their natural history except that they ebp – basal embolar process; c – copulatory duct; CAI are arboreal forest dwellers. Baehr & Baehr (1993) – carapace index; ci – consistency index; CL – cara- discussed their origin and possible relationship with pace length; CLL – clypeus length; CW – carapace Promurricia Baehr & Baehr from Sri Lanka. The pos- width; el – epigyne length; ew – epigyne width; Fe – sibility existed that Murricia is a junior synonym of femur; MOQ – median ocular quadrangle; MOQ- Hersilia with Promurricia intermediate between the AW – MOQ anterior width; MOQ-PW – MOQ pos- first two genera (Baehr & Baehr 1993; Rheims & terior width; MOQL – MOQ length; Mt – metatar- Brescovit 2004). Baehr & Baehr (1993) suggested sus; Pat – patella; PER – posterior eye row; PLE – that Murricia originated from ground-living hersili- posterior lateral eyes; PME – posterior median eyes; ids based on the genus’s restricted distribution in OAL – ocular area length; ri – retention index; Ta – the Oriental Region and the synapomorphy that tarsus; Tib – tibia; tS – terminal segment of poste- Promurricia shares with Murricia, namely the medio- rior lateral spinneret. posterior part of the female epigyne having a wide, Measurements: Measurements were made under a oval median plate slightly separated from the lateral stereomicroscope using an ocular micrometer with up lobes (Baehr & Baehr 1993). to 50× magnification. All measurements are given in The Afrotropical Region represented the last region millimetres with the observed ranges in parentheses. for which no contemporary revision was available. Material was received on loan from the following Such a revision would clarify generic delimitations institutions: in the family Hersiliidae (e.g. Hersilia, Murricia AMNH – American Museum of Natural History, Simon and Promurricia Baehr & Baehr) as well as New York, USA (N.I. Platnick); BMNH – The Nat- the phylogenetic relationships of the newly described ural History Museum, London, UK (J. Beccaloni); genus Tyrotama. This paper therefore attempts to CAS – California Academy of Sciences, Golden Gate reconstruct Afrotropical hersiliid phylogeny based Park, San Francisco, California, USA (C. Griswold); on a revision that included 46 Afrotropical hersiliid CDO – Collection C. Deeleman, Ossendrecht species (Foord & Dippenaar-Schoeman 2005a; (C. Deeleman); DNSM – Durban Natural Science Foord & Dippenaar-Schoeman 2005b; Foord & Museum, Durban, South Africa (T. Crouch); MCZ – Dippenaar-Schoeman 2006). Museum of Comparative Zoology, Cambridge, Mass- achusetts (G. Giribet and L. Leibensperger); MNHU – Museum für Naturkunde der Humboldt Universität, METHODS Berlin, Germany (J. Dunlop); MHNG – Muséum d’Histoire naturelle, Genève (P.J. Schwedinger), Study area: The area covered by this study is the MNHN – Museum National d’Histoire naturelle, Afrotropical Region, including the following islands Paris, France (C. Rollard); MRAC – Koninklijk in both the Indian and Atlantic Oceans: St. Helena, Museum voor Midden-Afrika, Tervuren, Belgium Comoros, Madagascar, the Seychelles, Zanzibar, (R. Jocqué); MSNG – Museo Civico di Storia Aldabra Islands and Cape Verde but excluding Yemen Naturale Giacomo Doria, Genova (G. Doria); MZBS

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– Museo de Zoologia, Universidad de Barcelona, – Neotropical, Ypypuera crucifera (Vellard) – Neotrop- Barcelona (C. Ribera); NCA – National Collection of ical, Yabisi habanensis (Franganillo) – Neotropical. Arachnida, Agricultural Research Council – Plant The outgroup consisted of Uroctea durandi (Latreille, Protection Research Institute, Pretoria, South Africa 1809) from the Mediterranean, Stegodyphus dumi- (A.S. Dippenaar-Schoeman); NM – Natal Museum, cola Pocock, 1898 (1 Ω 3 æ, NCA) from South Africa Pietermaritzburg, South Africa (D. Jennings); NMB and Dresserus colsoni Tucker, 1920 (2 Ω 2 æ, NCA) – National Museum, Bloemfontein, South Africa based on Lehtinen (1967), Coddington & Levi (L. Lotz); NRM – Swedish Museum of Natural His- (1991) and Rheims & Brescovit (2004). All species tory (T. Kronestedt); QM – Queensland Museum, for which one of the sexes was unknown were Brisbane, Australia (B. Baehr); SAM – Iziko Museum excluded from the analysis. of Cape Town, South Africa (M. Cochrane); NMN – The preferred tree must be the shortest, most resolved National Museum of Namibia, Windhoek, Namibia tree possible, i.e. make the most predictions that can (T. Bird); SMF – Forschungsinstitut und Naturmu- be tested given only unambiguous branch support seum Senckenberg, Federal Republic of Germany (Griswold & Ledford 2001). Preference is, therefore, (P. Jaeger); TMSA – Northern Flagship Institute (Trans- given to synapomorphic characters in the construc- vaal Museum), Pretoria, South Africa (K. Manamela). tion of a phylogeny as they have the largest predictive A data matrix comprising 40 hersiliid species was power (Farris 1985: in Griswold & Ledford 2001). constructed. This data set included 27 Afrotropical Analyses of the equally weighted and unordered char- species and 13 hersiliid species from all the other acters were subjected to heuristic searches in Nona geographical regions where hersiliids are known to 2.0 (Goloboff 1997) using h/1000, mult*1000, occur. Three non hersiliid species were included in jump50. Equally weighted characters cannot be con- the outgroup. The selection of Afrotropical species sidered unweighted and weighting will give more was based on recent revisions (Foord & Dippenaar- weight to characters more consistent with initial Schoeman 2005a; Foord & Dippenaar-Schoeman heuristic cladograms (Farris, 1969: in Bosselaers & 2005b; Foord & Dippenaar-Schoeman 2006). Jocqué 2001). Successive weighting with the squared Species for which only one sex are known were consistency index was therefore done in NONA enter- excluded from the analysis. This resulted in the inclu- ing the following command sequence: run swt.run sion of 27 Afrotropical hersiliid species (Appendix 1). h10000 h/1000 mult*1000 jump 50. Pee-Wee 2.6 Where possible, character variation based on the (Goloboff 1997) was used to obtain trees that max- examination of type specimens in the following non- imise implied weights across all characters (Goloboff Afrotropical taxa were included in the matrix. The 1993a), with all characters equally weighted (wt = 1) Oriental Region: Hersilia albumaculata Wang & Yin, and for both ordered and unordered multi-states 1 Ω from Hong San, S.E. Kiangsi, China, MCZ (h10000;h/1000;mult*1000;jump50;). Pee-Wee max- æ 67908; Hersilia tibialis Baehr & Baehr: 2 from Anu- imises total fit F=∑fi where fi = k/(k+Esi); k is a con- radhapura, Ceylon, SMF 38432 (4 æ, SMF), Hersilia stant of concavity and can be varied between 1 and savignyi Lucas: 5 Ω and 3 æ from Kandy, Ceylon, SMF 6 with the ‘concN’ command, allowing less (1) or 38429 (5 Ω 3 æ, SMF), Hersilia asiatica Song & more (6) influence for homoplasious charactes. Con- Zheng: 2 Ω and 2 æ from Tsin Leong Sam, E. Kwan- cavity constants 1, 3 and 6 were used in this analysis. tung, China, MCZ 67899 & 67900 & 67911; Phylogenetic signal for equally weighted trees was Hersilia sundaica Baehr & Baehr: 1 Ω, 1 æ from assessed with branch support indices (Bremer 1994). Djojosudharmo, Lombok, 12-14.i.1990, Koze Branch support was calculated with NONA 2.0 kansika, CDO (1 Ω 1 æ, CDO) Hersilia flagilifera (Goloboff 1997) using the options h10000; bsup- Baehr & Baehr: 2 Ω and 1 æ paratypes from Ketambe, port 100. The Bremer Support for a given node in Gn. Leuser, Nothern Sumatra, 20.ii.1985, Suharto, the shortest unconstrained tree is the number of CDO; Hersilia kinabaluensis Baehr & Baehr: Ω holo- additional steps required in the shortest trees for type and æ paratype from Kinabalu National Park, which that node collapses (only for unweighted Northern Borneo, ix.1980, C. Deeleman, MHNG; analysis). A branch present in one of the most Neotama variata (Pocock): 1 Ω from Dambulla, Sri parsimonious trees is more strongly supported by Lanka, G. Schmidt, xii.1981, SMF 36782. Palearc- the data if a large increase in length of additional tic:. Tama edwardsi Simon: 1 Ω, 1 æ from Ramble del trees is required before that branch is lost in the con- Pouis, Sierra de Carrascoy, Sangonera, Spain, sensus (Griswold & Ledford 2001). Branch support 1.iv.1995, rocky wall, J. Miñano, MZBS. Australia: was truncated at values of 100 and Bremer support Tamopsis eucalypti Baehr & Baehr, 1 Ω and 1 æ, QM. values reported range from 0 and greater. Bremer The following four species were coded with refer- support values for successive weighting trees were ence to their descriptions Hersilia jajat Rheims & obtained in Nona 2.0. Tree editing was done in Win- Brecovit – Malaysian, Ivivaiva pachyura (Mello-Leitão) clada 1.0 (Nixon 2002).

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DISCUSSION CI = 52; RI = 80) and five nodes collapsed in the strict consensus of these trees. The number of trees found With six genera, the Afrotropical Region now has under implied-weighting in PeeWee was constant the highest generic level diversity of all biogeographic with 8 trees for k = 3 (l = 201; CI = 52, RI = 80) regions. Heuristic searches of the matrix in NONA (Fig. 1a), 8 trees for k = 1 (l = 204; ci = 51, ri = 79) with characters equally weighted resulted in 56 most and 8 trees for k = 6 (l = 201, CI = 52, RI = 80). Trees parsimonious trees (l = 200; CI = 52; RI = 80). Twelve constructed with different character weighting pro- nodes collapse in the strict consensus of these trees cedures are largely congruent except for relationships (Fig. 1b). Successive weighting of the unordered multi- within the genus Hersilia and more specifically clade state matrix in NONA, generated 7 trees (l = 200; 17. Relationships and support for this clade itself is

Figure 1a. Preferred, strict consensus cladogram for 40 Hersiliidae species obtained under implied weighting (l = 200, ci = 52, ri = 79) collapsing five nodes. State changes that require only the minimum number of steps on the tree are indicated in black, state changes that require additional steps are white. Nodes are numbered on the tree. Values above branches are Bremer support values (in PeeWeefit values). Species in parenthesis are not Afrotropical.

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Figure 1b. Strict consensus of 42 most parsimonious trees found under equal weighting, with unambiguous support, for 40 Hersiliidae species. Figures above branches are Bremer support values (in steps). Species in paranthesis are not from the Afrotropical Region.

low and confirmed by the alternate relationships pro- & Jocqué 2000) (Fig. 1). Character states are found posed by different weighting procedures. They dif- in Appendix 1. All characters discussed are uniquely fered in the relationships and placement of clade 18 derived unless stated otherwise. and the extent to which the relationships of H. asiat- The proposed phylogeny is largely congruent with ica and H. saignyi, and H. madagascariensis and H. kin- that of Baehr & Baehr (1993a) and Rheims & abaluensis are resolved. The weak support for latter Brescovit (2004) except that the genus Tyrotama, not relationships are also confirmed by low Bremer sup- included in their analyses, is a sister clade to Hersili- port values (Figs 1a-b). Three clades are, however, ola. A basal dichotomy in the ingroup separates retained in all of the trees viz. clade 15, 18 and 22. clade 2, consisting of Hersiliola and Tyrotama, from The trees based on implied weighting (k = 3) are pre- all other hersiliids and is characterised by longitudi- ferred as these trees save steps for the less homopla- nal grooves dorsally on leg setae (character 8 [1]), sious characters (Platnick et al. 1991: in Bosselaers very small inconspicuous sigilla (character 15 [1]),

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and the construction of a retreat (character 22[1,2]). between the lateral borders and median plate of the Clade 3 is a very distinct subclade within clade 2, epigyne (character 46[1]). A narrow flexible zone composed of representatives of the genus Tyrotama on metarsi I, II and IV (character 4[2]), is a feature that are restricted to the arid regions of southern that is paralleled in the genus Ypypuera. Neotama Africa and firmly supported by several synapomor- has the widest distribution of all hersiliid genera phies: the construction of a circular retreat that (Afrotropical, Neotropical and Oriental). The pres- incorporates bits of debris (character 22[1]), a ence of Neotama in the Neotropical Region could sperm duct with a variable number of inward and be explained by the theory of ousted relics (Eskov outward bends (character 32[6]), an epigynal atrium & Golovatch 1986). Based on this theory Penney (character 52[1]) and the presence of a large hya- (2006) suggested that Hersiliidae could have had a line seminal receptacle (character 64[4]). This con- pancontinental distribution based on the presence firms Baehr & Baehr’s (1993) suggestion that the of fossil representatives that predicts a much wider Afrotropical representatives of Tama sensu lato are distribution of the family than at present. not related to Tama, sensu stricto. As such, the new Tamopsis (Australia) is basal in clade 8 with Prima genus Tyrotama was described to include Hersiliola gen. nov. (Madagascar), Ypypuera (Neotropical australis, H. fragilis, Tama arida, T. bicava, T. incerta Regiona), Iviraiva and Yabisi (Neotropical Region) and the new species T. abyssus, T. soutpansbergensis sequentially branching off. The new genus Prima is and T. taris (Foord & Dippenaar-Schoeman 2005a). characterised by two synapomorphies: a sperm duct, The remaining hersiliids are united at clade 4 by starting from the base of the embolus, basally with a leg IV that is less than 0.4× the length of leg I (char- hairpin outward bend followed by a bend inwards acter 2[1]), the presence of some form of articular medially (character 32[5]) and a bifid median apoph- modification on metatarsi I, II and IV (charac- ysis (character 36[1]). ter 4[1,2,3]), metarsus I more than 8× the length Clade 12 is the sister clade of clade 6 and consists of of tarsus I (character 6[1]), a tibial palp that is all hersiliids that have a biarticulation on metatarsi I, longer than the palpal patella (character 25[1]), the II and IV (character 4[3]). This clade has two sister median plate of the epigyne that is partially covered clades. The first of these, clade 13, is characterised by by the lateral borders (character 47[0]) and the pres- a sperm duct that, starting from the base of the ence of cylindrical seminal receptacles (charac- embolus, has an inward bend medially followed by a ter 65[4]). Tama edwardsi, in the monotypic genus large outward bend, that in turn is followed by a Tama, is a ground-dwelling species restricted to the medial inward bend and a basal loop (charac- Mediterranean and share all these synapomorphies ter 32[4]), an oval epigynal plate (character 51[1]) with arboreal hersiliids, suggesting that it was and an elongate, circular looped copulatory duct preadapted to tree living (Rheims & Brescovit 2004). (character 56[1]). There is a single representative of Clade 5 unites all arboreal hersiliids that are charac- this clade in the Afrotropical Region and although terised by the terminal segment of the posterior lat- Baehr & Baehr (1993) did not describe any males for eral spinnerets more than 3× the length of the basal the genus Murricia this species has several characters segment (character 18[2]) and the absence of a in common with species in this genus from the Ori- retreat (character 21[1]). ental Region viz. lateral eyes that are on a distinct eye Arboreal hersiliids are divided into two large sister tubercle, large posterior lateral eyes, a triangular to clades. Both of these clades’ support is rather low trapezoid abdomen, and a elongate complexly coiled as evidenced by their Bremer support (Figs 1a-b). seminal receptacle. Rheims & Brescovit (2004) The first of these, clade 6, groups six genera distrib- described Hersilia jajat from Borneo and although uted throughout the tropical and subtropical areas of the species does not share the latter characteristics the world and is characterised by several homopla- with Murricia, synapomorphies as listed for clade 12 sious characters, including sperm ducts with a sin- and proposed for the genus Murricia in this study is gle basal loop (character 32[2]) that is secondarily present in Hersilia jajat. lost or modified in Prima, Ypypuera and Yabisi; an Clade 14, the sister taxon of clade 13, includes all embolus that originates basally on the bulbus (char- representatives of Hersilia except Hersilia jajat. All acter 40[2]) that reverses to an apical origin in the species in this genus have spermathecae and sem- clade 9 and cylindrical spermathecae (charac- inal receptacles that are clearly separated from the ter 61[1]) that reverses into spherical spermatheae copulatory and fertilisation ducts (character 63[1]) in clade 10. Neotama (clade 7) is basal in this clade, and the presence of a distal tubercle or glandular and the monophyly of this genus is supported by two area on the seminal receptacle (character 66[1]). synapomorphies: a palpal tibia that is more than Clade 12 that unites Murricia and Hersilia has less twice as long as the palpal patella (character 25[2]), Bremer support than clade 13. The genus Murricia and the presence of oval slit-like lateral openings is therefore retained. Furthermore, it is obvious from

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the low Bremer support values for clade 17, and the CHARACTER DESCRIPTIONS uncertain placement of clade 18 that it would not AND INTERPRETATIONS be advisable to split the genus. However, clades 15, 18 and 25 are well supported. Character 1 Plumose setae (0) absent; (1) pres- Based on the strict consensus of eight trees found ent (Fig. 2). Griswold et al. (1999: 56, character 18) under implied weighting, Hersilia consists of two considered the presence of plumose setae as derived. clades, 15 and 17. Clade 15 comprises eight species This character is a synapomorphy for all hersiliids at that are characterised by the median plate of the clade 1 (Fig. 1a). epigynum with lateral sclerotised projections that LEGS presumably act as fixing structures during copula- tion (character 48[1]) and lateral borders with trian- Character 2 Relative length of leg III to leg I gular projections (character 49[1]). Species in this (0) = more than 0.6; (1) less than 0.4. The cladogram clade are widely distributed throughout the suggests that a long leg III is plesiomorphic and Afrotropical Region. Two species, H. sericea and H. relative shortening of leg III is derived. There is, setifrons from South Africa, are sympatric and syn- however, uncertainty whether it was owing to a short- topic in the savannah and have never been collected ening of leg III or lengthening of leg I, II and IV in forests. This seems to be true for all the other (Baehr & Baehr 1993a; Rheims & Brescovit 2004). species in this clade except the enigmatic H. sigillata The relative shortening of leg III is a synapomorhy for that has been collected in forests of Central Africa. the clade of all arboreal hersiliids and Tama edwardsi The core of clade 15, clade 16, is characterised by the (clade 4, Fig. 1a). presence of glands basally on the spermathecae Character 3 Relative length of leg IV, legIV/legI (character 59[1]) and kidney-shaped seminal recep- (0) less than one; (1) more than one. tacles (character 65[2]). The clade comprises three The cladogram suggests that an elongate leg IV species, two of which are endemic to Madagascar. was derived at the node that unites Uroctea and The third species, H. sigillata, is peculiar, and has Hersiliidae and reverted back to a relatively shorter several autapomorphies that include a hook-shaped leg IV in clade 4 (Fig. 1a) that includes Tama and all median apophysis that is distally truncate (charac- arboreal hersiliids. ter 37[2]), a very short, truncate cymbium (charac- Character 4 Metatarsi articulation of leg I, ter 30[0]), first three pairs of sigilla, very large and II and III (0) uniarticulate; (1) elongate flexible zone similar in size (character 15[2]), the interdistance (Fig. 28); (2) short flexuose zone; (3) biarticulate of AME-ALE is more than twice that of AME diam- (Fig. 27). Baehr (1998) suggested a possible evolu- eter (character 11[1]), a characteristic shared with tionary sequence in which an elongate flexible zone the genus Yabisi from the Neotropical Region, and is reduced in length to form a narrow flexible zone lateral eyes that are on prominent tubercles (char- and ultimately develop into a biarticulation. The elon- acter 12[2]), which is also shared with Yabisi and gate flexible zone is derived in clade 4 (Fig. 1a). Murricia. A narrow flexible zone evolved independently in Clade 17 is the second and largest clade in the genus Neotama (clade 7, Fig. 1a) and Ypypuera whereas Hersilia. The core of this clade is formed by nine a biarticulate metatarsus unites Murricia and Hersilia species from Central Africa (clade 25). Clade 25 is (clade 12, Fig. 1a). The uniarticulate metatarsus of well supported by the following synapomorphies: the genus Yabisi is a reversal. bulbus with a basal swelling (charater 33[1]), the Character 5 Metatarsal trichobothria (0) one to embolus originate retrolaterally on the bulbus (char- two; (1) four Griswold (2004: 56, character 5) noted acter 41[3]) and the presence of a basal embolar that the presence of more than two metatarsal process (character 45[1]). Clade 17 has several species trichobothria is derived for entelegyne spiders. The that branches off on consecutive, weakly supported presence of four trichobothria distad on the metatarsi branches, basal in position. These include several of hersiliids represents a synapomorphy for the fam- Oriental species such as H. flagilifera, H. kinabaluen- ily (clade 1, Fig. 1a). sis and H. sundaica. Notably, three Afrotropical species Character 6 Metatarsus/tarsus (0) less than four; are also included in this list: H. madagascariensis and, (1) more than eight. An elongate metatarsus is in clade 18, H. incompta and H. scrupulosa. a synapomorphy for all arboreal hersiliids and Tama The distribution of the genus Murricia and clade 17 edwardsi (clade 4, Fig. 1a). throughout the Oriental and Afrotropical Regions Character 7 Claw tuft hairs (0) absent; (1) pres- point to the colonisation, presumably repeated, of ent (Fig. 5). Rheims & Brescovit (2004: 195, charac- hersiliid species from the Afrotropical Region to the ter 8) considered the thick claw tufts found in all her- Oriental Region and vice versa after the closure of siliids to be a synapomorphy for the family and is the Thetys Sea (Sierwald 1997). confirmed by this cladogram.

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Character 8 Leg spine microstructure (0) scale- (Fig. 16). The presence of two pairs of tubercles lat- like projections (Fig. 9); (1) longitudinal grooves dor- erally on the abdomen is a putative synapomorphy sally (Fig. 2. 6-8). Electron micrographs of large for Hersilia scrupulosa and H. incompta (clade 21). spines on the legs of specimens suggest that scale- A smooth abdomen is considered plesiomorphic. like projections on the surface of these spines are ple- Character 15 Sigilla (0) large, four or more siomorphic. The longitudinal grooves dorsally on the (Fig. 16); (1) small, inconspicuous, less than four spines of Hersiliola and Tyrotama are derived (clade 2, (Fig. 10). The inconspicuous sigilla is a putative Fig. 1a). synapomorphy for the clade comprising Tyrotama Character 9 Legs spination, ventral surface (0) and Heriliola (clade 2). pair of parallel rows of strong spines along the entire Character 16 V-shaped ventral muscular pits leg; (1) pair of parallel rows of strong spines only (0) absent; (1) present. The cladogram suggests that along femora; (2) strong spines absent. Rheims & the presence of V-shaped ventral muscular pits Brescovit (2004) noted the presence of two rows of on the opisthosoma is derived. This character is strong spines ventrally along the entire length of secondarily lost in Tyrotama and Hersiliola (clade 2). Uroctea legs. In the ground active representatives of Its adaptive significance could be correlated with that Hersiliidae these rows are restricted to the femora of the dorsal sigilla. The abdomens of those taxa and they are absent in arboreal hersiliids and the rep- that have the sigilla are dorso-ventrally flattened. In resentatives of Eresidae. the case of arboreal hersiliids this could aid in their ability remain cryptic on the surface of bark. PROSOMA Character 17 (0) present; (1) absent. The absence of a cribellum is a synapomorphy for Character 10 ALE/AME (0) ALE as large as or the clade comprising Uroctea and Hersiliidae. larger than AME; (1) ALE < 0.7 × AME Rheims & Character 18 – PLS: tS/bS (0) < 0.5; (1) close Brescovit (2004: 194, character 2) identified the to one; (2) > 3. A very elongate terminal segment of enlarged ALE, that is usually small and barely con- the posterior lateral spinneret, at least three times spicuous in other hersiliids as a reversal in the genus the length of the basal segment, is derived and found Yabisi and it is supported by this phylogeny. in all arboreal hersiliids (clade 5) (Baehr & Baehr Character 11 Interdistance AME-ALE (0) equal 1993a: 84, character 21), whereas an elongate or smaller than 1.5 AME diameter; (1) more than terminal segment is derived for Uroctea and all twice AME diameter (Fig. 19). Rheims & Brescovit hersiliids. (2004: 1945, character 3) observed the projection of the AME anteriorly on the carapace and postulated HABIT it as a synapomorphy for all members of Yabisi. In this tree topology it is also an autapomorphy for Her- Character 19 Habitat (0) ground-dwelling; silia sigillata (Fig. 19). (1) arboreal. Both Baehr & Baehr (1993a) and Character 12 Lateral eyes (0) on slightly conspic- Rheims & Brescovit (2004) suggested that ground uous tubercles (Figs 13, 15, 17); (1) on prominent active behaviour is plesiomorphic in Hersiliidae. tubercles (Figs 19, 21, 83). Baehr & Baehr (1993: 84, The latter authors also noted that Tama edwardsi has character 4) proposed that eyes on prominent tuber- elongated spinnerets and modified metatarsi, sug- cles is a synapomorphy for the genus Murricia. gesting that this species were pre-adapted for an arbo- Rheims & Brescovit (2004: 194, character 1) clado- real life style. gram suggests that it is a synapomorphy for Yabisi. Character 20 Prey capture (0) do not encircle In this analysis it is an autapomorphy for H. sigillata prey; (1) encircle prey, fixing it to substrate with elon- and M. uva, respectively. Eyes on slight tubercles are gate PLS. This is a behavioural adaptation shared by a synapomorphy for the family Hersiliidae (clade 1). Uroctea and hersiliids. Character 13 Cheliceral teeth (0) present (Fig. 4); Character 21 Retreat (0) absent; (1) present. All (1) absent. If present, there are usually three large taxa that are basal in this cladogram construct a cheliceral teeth on the promargin. The loss is derived retreat that is used in the capture of prey and that are for all ground-dwelling hersiliids and eresids in this occupied for certain periods of time. The character cladogram with a reversal for the clade of all arbo- is absent in all arboreal hersiliids (clade 5). real hersiliids (clade 5). Character 22 Retreat shape (0) dome-shaped; (1) pholcid-like; (2) circular. OPISTHOSOMA The construction of a circular retreat is a synapo- morphy for the genus Tyrorama (clade 3) (Jocqué & Character 14 Female opisthosoma with two Dippenaar-Schoeman 2006). Whereas a pholcid-like pairs of tubercles laterally (0) absent; (1) present retreat is derived in clade 2.

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Character 23 Incorporate bits of debris into rounds the median apophysis (Figs 45, 50). The retreats and around egg sacs (0) absent; (1) pres- character is arises as a synapomorphy in clade 19. ent. This behaviour is shared by all hersiliids and Character 32 Sperm duct (0) no basal modifica- Uroctea (Smithers 1945). tions (Rheims & Brescovit 2004:Fig. 35); (1) with basal curvature (Fig. 22); (2) with single basal loop MALE GENITALIA (Fig. 23); (3) with two basal loops (Rheims & Brescovit 2004:Fig. 38); (4) starting from the base Character 24 Palpal patella with spinose ridge of the embolus, medially with one inward bend fol- (0) absent; (1) present. lowed by a large outward bend, that in turn is fol- The possession of a spinose ridge on the palpal patella lowed by a medial inward bend and a basal loop (Baehr & Baehr 1993:26, Fig. 20d) is derived in (Fig. 25); (5) starting from the base of the embolus, clade 24 that includes H. asiatica and H. savignyi. basally with a hairpin outward bend followed by a Character 25 Tibia/patella male palp (0) short, bend inwards medially (Fig. 26); (6) Starting from close to one; (1) elongate, longer than one; (2) very the base of the embolus, the sperm duct has a vari- elongate, twice as long as patella. An elongate palpal able number of inward and outward bends across the tibia evolves once in clade 4 and reverses back to bulbus (Fig. 24). In the ventral view of the left palp, a shorter tibia several times in this tree topology. the duct has to be followed in a clockwise direction, A very elongate tibia is a synapomorphy for the genus starting from the embolus base. Not all species allow Neotama (clade 7). the entire course of the duct to be followed without Character 26 Male palpal tibia with angular clearing. A sperm duct with a basal curvature is projection (0) absent, palpal tibia cylindrical derived in clade 1, whereas a single basal loop is (Figs 37-38, 41-42); (1) present, palpal tibia with a synapomorphy for clades 2 and 6, respectively. angular dorsal projection (Figs 45-46, 61-62). An Two basal loops is an autapomorphy for the genus angular projection on the palp is independently Yabisi, a sperm duct that medially has one inward derived in clades 20 and 26 bend followed by a large outward bend, that in turn Character 27 Palpal tibia, dorsal spines is followed by a medial inward bend and a basal loop (0) absent (Fig. 65); (1) present (Figs 46, 54). is a synapomorphy for the genus Murricia. A hairpin Most Hersiliidae have no dorsal spines on the pal- outward bend followed by a bend inwards medially pal tibia and this character is only found in represen- is an autapomorphy for Prima ansiae. The presence of tatives of Hersilia. These spines arise independently an unmodified sperm duct is a reversal in the genus in clade 15, clade 20 and H madagascariensis Ypypuera. (clade 27). Character 33 Bulbus with a basal swelling Character 28 Cymbium and alveolus shape 0 = absent; 1 = present (Figs 58, 62). (0) short cymbium with circular alveolus (Rheims & The presence of a basally swollen bulbus unites all Brescovit 2004:Fig. 32); (1) elongate cymbium with taxa in clade 25. oval alveolus (Rheims & Brescovit 2004:Fig. 34). An Character 34 Median apophysis (0) present elongate cybium with oval alveolus is an autapomor- (Figs 30, 37, 41, 45, 53, 65, 73, 86, 88); (1) absent phy for the genus Tamopsis. (Figs 33, 49-50, 58, 62, 69). Rheims & Brescovit Character 29 Cymbium distally (0) narrows grad- (2004) suggested that the presence of a median ually (Rheims & Brescovit 2004:Fig. 33); (1) narrows apophysis is plesiomorphic based on Griswold et al.’s abruptly (Rheims & Brescovit 2004:Fig. 31). (1999) assessment of the secondary repeated loss of Rheims & Brescovit (2004) found that this character the median apophysis in several spider families e.g. unites Neotama cunhabebe and two other Neotropi- Eresidae and Filistatidae. The character is very homo- cal Neotama species as well as Neotama variata from plasious and is secondarily lost several times through- Sri Lanka but is uniformative in this cladogram as out the topology. The loss of a median apophysis only the latter species is included here. unites a large number of species in clade 22. Character 30 Length of cymbium distally Character 35 Median apophysis, insertion (0) short, < 0.1 × cymbium length (Figs 30, 45); (0) medially (Fig. 41); (1) basal (between five and (1) elongate, > 0.3 × cymbium length (Figs 29, 69, seven o’clock) (Rheims & Brescovit 2004:Fig. 103); 80). An elongate distal part of the cymbium is inde- (2) distally (between eleven and one o’clock) (Figs 53, pendently derived in Hersiliola and at clade 5, it 80). The phylogeny suggests that a medially inserted reverses in H. sigillata and clade 21 (H. incompta and median apophysis is plesiomorphic. A distally H. scrupulosa). inserted median apophysis unites Yabisi habanensis Character 31 Tegulum with ridged modifica- and Prima ansiae. It is synapomorphy for genus Murri- tions (0) absent (Fig. 37); (1) present (Figs 45, 50). cia, and a autapomorphy for H. sericea and H mada- The bulbus forms a ridged modification that sur- gascariensis. A basally inserted median apophysis

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unites H. savignyi and H. asiatica, and is an autapo- (3) straight, do not circle bulbus circumference. morphy of Ivaraiva pachyura. The retrolateral inser- A long embolus has independently evolved three tion is an autapomorphy of Ypypuera crucifera. times viz. Murricia, Tamopsis and Ypypuera. A very Character 36 Median apophysis, shape (0) elongate embolus is also homoplasious being inde- t-shaped; (1) bifid (Fig. 73); (2) concave, with lamel- pendently derived in Dresserus colsoni, the genus Her- lae distally (Fig. 42); (3) hook-shaped, convex siliola and Hersilia flagilifera. A straight embolus that (Figs 29, 37, 41); (4) lamellae basally (Baehr & does not encircle the bulbus is a synapomorphy for Baehr 1993, Figs 22c-d); (5) brush-like apophysis clade 25. (Baehr & Baehr 1993, Figs 20c-d); (6) large, Character 42 Embolus form (0) filiform, cylin- extremely intricate (Baehr & Baehr 1993, Figs 38c- drical (Fig. 30); (1) medial projections (Fig. 87); d); (7) flap-like (Figs 53-54). A T-shaped median (2) wide, hollowed, tip wide (Figs 45-46); (3) sinu- apophysis is considered to be plesiomorphic in this ate or forked (Figs 54, 57, 62); (4) corkscrew-shaped tree topology. A bifid median apophysis is an H. kinabaluensis (Baehr & Baehr 1993:Fig. 19c). The autapomorphy for Prima ansiae. The presence of character is ambiguous in clade 14. A wide, hollowed a hook-shaped median apophysis is derived in embolus independently evolved in clades 10 and 19. the clade that unites all hersiliids with Uroctea. An embolus with medial projections unites H. cau- A concave and hollowed median apophysis is a data, H. albicomis and H. occidentalis. The presence synapomorphy for clade 17. of a sinuate or forked embolus evolved in clade 22 Character 37 Hook-shaped median apophysis and a corkscrew embolus is an autapmorphy for form (0) smooth, without appendages, distally H. kinabaluensis. pointed (Fig. 37); (1) smooth, distally pointed with Character 43 Basal embolar process (0) absent basal projection (Figs 41-42); (2) smooth, without (Fig. 37); (1) present (Figs 57-58, 61-62). This is a appendages, tip blunt (Fig. 65). The presence of a synapomorhy for clade 23. distally pointed median apophysis with a basal pro- Character 44 Sclerotised tegular projection jection is an autapomorphy of Hersilia aldabrensis, (0) absent; (1) present (Figs 61-62). This straight pro- a characteristic shared with Neotropical representa- jection arises medially on the bulbus and it defines tives of Neotama that were not included in this analy- the clade containing H. mowomogbe, H. alluaudi and sis. A distally truncate median apophysis is an autapo- H. sagitta. morphy of Hersilia sigillata. Character 38 Embolus in relation to median FEMALE GENITALIA apophysis (0) disposed around median apophysis (Fig. 29); (1) adjacent to median apophysis (Fig. 45). Character 45 Epigynal plate (0) entire (Figs 55, The phylogeny suggests that an embolus that is dis- 59, 63); (1) divided, median plate with pair of lateral posed around the median apophysis is plesiomorphic, borders (Figs 31, 39). Rheims & Brescovit (2004) whereas an adjacent embolus is a synapomorphy for tree topology suggested that an entire epigynal plate clades 8 and 19, respectively. is plesiomorphic whereas a divided epigynal plate is Character 39 Slide-like lateral projection (0) derived and confirmed by this analysis. absent; (1) present. The presence of a slide-like lateral Character 46 Oval lateral openings between projection is homoplasious and is derived in the genus median plate and lateral borders (0) absent Hersiliola, Neotama and clade 15, respectively. (1) present (Fig 72). This is a synapomorphy for the Character 40 Embolus, position from which genus Neotama (clade 7). embolus originates on bulbus (0) apical (between Character 47 Median plate (0) partially covered eleven and one o’clock) (Fig. 33); (1) prolateral by lateral borders (Figs 31, 39, 43); (1) free (Figs 47, (between eight and ten o’clock) (Fig. 45); (2) basal 67, 71). This character was also proposed by Rheims (between five and seven o’clock) (Fig. 29); (3) retro- & Brescovit (2004). A free median plate is an lateral (between two o’clock and four o’clock) autapomorphy for Hersiliola, Neotama corticola and (Fig. 57). The tree topology suggests that an apically Hersilia sigillata, respectively, and a synapomorphy inserted embolus is plesiomorphic. A prolaterally for clade 15. inserted embolus is derived for Uroctea durandti and Character 48 Median plate laterally (0) smooth in clade 19. A retrolaterally inserted embolus is a (Fig. 31); (1) with lateral sclerotisations or fixing synapomorphy for clade 25 and a basally originating structures (Figs 39, 43); (2) rippled (Fig. 48). Fixing embolus is derived in clades 6 and 23 with reversals structures laterally on the median plate is a synapo- to an apically origin in the clade 9. morphy for clade 15. A rippled lateral border of the Character 41 Segment of bulbus circumference median plate is a syanpomorphy for clade 19. circled by embolus (0) at most 270° (Fig. 37); (1) Character 49 Lateral borders with triangular 270°-360° (Figs 80-81); (2) more than 360° (Fig. 29); pockets (0) absent; (1) present (Figs 39, 44). The

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presence of triangular pockets on the lateral borders large spermathecae (Figs 32, 40). Forster (1980) is derived in clade 15. suggested a possible evolutionary sequence starting Character 50 Epigynal atrium (0) absent; with Grandungula sorenseni Forster (Gradungulidae), (1) present (Fig. 35). An epiginal atrium is a synapo- where the invagination of bursal wall with several morphy for the genus Tyrotama (clade 3). small spermathecae forms a stable bilobed arrange- Character 51 Copulatory openings (0) incon- ment of the spermathecae and a seminal receptacle. spicuous, (1) visible as circular or oval openings that The possible loss of these secondary spermathecae are either sclerotised or not (Fig. 55). This character would then give rise to the bilobed structure found in is a synapomorphy for clade 22. Hersilia sigillata group. The presence of multiple – Character 52 Epigynum with oval median plate more than 30 – small spermathecae are most appar- (0) absent; (1) present (Figs 42e-f). Baehr & Baehr ent in Murricia uva (Fig. 49). (1993) proposed an oval central field on the epigy- Character 61 Spermathecae shape (0) globose num as a synapomoprhy for the genus Murricia. (Fig. 40); (1) cylindrical (Fig. 44); (2) kidney-shaped It is confirmed by this cladogram. (Fig. 68). Kidney-shaped spermathecae is an autapo- Character 53 Copulatory openings relative to morphy of the genus Yabisi. A cylindrical spermath- spermathecae (0) posterior (Fig. 32); (1) median ecae is derived in clades 6, 16, 21 and 25 and reverses (Fig. 79); (2) anterior (Fig. 36). A copulatory open- in clade 10. ing that opens anteriorly on the epigynal plate is Character 62 Spermathecae stalk (0) absent a synapomorphy for Tyrotama species (clade 3) and (Figs 41, 65); (1) present (Fig. 64). Rheims & the clade comprising the genera of Iviraiva and Yabisi Brescovit (2004) proposed stalked spermathecae as (clade 11), whereas a copulatory opening that opens synapomorphies for Hersilia and Hersiliola, respec- mesally is derived in clade 9. tively. This tree topology supports this proposal for Character 54 Copulatory ducts (0) narrow Hersiliola but not for all Hersilia. A stalked spermath- (Fig. 32); (1) wide, exceeding diameter of fertilisa- ecae is derived in clade 12 and reverses in clades 21 tion ducts (Fig. 72). Wide copulatory ducts were and 25. independently derived in Neotama (clade 7) and Character 63 Spermathecae and seminal recep- Ypypuera. tacles clearly separated from ducts (0) absent Character 55 Length of copulatory duct (0) very (Fig. 79); (1) present (Fig. 40). A spermathecae that long, at least three times spermathecae length is clearly separated from both the copulatory and (Fig. 32); (1) median, twice as long as spermathecae fertilisation ducts is a synapomorphy for the genus length (Fig. 48); (2) short, equal (Fig. 40). The length Hersilia (clade 14). of the copulatory duct is very homoplasious. A short Character 64 Number of seminal receptacles copulatory duct is derived in clade 4 but reverts back (0) absent; (1) several; (2) two; (3) one (Fig. 40); to an elongate duct several times. Ducts of median (4) one, large hyaline (Fig. 36) (Rheims & Brescovit length independently evolved in Murricia, clade 10, 2004). The definition of a seminal receptacle is taken clade 20 and clade 25. from (Sierwald 1999: in Rheims & Brescovit 2004) Character 56 Copulatory duct (0) simple, straight as opening directly into the spermathecae and its (Fig. 40); (1) circular, looped (Figs 85-86); (2) regu- stalk with no connection to the fertilisation duct or larly spirally coiled (Fig. 32); corkscrew-shaped copulatory duct. The presence of a seminal recepta- (Baehr & Baehr 1993:Fig. 19f). A looped copulatory cle unites all hersiliids (clade 1). A large hyaline sem- duct is a synapomorphy for the genus Murricia inal receptacle is a synapomorphy for Tyrotama (clade 13), whereas a corkscrew-shaped duct is an species (clade 3). Some representatives of the genus autapomorphy for Hersilia kinabaluensis. Neotama have two, whereas it is secondarily lost in Character 57 Spermathecae (0) visible; (1) incon- Yabisi and the presence of several seminal recepta- spicuous. Inconspicuous spermathecae is an autapo- cles is an autapomorphy for Iviraiva. morphy for the genus Iviraiva Character 65 Seminal receptacle shape (0) glo- Character 58 Spermathecae basally with bul- bose (Figs 32, 36, 40); (1) irregular with apohysis bous sac (0) absent; (1) present (Fig. 56). A bulbous (Fig. 64); (2) kidney-shaped (Fig. 44) (3) tube-like, sac basally independently evolved in H. savignyi and complexly coiled (Fig. 86); (4) cylindrical. A cylin- H. madagascariensis. drical seminal receptacle is derived in clade 4. A tube- Character 59 Spermathecae with glands basally like, elongate and complexly coiled seminal recepta- (0) absent; (1) present (Fig. 44). cle is a synapomorphy for the genus Murricia The presence of these glands is a synapomorphy for (clade 13). A kidney-shaped seminal receptacle is a the clade 16. synapomorphy for clade 16 and an irregularly shaped Character 60 Number of spermathecae (0) sev- seminal receptacle with apophyses is a synapomorphy eral spermathecae, more than 30 (Fig. 86); (1) one for clade 25.

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Character 66 Seminal receptacle borders Description Size: small to medium (3-13 mm). (0) smooth (Figs 32, 36, 48, 72, 79, 86); (1) with dis- Colour: varies from golden brown to white to almost tal tubercle (Figs 39, 44, 55, 64, 66, 68). black with mottled appearance. Baehr & Baehr (1993a) referred to this character as Carapace: ovoid and flattened, as wide as long; glandular parts covered by threads. This character is narrow longitudinal fovea with three pairs of radi- synapomorphy for the genus Hersilia (clade 14). It ating striae; densely covered with plumose setae. reverses in H. vanmoli and H. furcata. Cephalic region moderately elevated, thoracic Character 67 Fertilisation duct (0) short, shorter region inclined or raised, higher than thoracic than spermathecae length (Fig. 40); (1) long, longer region and thoracic region flattened. Eyes: eight in than spermathecae length (Figs 60, 64); two very two strongly recurved rows (Fig. 10); anterior long, twice as long as spermathecae length (Fig. 48). median eyes often largest; lateral eyes on slight or A long fertilisation duct is a synapomorphy for prominent tubercle (Figs 17, 19). Clypeus length clade 8 and reverses in clade 11 (Iviraiva and Yabisi). variable, short (Fig. 15) or very elongate (Fig. 13) It is also derived in clade 25. Very long fertilisation Chelicerae: unarmed (Tyrotama Tama and Hersi- ducts is derived in clade 18. liola); armed with three large teeth on promargin and 5-11 minute teeth on retromargin (Hersilia, Murricia, Neotama and Prima); labium free with rounded tip; endites oblique often touching each other. Sternum: heart-shaped, anterior edge straight Family Hersiliidae Thorell 1870 (Figs 4, 5) or slightly convex. Abdomen: dorsoventrally flattened, with 4-5 distinct Herséliensis Simon, 1864: 343 (Lycosiformes, in dorsal muscular pits (Hersilia, Neotama, Prima and part). Tama) (Fig. 10) or convex with four or less indistinct Hersilioidae Thorell, 1870: 109, 114; Ausserer, dorsal muscular pits (Tyrotama, Hersiliola) (Fig. 12); 877: 110. abdomen oval to triangular in shape; densely covered Chalinuroidea Thorell, 1873: 605; Kaston, 1938: with plumose setae (Fig. 2); respiratory system: two 640 (Chalinuroidae = Hersiliidae). booklungs; tracheal spiracle opening close to spin- Hersilidae Simon, 1874: 14, 15 (lapsus). nerets; anal tubercle small, reduced. Spinnerets: Hersiliidae Simon, 1882: 255; Lucas, 1869: 1; Simon, six spinnerets; posterior lateral spinnerets longer than 1893: 440; Smithers, 1945: 1; Benoit, 1967: 1; Baehr prosoma width (Hersilia, Murricia, Neotama and & Baehr, 1987: 351; 1993a: 3; Dippenaar-Schoeman Prima), shorter than prosoma width (Tyrotama Tama & Jocqué, 1997: 181; Levi, 2003: 1; Rheims & and Hersiliola); cylindrical with elongated and taper- Brescovit, 2004: 202; Wunderlich 2004: 814. ing terminal segment; inner surface with series of long spinules (see spinneret terminology in Codding- Type genus: Hersilia Audouin, 1827. ton 1999) producing the thin silk threads (Fig. 3). Legs: three claws with modified claw tufts surround- Diagnosis Posterior lateral spinnerets long, terminal ing claws (Fig. 5); unpaired claw simple with two segment tapering, longer than basal segment; longest teeth; paired claws with 5-12 minute teeth, legs very leg at least 1.4 times total body length. Hersiliidae long; leg III shortest; all legs except leg III with resembles by the presence of elongate metatarsi uniarticulate (Tyrotama, Hersiliola and posterior lateral spinnerets and the way in which prey Yabisi), elongate flexible zone distad (Prima, Tama, is captured: the spider rapidly encircles the prey and Tamposis and Iviraiva), narrow flexible zone (Neo- enswathes it by holding its spinnerets over the prey tama), biarticulate (Hersilia and Murricia); 3-5 tri- and fixing it to the substrate. They are distinguished chobothria distad on metatarsi; legs with few spines by the reduced anal tubercle; plumose setae that cover of which scale-like projections covering spine surface prosoma and abdomen; modified claw tufts sur- (Figs 6, 8, 9) (Hersilia, Murricia, Neotama, Prima), or rounding tarsal claws (Fig. 5); 3-5 trichobothria dis- spines dense, longitudinal grooves dorsally (Fig. 7) tally on metatarsi. (Tyrotama, Tama and Hersiliola); autospasy occurs at Phylogenetics Six unambiguous synapomorphies patella-tibia joint. are hypothesised for the family Hersiliidae: The pres- Epigyne: Epigynal plate entire (Fig. 60) or longitu- ence of plumose setae (character 1[1]), four dinally divided into three parts, a median plate and metatarsal trichobothria (character 5[1]), the pres- two lateral borders (Fig. 28); median plate free ence of claw tufts (character 7[1]), ALE < 0.7× AME (Fig. 44) or slightly covered by lateral borders (character 10[1]), posterior lateral eyes on slight or (Figs 36, 40); median plate smooth (Fig. 28) or with prominent tubercles (character 12[1&2]); presence sub-triangular or sickle-shaped fixing structures of one seminal receptacles (character 63[3]). laterally (Figs 36, 40), atrium surrounded by a

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sclerotised rim in Tyrotama (Fig. 32) or ripples Hersiliola from the Palearctic and Afrotropical (Fig. 45 dorsal view of epigynum). Copulation open- Regions; Murricia from the Oriental and Afrotrop- ings posterior (Fig. 29), medial (Fig. 80) or anterior ical Regions; Prima from the Afrotropical Region; (Fig. 33); copulatory ducts simple short (Fig. 37), Promurricia from the Oriental Region; Neotama simple, elongate, as long as epigynal plate (Fig. 45) from the Oriental, Neotropical and Afrotropical or coiled (Fig. 29). Spermathecae unique (Fig. 37), Regions; Tama from the Palearctic Region; Tyrotama several and small (Fig. 41) several and large (Fig. 49); from the Afrotropical Region; Ypypuera and Iviraiva spermathecae shape varies from globose (Fig. 37), from the Neotropical Region; Yabisi from the cylindrical (Fig. 41) to subtriangular (Fig. 69). Nearctic and Neotropical Regions. Seminal receptacle large, hyaline (Fig. 33), small sclerotised (Fig. 37); fertilisation ducts short, curved Key to the genera of Hersiliidae from the (Fig. 37), elongate, straight (Fig. 45), elongate with Afrotropical Region medial curve (Fig. 65). 1. Chelicerae unarmed; posterior lateral spinnerets Palps: tibiae short as long as patellae, apophysis and < carapace width; leg IV longest; thoracic region modified structures absent; cylindrical (Fig. 34) or of carapace sloping (Figs 10-11) ...... 2 with dorsal projection (Fig. 43), spines absent — Chelicerae armed; posterior lateral spinnerets (Fig. 35) or 2-7 strong spines present on promargin > carapace width; leg I or II longest; thoracic (Fig. 43); ventrally with several setae that extends in region of carapace dorso-ventrally flattened front of bulbus (Fig. 34). Cymbium covered with (Figs 12-21) ...... 3 short setae; 2-7 strong spines apically; cymbium 2. Seminal receptacle, small and sclerotised; bulbus compact, not projecting much beyond bulbus flattened with regularly curved sperm duct; embo- (Fig. 30); apex narrowing abruptly beyond bulbus lus coiled, elongate, complete more than one cir- (Fig. 75), digitate, (Fig. 26); bulbus, globose (Fig. 30), cle; median apophysis hook-shaped; tegular projec- flattened (Fig. 26), round (Fig. 35) or basally swollen tion, absent (Figs 29-30) ...... Hersiliola (Thorell) (Figs 59, 63); subtegulum semicircular, sclerotised — Seminal receptacle large, unsclerotised; bulbus and entire; tegulum with tegular projection (Fig. 63); globose, sperm duct with one or more switch conductor, if present, accompanies the embolus backs; embolus originate distally on bulbus, short (Figs 58, 62) or slide like tegular projection of does not complete a full circle; median apohysis varying sizes that accompany embolus often form- absent; tegular projection, if present, short stout ing a fulcrum in which the embolus lies (Figs 81-83); distad on bulbus (Figs 33-34) ...... Tyrotama embolus stout, hook-shaped (Fig. 50), filiform 3. Metatarsi distally with flexible zone on legs I, II (Fig. 38); ridged (Fig. 42); embolic projections usu- and IV (Fig. 28) ...... 4 ally absent, when present subapical (Fig. 81); embo- — Metatarsi biarticulate in legs I, II and IV lus arise apical on bulbus (Fig. 82), basal (Fig. 26), (Fig. 27) ...... 5 prolateral (Fig. 42), or retrolateral (Fig. 58); sperm 4. Distal third of metatarsus I, II, IV flexible (Fig. 28) duct with basal curvature (Fig. 22); basal loop ...... Prima gen. n. (Fig. 23), repeated curvature (Fig. 24); basal loop — Narrow flexible zone in distal third of meta- with medial curvature (Fig. 25). Median apophysis, tarsi I, II and III ...... if present, hook-shaped, convex, apex acute (Fig. 34) ...... Neotama Baehr & Baehr or hook-shaped, hollowed medially, apex acute or 5. Lateral eyes on slight tubercles, abdomen longer truncate with or without basal projection (Figs 38, than wide (Figs 12, 14, 16, 18), copulatory duct 83) or complex (Fig. 42) or bifid (Fig. 74); median straight simple, seminal receptacles simple apophysis insertion, medial (Figs 34, 38) or distal spherical to subquadrate with distal tubercle (Fig. 74). (Figs 40, 44) ...... Hersilia Audouin Composition Hersilia Audouin, 1826 – 73 species; — Lateral eyes on prominent tubercles, abdomen Hersiliola Thorell, 1870 – seven species; Murricia wider than long, subtriangular to subquadrate Simon, 1882 – five species; Tama Simon, 1882 – one (Fig 20), copulatory ducts looped, seminal recep- species; Tamopsis Baehr & Baehr, 1987 – 49 species; tacles cylindrical, very elongate and complexly Neotama Baehr & Baehr, 1993 – one species; Ypy- coiled (Figs 85-86) ...... Murricia Simon, 1882 puera Rheims & Brescovit, 2004 – three species; Ivivraiva Rheims & Brescovit, 2004 – two species; Yabisi Rheims & Brescovit, 2004 – two species; Tyro- Prima gen. n. tama – nine species; Prima – one species. Distribution Tropical and subtropical zones, Type species: Prima ansiae spec. n. worldwide. The genus Hersilia from the Palearctic, Etmology Prima was the daughter of Hersilia, Afrotopical, Oriental and Australian Regions; Roman mythology. The gender is feminine.

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Figures 2-9. Scanning electron micrographs. Hersilia sericea Pocock. 2, Plumose setae; 3, posterior lateral spinnerets, ags = aciniform gland spigots; 4, cheliceral teeth; 5, tarsal claw; 6, microstructure on surface of leg spines. Tyrotama abyssus Foord & Dippenaar- Schoeman. 7, Microstructure on surface of leg spines. Prima ansiae gen. n. 8, Microstructure on surface of leg spines. Uroctea sp. 9, Microstructure on surface of leg spines.

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Figures 10-19. Tyrotama abyssus. 10, 11, Habitus (10), opisthosoma lateral view (11). Hersilia caudata. 12, 13, Habitus (12), opistho- soma lateral view (13). Hersilia mboszi. 14, 15, Habitus (14), opisthosoma lateral view. (15). Hersilia incompta. 16, 17, Habitus (16), opisthosoma lateral view (17). Hersilia sigillata. 18, 19, Habitus (18), opisthosoma lateral view (19). Murricia uva. 20, 21, Habitus (20), opisthosoma lateral view (21).

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Figures 22-28. Hersilia sericea Pocock. 22, Sperm duct. Neotama corticola (Lawrence). 23, Sperm duct. Tyrotama australis (Simon). 24,, Sperm suct. Murricia uva sp. n. 25, Sperm duct. Prima ansieae gen. sp. n. 26, Sperm duct. Hersilia sericea Pocock. 27, Leg I, lateral view. Prima ansieae gen. sp. n. 28, Leg I, lateral view.

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Figures 29-36. Hersiliola macullulata (Dufour). 29, 30, Male palp ventral view (29), prolateral view (30); 31, 32, epigyne ventral view (31), dorsal view (32). Tyrotama arida (Smithers, 1945). 33, 34, Left palp ventral view (33), prolateral view (34); 35, 36, epigyne ventral view (35), dorsal view (36); a = atrium; co = copulatory opening; c = copulatory duct; f = fertilisation duct; lb = lateral borders; mp = median plates = spermatheca; sr = seminal receptacle.

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Figures 37-44. Hersilia albicomis Simon, 1887. 37, 38, Male palp (right palp) ventral view (37), prolateral view (38); 39, 40, epigyne ventral view (39), dorsal view (40). Hersilia aldabrensis Foord & Dippenaar-Schoeman, 2006. 41, 42, Male palp ventral view (41), ven- tral view (42); 43, 44, epigyne ventral view (44), dorsal view (44). e = embolus; ma = median apophysis; fs = fixing structures; s = sper- mathecae; sr = seminal receptacle; f = fertilization duct.

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Figures 45-52. Hersilia scrupulosa Foord & Dippenaar-Schoeman, 2006. 45, 46, Male palp ventral view (45), prolateral view (46); 47, 48, epigyne ventral view (47) dorsal view (48). Hersilia aborea Lawrence, 1928. 49, 50, Male palp ventral view (49) prolateral view (50); 51, 52. epigyne ventral view (50), dorsal view (51); c = copulatory duct; f = fertilization duct; r = ripples.

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Figures 53-56. Hersilia madagascariensis (Wunderlich 2004). 54, 55, Male palp ventral view (54), prolateral view (55); 56, 57, epigyne ventral view and dorsal view.

Diagnoses Species of the genus Prima resemble sal muscular pits; antero-lateral border dark brown; those of Tama, Tamopsis and Iviraiva by the presence venter pale grey to mottled white; femora and tibiae of an elongate flexible zone on metatarsi I, II and IV. pale yellow with faint to dark brown annulation and The males are distinguished by an apically inserted, lateral stripes. bifid median apophysis, short stout embolus (Fig. 74) Carapace: as wide as long; thoracic region widest; and a sperm duct, starting from the base of the embo- cephalic region narrow; dorso-ventrally flattened; lus, basally with a hairpin outward bend followed by fovea longitudinal with radial striae; covered with a bend inwards medially (Fig. 26). The females plumose setae; clypeus truncate in dorsal view, resemble that of Ypypuera by the median copulatory sloping, short, varies in length from 0.46-0.97 × openings on the epigynal plate and that of Promurri- median ocular quadrangle length; eye area cia by the sub-triangular median plate and indistinct depressed to slightly raised (Figs 85, 86); ALE separation of the spermathecae and seminal recep- smallest; AME ≥ PME ≥ PLE >> PLE with ratio tacles. The absence of a biarticulation on metatarsi I, range AME:ALE:PME:PLE = 1: 0.38-0.83: 0.77- II and IV distinguishes it from Promurricia (Fig. 80). 1.25: 0.54-1; chelicerae stout, retromargin with one Phylogenetics Three unambiguous synapomorphies row of minute teeth, promargin with three large are hypothesised for the clade Prima: a sperm duct, teeth; sternum heart-shaped; labium half the length starting from the base of the embolus, basally with a of endite, triangular to crescent-shaped; endites hairpin outward bend followed by a bend inwards elongate, rectangular. medially (character 32[5]), a bifid median apophysis Abdomen: as long as wide, circular (Fig. 77); dorso- (character 36[1]) and an elongate cymbium narrow- ventrally flattened; dorsum with lancet-shaped heart ing abruptly at the distal margin of the alveolus (char- mark; four small dorsal muscular pits that vary in acter 29[1]). size; second pair largest; ventrum with V-shaped mus- Description Female Size: small, range (4.5-6). cular pits. Spinnerets: posterior lateral spinnerets Colour: Carapace pale yellow; clypeus pale yellow long, at least 2 × longer than carapace width; termi- with dark or white markings; eye area dark around nal segment > 4 × length of basal segment; spinules median ocular quadrangle and around posterior eye on posterior lateral spinnerets conical, tapering dis- row, white marks posteriad of eye tubercle in some tally, apex truncate or acute. species; sternum, labium and endites pale grey. Legs: either leg I or II longest; very long, leg I at least Abdomen: dorsum mottled white; heart mark dark 3.23 × longer than total length of body; leg formula brown, lancet-shaped extends up to third pair of dor- I:II:IV:III; leg III very short, < 0.4 × length of leg I;

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Figures 57-64. Hersilia furcata Foord & Dippenaar-Schoeman, 2006. 57, 58, Male palp ventral view (57) prolateral view (58); 59, 60, epigyne ventral view (59), dorsal view (60). Hersilia mowomogbe Foord & Dippenaar-Schoeman, 2006. 61, 62, Male palp ventral view (61) and prolateral view (62); 63, 64, epigyne ventral view (63) dorsal view (64); ebp = basal embolar process, stp = sclerotized tegu- lar projection.

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Figures 65-72. Hersilia sigillata Benoit, 1967. 65, 66, Male palp ventral view (65), prolateral view (66); 67, 68, epigyne ventral view (67), dorsal view (68). Neotama corticola (Lawrence 1973). 69, 70, Male palp ventral view (69), prolateral view (70); 71, 72, epigyne ventral view (71) dorsal view (72).

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metatarsus I > 6 × longer than tarsus I; metatarsi I, Description Male Carapace and clypeus pale II and IV with elongate, flexible zone distally. Spines brown; eye area dark with tridentate white spot pos- short spines < leg diameter; spine formula rather sim- teriad on eye tubercle; abdomen white with antero- ilar between species, variations of the following: I- lateral dark border; cardiac impression linear broad- Fe 1r1p-1d-1d-1r1p1d, Pat, Tib 1p, Mt 1p1r; II- Fe ening posteriorly; posterior half of dorsum with dark 1r1p-1d-1d-1r1p1d-1r-1p-1d, Pat, Tib, Mt 1p1r; III- chevron markings (Fig. 77), distinct transverse lines; Fe 1d-1d-1d, Pat 1d, Tib, Mt; IV- Fe 1d-1d-1d-1r- ventrum white; posterior lateral spinnerets with 1p1d, Pat, Tib 1p, Mt 1r1p. Spine microstruture with annulation; legs pale brown; femora and palps annu- irregular patterns of lancet-shaped scales; tarsal claws late. Total length 5.36; carapace: 2.4 long, 2.4 wide; with seven tarsal teeth. clypeus: 0.39 high, short, 4.88 × OAL; chelicerae Epigyne (Figs 78-79): Epygynum entire, white; cop- short (l/w 0.39), cheliceral groove with eight dentic- ulatory openings medially of spermathecae; copula- ules on the retromargin; eye measurements: diame- tory ducts with medial curve; spermathecae and sem- ters: AME 0.26; ALE 0.07; PME 0.13; PLE 0.2; eye inal receptacles indistinctly separated; fertilisation ratio: 1: 0.25: 0.5: 0.75; interdistances: AME-AME duct medially curved. 0.08; PME-PME 0.08; PME-PLE 0.07; AME-PME Male Size: Small (4.5). Resembles female in shape 0.47; legs: I – Fe 7.2; II – 8; III – 3.12, 2.4, 2.88, 0.8, and colour but differs structurally as follows: smaller total 9.2; IV – 7.6; Palp – 1.33, 1.61, 0.91, total 3.85; in size; abdomen more slender. opisthosoma length: 3.72, width: 2.4; DMP round; Palps (Figs 73, 74): tibia elongate; cymbium elongate, ventral muscular pits V-shaped; PLS, bS: 0.8, tS: 4, narrows abruptly, digitate, one spine apically; alveo- tS/bS 5, 2 × CW. lus oval; bulbus flattened distally; sperm duct with Palps (Figs 73, 74): Tibia short, 0.53 × patella, sim- medial curve; median tegular apophysis apically ple, dorsal tibial spines absent; cymbium digitate, 2.8 inserted; embolus short, stout. × longer than wide, one apical spine; row of long Distribution Madagascar. white setae on prolateral border of cymbium extends across bulbus; bulbus distally truncate, sperm duct circular with basal and medial curvature (Fig. 73); Prima ansieae new species embolus short, slightly curved, apex acute; median Figs 73-79 tegular apophysis apically attached; hollowed, bifid, laterally with concave flap and medially with curved Holotype: Ω: MADAGASCAR, Fianarantsoa process excised at apex; embolus short, stout, apex Province: Parc Nationale, Ranomafana: Talatakely acute. (21°14'S 47°25'E), 5-18.iv.1998, C.E. Griswold, D.H. Female Size (n = 3). Colouration and pattern as Kavanaugh, N.D. Penny, M.J. Raherilalao, J.S. Rano- in males. Total length 5.62 (5.25-6); carapace: rianarisoa, J. Scweikert, D. Ubick, (CAS). 2.06 (1.84-2.16) long, 2.14 (2-2.16) wide; clypeus: Paratypes: 1 æ, 8 juveniles: same data; 1 æ: Parc 0.28 (0.26-0.3) high, clypeus short, 2.39 × OAL; Nationale Ranomafana: 23 km N. Vohiparona chelicerae: (l/w 1.67); eye measurements: diame- village, (21°12'S 47°23'E), c.a. 1100 m, 10- ters: AME 0.14; ALE 0.11; PME 0.2; PLE 0.15; eye 11.iv.1998, C.E. Griswold, D.H. Kavanaugh, N.D. ratio: 1: 0.68: 1.04: 0.82; interdistances: AME-AME Penny, M.J. Raherilalao, J.S. Ranorianarisoa, J. 0.21; PME-PME 0.13; PME-PLE 0.14; AME-PME Scweikert, D. Ubick, (CAS); 1 æ: Antsiranana 0.11; legs: I – Fe 4.73, Pat + Tib 5.23, Mt I 6.54, Ta Province; PN Montagne d’Ambre, 2.79 km NE of 0.81, total 16.88; II – 4.86, 6.12, 7.69, 0.85, total park entrance (12°32'S 49°10'E), 21-30.xi.1993, 18.79; III – 1.66, 1.96, 1.73, 0.7, total 5.71; IV – J. Coddington, C. Griswold, N. Scharff, S. Larcher, 4.69, 3.78, 5.4, 0.9, total 14.31; Palp – 1.09, 0.98, R. Adrianmazinama, (CAS); 1 æ: Androrona 0.91, total 2.98; legs long, 3.23 × body length; leg (15º50'S 49º31E), x.1970, A. Lambillon, (MRAC ratio: 1: 1.11: 0.34: 0.96; opisthosoma length: 3.16, 42940). width: 3.16; DMP round; ventral muscular pits Etymology The specific epithet is a patronym in V-shaped; PLS bS: 0.91, tS: 3.93, tS/bS 4.37, 2.26 honour of Dr Ansie Dippenaar-Schoeman for her × CW. contribution to the knowledge of African spiders. Epigynum (Figs 79, 80): wide (el/ew 0.5); epigyne Diagnosis Males are distinguished by the medial with trapezoid central unpigmented field, copulatory curve of the sperm duct; median tegular apophysis, ducts with medial, fertilisation ducts with basal loop apically attached, bifid with concave flap laterally, duct, medially curved distally. medially with a hook-shaped process excised at Distribution Madagascar (Fig. 94). apex (Fig. 73); females with copulatory duct with Natural history Specimens were collected at medial bend, fertilisation ducts with basal loop 1100 m above sea level; females collected in Novem- (Fig. 79). ber and April, males in April.

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Figures 73-79. Prima ansiae gen. n. sp. n. 73, 74, Male palp ventral view (73); prolateral view (74); 75, 76, opisthosoma lateral view (75) anterior view (76); 77, Female habitus; 78, 79, epigyne ventral view (78), dorsal view (79); c = copulatory duct, e = embolus, f = fertilisation duct, ma = median apophysis, s = spermathecae, sr = seminal receptacle.

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Murricia Simon, 1882 with row of four minute teeth, promargin with three large teeth. Sternum: heart-shaped; labium: Murricia Simon, 1882: 255; 1893: 446; Pocock, sub-triangular, two-thirds the length of endites; 1900: 242; Baehr & Baehr 1993: 64. endites stout, subquadrate, broadening anteriorly. Type species: Murricia indica (Lucas, 1836). Abdomen: wider than long, sub-triangular; dorso- Diagnosis Murricia resembles Hersilia by the pres- ventrally flattened; four to five pairs of distinct dor- ence of a biarticulation on metatarsi of legs I, II sal muscular pits that vary in size (Fig. 20); venter and IV. The males are distinguished by a long with V-shaped muscular pits. Spinnerets: posterior embolus, completing one circle, which is positioned lateral spinnerets long > 1.5 × carapace width; ter- apically on the bulbus. The females are distin- minal segment ≥ 4 × length of basal segment. guished by the presence of wide, elliptical poste- Legs: rather short ≤ 2.22 × total body length; either rior-median field, very long and looped copulatory leg I or II longest; leg formula I:II:IV:III; leg III very ducts, and the presence of one tube-like, elongate short, < 0.4 × length of leg I; metatarsus I > 6 × and complexly coiled seminal receptacle. The longer than tarsus I; distal segment of metatarsus opisthosoma is often subtriangular to subquadrate. > 0.7 × length of proximal segment; femur, patella, Phylogenetics Six unambiguous synapomorphies and metatarsus with spines, spine formula tends to are proposed for the genus Murricia: a sperm duct be rather similar between species and were varia- that, starting from the base of the embolus, has an tions of the following: I- Fe 1p1d1r-1p1d1r- inward bend medially followed by a large outward 1p1d1r-1p1r, Pat 1d1d, Tib 1d-1p-1d1r-1p1d1r, bend, which in turn is followed by a medial inward Mt 1p1r-1d; II- Fe 1p1d1r-1p1d-1r-1p1d-1p1r, Pat bend and a basal loop (character 32[4]), an distally 1p1r-1d, Tib 1d-1p1r-1d-1p1d1r, Mt 1p1r-1p-1d; inserted median apophysis (character 35[3]), a long III- Fe 1d-1d-1d-1r, Pat 1d-1d, Tib 1d-1v-1d, Mt embolus that completes at least one circle (charac- 1d-1d; IV- Fe 1p1d-1d1r-1d-1p1r, Pat 1d-1r-1d, ter 42[1]), a copulatory duct that is twice the length Tib 1d1r1p-1d-1p1r1d, Mt 1r1p-1r1p-1d; spine of the spermathecae (character 54[1]), a looped, microstrucutes with randomly arranged conical circular copulatory duct (character 55[1]) and scales covering spine surface. tube-like, complexly coiled seminal receptacles Epigynum: ovate depressions posterior of copula- (character 64[3]). tory openings (Fig. 85); copulatory ducts elongate, Description Female Size: small to medium, range looped; seminal receptacles cylindrical, complexly (4.52-6.38). coiled (Fig. 86); fertilisation duct curved medially. Colour: Carapace varies from pale yellow to dark Remark Forster (1980) suggested a possible evo- brown, with dark rim of varying thickness around lutionary sequence starting with Grandungula border; clypeus pale with dark or white markings; sorenseni (Grandungulidae) where the invagination eye area dark, white mark posteriad on eye tuber- of bursal wall with several small spermathecae form cle; sternum pale with mottled white markings; stable bilobed spermathecae. The numerous sec- labium and endites pale. Abdomen: dorsum with ondary spermathecae found in Afrotropical females mottled white background; heart mark narrow, of Murricia are hypothesised to be homologous to elongate, along entire length of abdomen; antero- these small spermathecae. lateral border dark brown; venter pale to mottled Male Size: small (4.88-5.03). Resemble female in white. Legs: femora and tibiae pale with faint to shape and colour; differs structurally as follows: dark annulation and lateral striae. smaller in size; abdomen more slender, sub- Carapace: varying between longer than wide to as quadrate, widest posteriorly; distal segment of wide as long; thoracic region widest, cephalic metatarsus > 0.6 × length of proximal; metatarsus region narrow; dorso-ventrally flattened; fovea lon- I > 10 × tarsus I, legs relatively longer, 3.45 × total gitudinal with radial striae; clypeus truncate in dor- body length. sal view, not projecting much beyond eye area, Palps (Figs 80-81): tibia stout, dorsal spines absent; varies in length from 0.88-0.94 × median ocular cymbium compact; bulbus large, round, median quadrangle length; eye tubercle slightly raised; eyes: tegular apophysis, if present small, triangular, hook- PER recurved seen from above and in front; AER shaped, distally attached; embolus circular, apex recurved seen from above and in front; ALE small- acute, originate distally on bulbus. est, translucent; PLE > PME <> AME >> PLE with Composition Murricia uva sp. n. (Kenya, Uganda, ratio range AME:ALE:PME:PLE = 1: 0.27-0.42: Rwanda, Cameroon), M. cornuta Baehr & Baehr 0.92-1.3: 1.28-2; median ocular quadrangle wider 1993 (Singapore), M. crinifera Baehr & Baehr 1993 than long; MOQ-AW = MOQ-PW; chelicerae (Sri Lanka), M triangularis Baehr & Baehr 1993 rather stout, 1.61 × longer than wide, retromargin (India).

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Murricia uva new species 3.68, 4.2, 3.75, 2.1, 0.6, total 14.25; Palp – 0.9, 0.83, Figs 80-86, 95 0.75, total 2.55; legs long, 3.33 × body length; leg ratio: 1: 1.03: 0.32: 0.88; opisthosoma length: Holotype: 1 Ω: KENYA, from Rift Valley Province, 2.78, width: 2.85; five dorsal muscular pits; DMP Marich Pass, Field Studies Centre (1º32'S, 35º27'E), elliptic, second pair by far the largest, third pair the 7.vi.1999, W.J. Pulawski, J.S. Schweikert (CAS). smallest; ventral muscular pits form two converg- Paratypes: CAMEROON: 1 Ω: Chabal Mbabo, SW ing lines; bS: 0.75, tS: 3, tS/bS 4, 1.92 × CW. slope (07º25'N, 12º49'E), 1300 m a.s.l., grassland, Palp (Figs 80-81): Tibia short, as long as patella, 12.iii.1981, Bosmans & van Stalle, (MRAC tibia simple; cymbium with three apical spines; 162559); CONGO D.R.: 1 Ω: Avakubi (01º19'N, embolus circular, filiform, apex acute; median 27º33'E), vii.1932, B. Randoux (MRAC 137384); apophysis distally inserted (Fig. 80), apex acute, pro- UGANDA: 1 æ: Semliki Forest (0º48'N, 30º8'E), laterally directed with rugous surface (Fig. 81). 5.-12.ii.1997, Wagner, deposited in MNHU; Female Size (n = 4). Carapace pale yellow, sparsely RWANDA: 1 æ: PN Akagera, 6 km S de la pêcherie, covered with white setae; clypeus pale yellow; foret seche a Sansevieria (15°5'S, 30°45'E), 14.xi.- median ocular quadrangle dark; opisthosoma later- 3.xii.1985, R. Jocqué, Nsengimana, Michiels ally covered with dense long white setae; mottled (MRAC 165429); 1 æ: PN Akagera, 50 km Nde la white with dark markings postero-laterally; dor- pêcherie Ihema, pres du lac Mihindi (13°2'S, sum with narrow lancet-shaped cardiac impression 30°43'E), 6.xii.1985, R. Jocqué, Nsengimana, (Fig. 20); posterior half of opisthosoma with tri- Michiels, (MRAC 165058). angular darkly mottled area; legs and palps pale yel- low, without annulations. Total length 5.56 (4.52- Etymology The specific epithet is from latin (uva-um 6.38); carapace: 2.13 (2.03-2.25) long, 2.22 = ‘bunch of grapes’) and refers to the large number of (2.03-2.48) wide; clypeus: 0.39 high, short, 3.11 × spermathecae abutting a central stalk, adjective. OAL; chelicerae long: (l/w 1.61), cheliceral groove Diagnosis The males of M. uva sp. n. resemble that with ten retromarginal denticules; eye measure- of H. jajat Rheims & Brescovit (2004:2852, Figs 1- ments: diameters: AME 0.14; ALE 0.04; PME 0.16; 4 (figures of palp was used for this comparison)) in PLE 0.23; eye ratio: 1: 0.36: 1.1: 1.63; interdis- the elongate and apical embolus and truncated cym- tances: AME-AME 0.16; PME-PME 0.13; PME- bium (Fig. 80). They are distinguished by the pres- PLE 0.2; AME-PME 0.13; legs: I – Fe 3.9, Pat + Tib ence of a median apophysis (Fig. 81). Females of 3.86, Mt I 2.81, II 2.06, Ta 0.64, total 13.13; II – M. uva sp. n. resemble that of H. jajat sp. n. Rheims 4.01, 4.14, 2.81, 2.06, 0.6, total 13.35; III – 1.53, & Brescovit (2004b:2852, Figs 5-6) by the long and 1.57, 1.08, 0.55, total 4.77; IV – 3.57, 3.39, 2.89, circular copulation duct and the long tube-like 1.8, 0.56, total 12.19; Palp – 0.89, 1.11, 0.75, total seminal receptacles. They are distinguished by the 2.63; legs short, 2.22 × body length; leg ratio: 1: several, spherical spermathecae abutting a central 1.02: 0.37: 0.93; opisthosoma length: 3.26, width: stalk (Fig. 86). 4.43; five dorsal muscular pits; DMP round; ventral Description Male Size (n = 2). Carapace red- muscular pits V-shaped; bS: 0.78, tS: 3.23, tS/bS brown to pale yellow, dark broad posterior border, 4.1, 1.71 × CW. distinct longitudinal fovea; clypeus with dark medial Epigynum (Figs 85-86): Elongate, (el/ew 0.9); line; eye area dark; opisthosoma: dorsum white with externally with two longitudinal oval depressions; dark markings, dark antero-lateral border; cardiac fertilisation duct short, medially curved. impression lancet-shaped, as long as opisthosoma (Fig. 84); dorsum pale; posterior lateral spinnerets Distribution Cameroon, Kenya, Democratic with strong annulation; legs pale with striations on Republic of the Congo, Rwanda, Uganda (Fig. 95). margins, patellae dark; palps with tarsi dark. Total length 4.95 (4.88-5.03); carapace: 2.25 long, 2.03 Natural history The male paratype was caught at (1.95-2.1) wide; clypeus: 0.32 high, short, 2.47 × an altitude of 1300 m a.s.l., in grassland. Some of OAL; chelicerae long (l/w 1.85), cheliceral groove the type specimens were collected from trees in the with four denticules on the retromargin; eye meas- forests biome. urements: diameters: AME 0.2; ALE 0.08; PME 0.17; PLE 0.18; eye ratio: 1: 0.41: 0.85: 0.9; interdis- tances: AME-AME 0.16; AME-ALE 0.16; PME- ACKNOWLEDGEMENTS PME 0.12; PME-PLE 0.25; AME-PME 0.5; legs: I I wish to thank the curators for loaning material for – Fe 4.28, Pat + Tib 4.8, Mt I 3.98, II 2.4, Ta 0.6, this study; the researchers and technicians of the Biosys- total 16.29; II – 4.35, 5.33, 4.13, 2.48, 0.6, total tematic Division/Arachnida Unit of the ARC Plant Pro- 16.51; III – 1.65, 1.76, 0.9, 0.38, total 4.65; IV – tection Research Institute for their support. Dr Ansie

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Figures 80-86. Murricia uva sp. n. 80, 81, Male palp ventral view (80) prolateral view (81); 82, 83, Opisthosoma lateral view (82), anterior view (83); 84, Male habitus; 85, 86, epigyne ventral view (85), dorsal view (86); c = copulatory duct; f = fertilisation duct; s = spermathecae; sr = seminal receptacle.

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Figures 87-93. Scanning electron micrographs of male palps. 87. H. occidentalis from Butar, Rwanda; 88, H. sericea from Pretoria, South Africa; 89, H. aldabrensis, right palp of male from Aldabra Island Group, Malabar Island; 90, Murricia uva, right palp of male from Chabal Mbabo; 91, Murricia uva, median apophysis; 92, H. madagascariensis (Wunderlich) from Ranomfana, Fianarantsoa Pro- vince, Madagascar; 93, H. sagitta from Shimba Hills Nature Reserve, Kenya.

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2005. This work was supported by the NRF and the Asso- ciation of Commonwealth Universities, Anastasious Christodoulou International Conference Award.

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