Cladistic analysis of the Afrotropical Hersiliidae (Arachnida, Araneae) with the first records of...

INTRODUCTION

Hersiliidae is a small spider family with 141 speciesand 11 genera excluding the results from this study(Platnick 2007, Rheims & Brescovit 2004). The fam-ily is characterised by conspicuously long posteriorlateral spinnerets, elongated legs and is limited to thetropical and subtropical regions of the world. All her-siliids are arboreal except for the representatives ofHersiliola Thorell, 1870, Tama Simon, 1882 and Tyro-tama Foord & Dippenaar-Schoeman, 2005. Thesegenera are all ground active, inhabiting arid regionsof Africa, the Mediterranean and the western partsof the Oriental Region (Smithers 1945, Levy 2003). The family has recently been the focus of extensiverevisions in the Australian (Baehr & Baehr 1987),Oriental (Baehr & Baehr 1993a; Chen 2007),Neotropical and Nearctic Regions (Rheims &Brescovit 2004), and the Afrotropical Region (Foord& Dippenaar-Schoeman 2005a, 2005b; Foord &Dippenaar-Schoeman 2006). Prior to these revisions,Tama was the most widely distributed genus in thefamily. However, Baehr & Baehr’s 1987 revision ofAustralian hersiliids resulted in the transfer of mostspecies previously attributed to Tama to the new

genus Tamopsis Baehr & Baehr, 1987. In this work,29 species, of which 93 % were new to science,were either described or redescribed. Subsequentpublications doubled the number of species knownfor the Australian Region to 55 in total (Baehr &Baehr 1988b, 1989, 1992, 1993b, 1995). In theirrevision of the Oriental Region, Baehr & Baehr(1993a) described 27 new species out of a total of33 taxa and proposed a tentative phylogeny for thefamily. Rheims & Brescovit (2004) described six newspecies, redescribed five species, and proposed threenew genera, Yabisi Rheims & Brecovit, 2004, IviraivaRheims & Brecovit, 2004, and Ypypuera Rheims &Brecovit, 2004, for the Neotropical and NearcticRegions. Once again all the Neotropical and Nearc-tic representatives of Tama were transferred to thelatter three genera and to Neotama Baehr & Baehr,1993.Members of the family Hersiliidae are foundthroughout the Afrotropical Region and include botharboreal (Foord & Dippenaar-Schoeman 2005b,2006) and ground active genera (Foord & Dippenaar-Schoeman 2005a). Descriptive work was donebetween 1827 and 1976. Smithers (1945) revised thethree genera in southern Africa and Benoit (1967)

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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

S.H. FOORD

ABSTRACT

FOORD, S.H. 2008. Cladistic analysis of the family Hersiliidae (Arachnida, Araneae) of the Afrotropical Region, with thedescription 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 theAfrotropical Region is presented, with the description of a new species, Murricia uva sp. n. from Central Africa. Thespecies 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 mostparsimonious trees. The preferred tree supports the monophyly of Tyrotama Foord & Dippenaar-Schoeman, 2005, andretrieves it as the sister-group of Hersiliola Thorell, 1870. These two genera form the most basal lineage of the familyHersiliidae. Tama Simon, 1882, is the sister-group of all the remaining hersiliids, again divided in two large lineages. Thefirst of these clades comprises all arboreal hersiliids without biarticulations on the metatarsi I, II and III. Neotama is basalin 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. Thefamily 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

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revised the genus Hersilia Audouin, 1826, of theAfrotropical Region excluding Madagascar. None ofthese studies attempted to determine phylogeneticrelationships of the hersiliid taxa. Before this studythe Afrotropical Region included records of 43species in four genera, Hersilia (31 species), Hersili-ola Thorell, 1870 (three species), Neotama (onespecies) and Tyrotama (eight species). All Afrotrop-ical species in the genus Tama have been transferredto the new genus Tyrotama (Foord & Dippenaar-Schoeman 2005a). Only Tama edwardsi (Lucas, 1846)remains in Tama and is restricted to the Mediter-ranean and the Palearctic Regions.

Both Tyrotama and Hersiliola occur in the warmerparts of the Afrotropical Region, where they are foundunder stones (Smithers 1945). Hersiliola construct a pholcid-like web and Tyrotama a circular retreat ofclosely woven webbing plastered with small pebbles,chips and vegetable debris. Anchor threads attachedto the substratum warn the spider of approachingprey (Smithers 1945; Dippenaar-Schoeman et al.1999; Jocqué & Dippenaar-Schoeman 2006). Theymove at great speed, overpower their prey and drag itback to their retreat where they feed (Lawrence 1964).Their egg sacs are attached to the underside of rocksand covered with stone chips (Dippenaar-Schoeman& Jocqué 1997). The absence of ground-dwelling representatives from the Oriental, Neotropical andAustralian regions is peculiar. Penney (2006) sug-gested that Yabisi guaba Rheims & Brescovit, 2004,might be a ground wanderer. However, this is refutedby the capture of specimens in the Mangrove Islandsoff the Florida Keys (Simberloff 1971). Hersilia is the most widely distributed genus in thefamily. Platnick (2007) catalogues the Afrotropical,Oriental and Australian Regions, and one species thatpenetrates into the Palaeartic. All Afrotropical speciesof this genus live on tree trunks. They are well cam-ouflaged with their flattened bodies pressed againstthe bark, and when disturbed they move at greatspeed. Although specimens are very cryptic, thereflection of sunlight on the guide threads left ontrunks is a good indicator of their presence (Dippe-naar-Schoeman & Jocqué 1997). Females cover theireggs with an oval-shaped layer of silk camouflagedwith bits of bark. They stand guard over the eggs inan upside-down position above the silk patch untilthe young hatch (pers. obs.).Neotama is represented in the Afrotropical Region byone species, Neotama corticola (Lawrence 1937),which is restricted to the south-eastern region ofSouth Africa (Foord & Dippenaar-Schoeman 2005b).Petrunkevitch (1963), Schawaller (1981) and Wun-derlich (2004) have described several fossil generawhose relationships with extant Hersiliidae remain

uncertain. Recent work on fossil spiders suggests thatthe family is an ancient taxon, and predicts the pres-ence of Hersiliidae in the early Cretaceous (Penney2002). Fossil evidence also points to a much widerdistribution in Europe during the Oligocene andMiocene (Penney 1999; 2006). Lehtinen (1967) con-sidered the Hersiliidae to be related to the Oecobi-idae based on similarity of genital structures, modi-fication of the carapace, eye pattern and spinnerets.Coddington & Levi (1991), in a cladistic analysis ofall spider families, grouped the hersiliids with theoecobiids based on the similarity of the elongatedposterior lateral spinnerets and the distinct prey cap-ture behaviour of the two families. While facing away,the spider rapidly encircles the prey and covers andenswathes it with silk from the long posterior lateralspinnerets. Although Coddington & Levi (1991)placed the hersiliids in the Eresoidea (Hersiliidae,Oecobidae and Eresidae), basal within the Entelege-nae, representative species of Hersiliidae were notincluded in their compilation of data. Baehr & Baehr (1993a) did not consider these rela-tionships well supported and based their phyloge-netic analysis on morphoclines. Baehr & Baehr’s(1993a) attempt at inferring phylogenetic relation-ships for hersiliids, included representatives of thefamily from Australia, the Oriental Region and NewGuinea. They subsequently also proposed a phy-logeny for Hersilia from Australia and New Guinea(Baehr 1998). Their hersiliid phylogeny has Hersiliolabasal, based on the following plesiomorphic charac-ters: short legs, shorter posterior lateral spinnerets, a convex opisthosoma and no flexible zone on legmetatarsi. Tama is defined by longer legs, long pos-terior lateral spinnerets as well as the presence of aflexible zone on the metatarsi of legs I, II and IV,whereas Hersilia has a biarticulate metatarsus, verylong I, II, and IV legs and armed chelicerae. Rheimsand Brescovit (2004) suggested the use of UrocteaDufour, 1820, as an outgroup based on the proposalby Lehtinen (1967) that Hersiliidae is a sister groupof ecribellate Oecobiids belonging to the genusUroctea and followed the principles set forth by Nixon& Carpenter (1993) for outgroup selection. In spiteof the differences in approach, their proposed phy-logeny corresponds with that of Baehr & Baehr(1993a) except for placing Neotama plus Hersilia asthe sister-group to Tamopsis. Baehr & Baehr (1993) suggested that the hersiliidshave an African origin based on the principle thatplesiomorphy and place of origin should correspond.The presence of the plesiomorphic hook-shapedmedian apophysis found on the male palp in someAfrotropical Hersilia species and the occurrence ofthe ground dwelling genera of Hersiliola in Africapoints in that direction. However, Penney (2006) has

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questioned this assertion with reference to fossil evi-dence of a much wider distribution of hersiliidsthroughout the northern hemisphere and invoked thealternative hypothesis of ousted relics (Eskov & Golo-vatch 1986). This would suggest that the current dis-junct ‘Gondwanan’ distribution of hersiliids is theresult of a formerly worldwide distribution followedby extinction of the northern hemisphere represen-tatives (Penney 2006).The taxonomic status of two Oriental genera, Murri-cia Simon, 1882, and Promurricia Baehr & Baehr,1993 has also been doubted (Rheims & Brescovit2004). Murricia Simon is a small genus in the familyHersiliidae, only known from females, with threespecies that have been recorded from India, Sri Lankaand Singapore, respectively (Platnick 2007). Thegenus is characterised by the lateral eyes that are ondistinct eye tubercles, the markedly triangularopisthosoma and the second pair of dorsal muscularpits that are transverse (Baehr & Baehr 1993). Littleis known about their natural history except that theyare arboreal forest dwellers. Baehr & Baehr (1993)discussed their origin and possible relationship withPromurricia Baehr & Baehr from Sri Lanka. The pos-sibility existed that Murricia is a junior synonym ofHersilia with Promurricia intermediate between thefirst two genera (Baehr & Baehr 1993; Rheims &Brescovit 2004). Baehr & Baehr (1993) suggestedthat Murricia originated from ground-living hersili-ids based on the genus’s restricted distribution in the Oriental Region and the synapomorphy that Promurricia shares with Murricia, namely the medio-posterior part of the female epigyne having a wide,oval median plate slightly separated from the laterallobes (Baehr & Baehr 1993).The Afrotropical Region represented the last regionfor which no contemporary revision was available.Such a revision would clarify generic delimitationsin the family Hersiliidae (e.g. Hersilia, MurriciaSimon and Promurricia Baehr & Baehr) as well asthe phylogenetic relationships of the newly describedgenus Tyrotama. This paper therefore attempts toreconstruct Afrotropical hersiliid phylogeny basedon a revision that included 46 Afrotropical hersiliidspecies (Foord & Dippenaar-Schoeman 2005a;Foord & Dippenaar-Schoeman 2005b; Foord &Dippenaar-Schoeman 2006).

METHODS

Study area: The area covered by this study is theAfrotropical Region, including the following islandsin both the Indian and Atlantic Oceans: St. Helena,Comoros, Madagascar, the Seychelles, Zanzibar,Aldabra Islands and Cape Verde but excluding Yemen

(see Rheims et al. 2004). Species from North Africa(countries outside the Afrotropical Region) are onlyincluded when such species also occur in theAfrotropical Region. Slide preparation: The epigyne of the female and theleft palp of the male (when available, otherwise theright) were removed and mounted temporarily onslides in Heinze’s modified PVA mounting medium(Smith-Meyer & Rodrigues 1966). References toillustrations in this publication are capitalisedwhereas references to figures from other papers arenoted in lower case.Illustrations: All illustrations were made with a stere-omicroscope Zeizz Stemi SV 6, using a cameralucida. Scanning electron micrographs were takenwith a JEOL (JSM 840) microscope.Abbreviations: The following abbreviations are usedin this paper: ALE – anterior lateral eye; AME – anterior medianeye; bS – basal segment of posterior lateral spinneret;ebp – basal embolar process; c – copulatory duct; CAI– carapace index; ci – consistency index; CL – cara-pace length; CLL – clypeus length; CW – carapacewidth; el – epigyne length; ew – epigyne width; Fe –femur; MOQ – median ocular quadrangle; MOQ-AW – MOQ anterior width; MOQ-PW – MOQ pos-terior width; MOQL – MOQ length; Mt – metatar-sus; Pat – patella; PER – posterior eye row; PLE –posterior lateral eyes; PME – posterior median eyes;OAL – ocular area length; ri – retention index; Ta –tarsus; Tib – tibia; tS – terminal segment of poste-rior lateral spinneret.Measurements: Measurements were made under astereomicroscope using an ocular micrometer with upto 50× magnification. All measurements are given inmillimetres with the observed ranges in parentheses. Material was received on loan from the followinginstitutions:AMNH – American Museum of Natural History,New York, USA (N.I. Platnick); BMNH – The Nat-ural History Museum, London, UK (J. Beccaloni);CAS – California Academy of Sciences, Golden GatePark, San Francisco, California, USA (C. Griswold);CDO – Collection C. Deeleman, Ossendrecht (C. Deeleman); DNSM – Durban Natural ScienceMuseum, Durban, South Africa (T. Crouch); MCZ –Museum of Comparative Zoology, Cambridge, Mass-achusetts (G. Giribet and L. Leibensperger); MNHU– Museum für Naturkunde der Humboldt Universität,Berlin, Germany (J. Dunlop); MHNG – Muséumd’Histoire naturelle, Genève (P.J. Schwedinger),MNHN – Museum National d’Histoire naturelle,Paris, France (C. Rollard); MRAC – KoninklijkMuseum voor Midden-Afrika, Tervuren, Belgium (R. Jocqué); MSNG – Museo Civico di Storia Naturale Giacomo Doria, Genova (G. Doria); MZBS

PHYLOGENY OF AFROTROPICAL HERSILIIDAE (ARANEAE)

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– Museo de Zoologia, Universidad de Barcelona,Barcelona (C. Ribera); NCA – National Collection ofArachnida, Agricultural Research Council – PlantProtection Research Institute, Pretoria, South Africa(A.S. Dippenaar-Schoeman); NM – Natal Museum,Pietermaritzburg, South Africa (D. Jennings); NMB– National Museum, Bloemfontein, South Africa (L. Lotz); NRM – Swedish Museum of Natural His-tory (T. Kronestedt); QM – Queensland Museum,Brisbane, Australia (B. Baehr); SAM – Iziko Museumof Cape Town, South Africa (M. Cochrane); NMN –National Museum of Namibia, Windhoek, Namibia(T. Bird); SMF – Forschungsinstitut und Naturmu-seum Senckenberg, Federal Republic of Germany(P. Jaeger); TMSA – Northern Flagship Institute (Trans-vaal Museum), Pretoria, South Africa (K. Manamela). A data matrix comprising 40 hersiliid species wasconstructed. This data set included 27 Afrotropicalspecies and 13 hersiliid species from all the othergeographical regions where hersiliids are known tooccur. Three non hersiliid species were included inthe outgroup. The selection of Afrotropical specieswas based on recent revisions (Foord & Dippenaar-Schoeman 2005a; Foord & Dippenaar-Schoeman2005b; Foord & Dippenaar-Schoeman 2006).Species for which only one sex are known wereexcluded from the analysis. This resulted in the inclu-sion of 27 Afrotropical hersiliid species (Appendix 1). Where possible, character variation based on theexamination of type specimens in the following non-Afrotropical taxa were included in the matrix. TheOriental Region: Hersilia albumaculata Wang & Yin,1 Ω from Hong San, S.E. Kiangsi, China, MCZ67908; Hersilia tibialis Baehr & Baehr: 2 æ from Anu-radhapura, Ceylon, SMF 38432 (4 æ, SMF), Hersiliasavignyi Lucas: 5 Ω and 3 æ from Kandy, Ceylon, SMF38429 (5 Ω 3 æ, SMF), Hersilia asiatica Song &Zheng: 2 Ω and 2 æ from Tsin Leong Sam, E. Kwan-tung, China, MCZ 67899 & 67900 & 67911; Hersilia sundaica Baehr & Baehr: 1 Ω, 1 æ from Djojosudharmo, Lombok, 12-14.i.1990, Kozekansika, CDO (1 Ω 1 æ, CDO) Hersilia flagiliferaBaehr & Baehr: 2 Ω and 1 æ paratypes from Ketambe,Gn. Leuser, Nothern Sumatra, 20.ii.1985, Suharto,CDO; Hersilia kinabaluensis Baehr & Baehr: Ω holo-type and æ paratype from Kinabalu National Park,Northern Borneo, ix.1980, C. Deeleman, MHNG;Neotama variata (Pocock): 1 Ω from Dambulla, SriLanka, G. Schmidt, xii.1981, SMF 36782. Palearc-tic:. Tama edwardsi Simon: 1 Ω, 1 æ from Ramble delPouis, Sierra de Carrascoy, Sangonera, Spain,1.iv.1995, rocky wall, J. Miñano, MZBS. Australia:Tamopsis eucalypti Baehr & Baehr, 1 Ω and 1 æ, QM.The following four species were coded with refer-ence to their descriptions Hersilia jajat Rheims & Brecovit – Malaysian, Ivivaiva pachyura (Mello-Leitão)

– Neotropical, Ypypuera crucifera (Vellard) – Neotrop-ical, Yabisi habanensis (Franganillo) – Neotropical.The outgroup consisted of Uroctea durandi (Latreille,1809) from the Mediterranean, Stegodyphus dumi-cola Pocock, 1898 (1 Ω 3 æ, NCA) from South Africaand Dresserus colsoni Tucker, 1920 (2 Ω 2 æ, NCA)based on Lehtinen (1967), Coddington & Levi(1991) and Rheims & Brescovit (2004). All speciesfor which one of the sexes was unknown wereexcluded from the analysis. The preferred tree must be the shortest, most resolvedtree possible, i.e. make the most predictions that canbe tested given only unambiguous branch support(Griswold & Ledford 2001). Preference is, therefore,given to synapomorphic characters in the construc-tion of a phylogeny as they have the largest predictivepower (Farris 1985: in Griswold & Ledford 2001).Analyses of the equally weighted and unordered char-acters were subjected to heuristic searches in Nona2.0 (Goloboff 1997) using h/1000, mult*1000,jump50. Equally weighted characters cannot be con-sidered unweighted and weighting will give moreweight to characters more consistent with initialheuristic cladograms (Farris, 1969: in Bosselaers &Jocqué 2001). Successive weighting with the squaredconsistency index was therefore done in NONA enter-ing the following command sequence: run swt.runh10000 h/1000 mult*1000 jump 50. Pee-Wee 2.6(Goloboff 1997) was used to obtain trees that max-imise implied weights across all characters (Goloboff1993a), with all characters equally weighted (wt = 1)and for both ordered and unordered multi-states(h10000;h/1000;mult*1000;jump50;). Pee-Wee max-imises total fit F=∑fi where fi = k/(k+Esi); k is a con-stant of concavity and can be varied between 1 and6 with the ‘concN’ command, allowing less (1) ormore (6) influence for homoplasious charactes. Con-cavity constants 1, 3 and 6 were used in this analysis.Phylogenetic signal for equally weighted trees wasassessed with branch support indices (Bremer 1994).Branch support was calculated with NONA 2.0(Goloboff 1997) using the options h10000; bsup-port 100. The Bremer Support for a given node inthe shortest unconstrained tree is the number ofadditional steps required in the shortest trees forwhich that node collapses (only for unweightedanalysis). A branch present in one of the most parsimonious trees is more strongly supported bythe data if a large increase in length of additionaltrees is required before that branch is lost in the con-sensus (Griswold & Ledford 2001). Branch supportwas truncated at values of 100 and Bremer supportvalues reported range from 0 and greater. Bremersupport values for successive weighting trees wereobtained in Nona 2.0. Tree editing was done in Win-clada 1.0 (Nixon 2002).

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DISCUSSION

With six genera, the Afrotropical Region now hasthe highest generic level diversity of all biogeographicregions. Heuristic searches of the matrix in NONAwith characters equally weighted resulted in 56 mostparsimonious trees (l = 200; CI = 52; RI = 80). Twelvenodes collapse in the strict consensus of these trees(Fig. 1b). Successive weighting of the unordered multi-state matrix in NONA, generated 7 trees (l = 200;

CI = 52; RI = 80) and five nodes collapsed in the strictconsensus of these trees. The number of trees foundunder implied-weighting in PeeWee was constantwith 8 trees for k = 3 (l = 201; CI = 52, RI = 80)(Fig. 1a), 8 trees for k = 1 (l = 204; ci = 51, ri = 79)and 8 trees for k = 6 (l = 201, CI = 52, RI = 80). Treesconstructed with different character weighting pro-cedures are largely congruent except for relationshipswithin the genus Hersilia and more specifically clade17. Relationships and support for this clade itself is


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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, statechanges 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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low and confirmed by the alternate relationships pro-posed by different weighting procedures. They dif-fered in the relationships and placement of clade 18and the extent to which the relationships of H. asiat-ica and H. saignyi, and H. madagascariensis and H. kin-abaluensis are resolved. The weak support for latterrelationships are also confirmed by low Bremer sup-port values (Figs 1a-b). Three clades are, however,retained in all of the trees viz. clade 15, 18 and 22.The trees based on implied weighting (k = 3) are pre-ferred as these trees save steps for the less homopla-sious characters (Platnick et al. 1991: in Bosselaers

& Jocqué 2000) (Fig. 1). Character states are foundin Appendix 1. All characters discussed are uniquelyderived unless stated otherwise. The proposed phylogeny is largely congruent withthat of Baehr & Baehr (1993a) and Rheims &Brescovit (2004) except that the genus Tyrotama, notincluded in their analyses, is a sister clade to Hersili-ola. A basal dichotomy in the ingroup separatesclade 2, consisting of Hersiliola and Tyrotama, fromall other hersiliids and is characterised by longitudi-nal grooves dorsally on leg setae (character 8 [1]),very small inconspicuous sigilla (character 15 [1]),

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

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and the construction of a retreat (character 22[1,2]).Clade 3 is a very distinct subclade within clade 2,composed of representatives of the genus Tyrotamathat are restricted to the arid regions of southernAfrica and firmly supported by several synapomor-phies: the construction of a circular retreat thatincorporates bits of debris (character 22[1]), asperm duct with a variable number of inward andoutward bends (character 32[6]), an epigynal atrium(character 52[1]) and the presence of a large hya-line seminal receptacle (character 64[4]). This con-firms Baehr & Baehr’s (1993) suggestion that theAfrotropical representatives of Tama sensu lato arenot related to Tama, sensu stricto. As such, the newgenus Tyrotama was described to include Hersiliolaaustralis, H. fragilis, Tama arida, T. bicava, T. incertaand the new species T. abyssus, T. soutpansbergensisand T. taris (Foord & Dippenaar-Schoeman 2005a).The remaining hersiliids are united at clade 4 byleg IV that is less than 0.4× the length of leg I (char-acter 2[1]), the presence of some form of articularmodification on metatarsi I, II and IV (charac-ter 4[1,2,3]), metarsus I more than 8× the lengthof tarsus I (character 6[1]), a tibial palp that islonger than the palpal patella (character 25[1]), themedian plate of the epigyne that is partially coveredby the lateral borders (character 47[0]) and the pres-ence of cylindrical seminal receptacles (charac-ter 65[4]). Tama edwardsi, in the monotypic genusTama, is a ground-dwelling species restricted to theMediterranean and share all these synapomorphieswith arboreal hersiliids, suggesting that it waspreadapted to tree living (Rheims & Brescovit 2004).Clade 5 unites all arboreal hersiliids that are charac-terised by the terminal segment of the posterior lat-eral spinnerets more than 3× the length of the basalsegment (character 18[2]) and the absence of aretreat (character 21[1]). Arboreal hersiliids are divided into two large sisterclades. Both of these clades’ support is rather lowas evidenced by their Bremer support (Figs 1a-b).The first of these, clade 6, groups six genera distrib-uted throughout the tropical and subtropical areas ofthe world and is characterised by several homopla-sious characters, including sperm ducts with a sin-gle basal loop (character 32[2]) that is secondarilylost or modified in Prima, Ypypuera and Yabisi; anembolus that originates basally on the bulbus (char-acter 40[2]) that reverses to an apical origin inclade 9 and cylindrical spermathecae (charac-ter 61[1]) that reverses into spherical spermatheaein clade 10. Neotama (clade 7) is basal in this clade,and the monophyly of this genus is supported by twosynapomorphies: a palpal tibia that is more thantwice as long as the palpal patella (character 25[2]),and the presence of oval slit-like lateral openings

between the lateral borders and median plate of theepigyne (character 46[1]). A narrow flexible zoneon metarsi I, II and IV (character 4[2]), is a featurethat is paralleled in the genus Ypypuera. Neotamahas the widest distribution of all hersiliid genera(Afrotropical, Neotropical and Oriental). The pres-ence of Neotama in the Neotropical Region couldbe explained by the theory of ousted relics (Eskov& Golovatch 1986). Based on this theory Penney(2006) suggested that Hersiliidae could have had apancontinental distribution based on the presenceof fossil representatives that predicts a much widerdistribution of the family than at present.Tamopsis (Australia) is basal in clade 8 with Primagen. nov. (Madagascar), Ypypuera (NeotropicalRegiona), Iviraiva and Yabisi (Neotropical Region)sequentially branching off. The new genus Prima ischaracterised by two synapomorphies: a sperm duct,starting from the base of the embolus, basally with ahairpin outward bend followed by a bend inwardsmedially (character 32[5]) and a bifid median apoph-ysis (character 36[1]). Clade 12 is the sister clade of clade 6 and consists ofall hersiliids that have a biarticulation on metatarsi I,II and IV (character 4[3]). This clade has two sisterclades. The first of these, clade 13, is characterised bya sperm duct that, starting from the base of theembolus, has an inward bend medially followed by alarge outward bend, that in turn is followed by amedial inward bend and a basal loop (charac-ter 32[4]), an oval epigynal plate (character 51[1])and an elongate, circular looped copulatory duct(character 56[1]). There is a single representative ofthis clade in the Afrotropical Region and althoughBaehr & Baehr (1993) did not describe any males forthe genus Murricia this species has several charactersin common with species in this genus from the Ori-ental Region viz. lateral eyes that are on a distinct eyetubercle, large posterior lateral eyes, a triangular totrapezoid abdomen, and a elongate complexly coiledseminal receptacle. Rheims & Brescovit (2004)described Hersilia jajat from Borneo and althoughthe species does not share the latter characteristicswith Murricia, synapomorphies as listed for clade 12and proposed for the genus Murricia in this study ispresent in Hersilia jajat. Clade 14, the sister taxon of clade 13, includes allrepresentatives of Hersilia except Hersilia jajat. Allthe species in this genus have spermathecae and sem-inal receptacles that are clearly separated from thecopulatory and fertilisation ducts (character 63[1])and the presence of a distal tubercle or glandulararea on the seminal receptacle (character 66[1]).Clade 12 that unites Murricia and Hersilia has lessBremer support than clade 13. The genus Murriciais therefore retained. Furthermore, it is obvious from


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the low Bremer support values for clade 17, and theuncertain placement of clade 18 that it would notbe advisable to split the genus. However, clades 15,18 and 25 are well supported. Based on the strict consensus of eight trees foundunder implied weighting, Hersilia consists of twoclades, 15 and 17. Clade 15 comprises eight speciesthat are characterised by the median plate of theepigynum with lateral sclerotised projections thatpresumably act as fixing structures during copula-tion (character 48[1]) and lateral borders with trian-gular projections (character 49[1]). Species in thisclade are widely distributed throughout theAfrotropical Region. Two species, H. sericea and H.setifrons from South Africa, are sympatric and syn-topic in the savannah and have never been collectedin forests. This seems to be true for all the otherspecies in this clade except the enigmatic H. sigillatathat has been collected in forests of Central Africa.The core of clade 15, clade 16, is characterised by thepresence of glands basally on the spermathecae(character 59[1]) and kidney-shaped seminal recep-tacles (character 65[2]). The clade comprises threespecies, two of which are endemic to Madagascar.The third species, H. sigillata, is peculiar, and hasseveral autapomorphies that include a hook-shapedmedian apophysis that is distally truncate (charac-ter 37[2]), a very short, truncate cymbium (charac-ter 30[0]), first three pairs of sigilla, very large andsimilar in size (character 15[2]), the interdistanceof AME-ALE is more than twice that of AME diam-eter (character 11[1]), a characteristic shared withthe genus Yabisi from the Neotropical Region, andlateral eyes that are on prominent tubercles (char-acter 12[2]), which is also shared with Yabisi andMurricia. Clade 17 is the second and largest clade in the genusHersilia. The core of this clade is formed by ninespecies from Central Africa (clade 25). Clade 25 iswell supported by the following synapomorphies:bulbus with a basal swelling (charater 33[1]), theembolus originate retrolaterally on the bulbus (char-acter 41[3]) and the presence of a basal embolarprocess (character 45[1]). Clade 17 has several speciesthat branches off on consecutive, weakly supportedbranches, basal in position. These include severalOriental species such as H. flagilifera, H. kinabaluen-sis and H. sundaica. Notably, three Afrotropical speciesare also included in this list: H. madagascariensis and,in clade 18, H. incompta and H. scrupulosa.The distribution of the genus Murricia and clade 17throughout the Oriental and Afrotropical Regionspoint to the colonisation, presumably repeated, ofhersiliid species from the Afrotropical Region to theOriental Region and vice versa after the closure ofthe Thetys Sea (Sierwald 1997).

CHARACTER DESCRIPTIONS AND INTERPRETATIONS

Character 1 Plumose setae (0) absent; (1) pres-ent (Fig. 2). Griswold et al. (1999: 56, character 18)considered the presence of plumose setae as derived.This character is a synapomorphy for all hersiliids atclade 1 (Fig. 1a).

LEGS

Character 2 Relative length of leg III to leg I(0) = more than 0.6; (1) less than 0.4. The cladogramsuggests that a long leg III is plesiomorphic and relative shortening of leg III is derived. There is, however, uncertainty whether it was owing to a short-ening of leg III or lengthening of leg I, II and IV(Baehr & Baehr 1993a; Rheims & Brescovit 2004).The relative shortening of leg III is a synapomorhy forthe clade of all arboreal hersiliids and Tama edwardsi(clade 4, Fig. 1a).Character 3 Relative length of leg IV, legIV/legI(0) less than one; (1) more than one.The cladogram suggests that an elongate leg IV was derived at the node that unites Uroctea and Hersiliidae and reverted back to a relatively shorterleg IV in clade 4 (Fig. 1a) that includes Tama and allarboreal hersiliids.Character 4 Metatarsi articulation of leg I, II and III (0) uniarticulate; (1) elongate flexible zone(Fig. 28); (2) short flexuose zone; (3) biarticulate(Fig. 27). Baehr (1998) suggested a possible evolu-tionary sequence in which an elongate flexible zoneis reduced in length to form a narrow flexible zoneand ultimately develop into a biarticulation. The elon-gate flexible zone is derived in clade 4 (Fig. 1a). A narrow flexible zone evolved independently in Neotama (clade 7, Fig. 1a) and Ypypuera whereas a biarticulate metatarsus unites Murricia and Hersilia(clade 12, Fig. 1a). The uniarticulate metatarsus ofthe genus Yabisi is a reversal. Character 5 Metatarsal trichobothria (0) one totwo; (1) four Griswold (2004: 56, character 5) notedthat the presence of more than two metatarsal trichobothria is derived for entelegyne spiders. Thepresence of four trichobothria distad on the metatarsiof hersiliids represents a synapomorphy for the fam-ily (clade 1, Fig. 1a).Character 6 Metatarsus/tarsus (0) less than four;(1) more than eight. An elongate metatarsus is a synapomorphy for all arboreal hersiliids and Tamaedwardsi (clade 4, Fig. 1a).Character 7 Claw tuft hairs (0) absent; (1) pres-ent (Fig. 5). Rheims & Brescovit (2004: 195, charac-ter 8) considered the thick claw tufts found in all her-siliids to be a synapomorphy for the family and isconfirmed by this cladogram.

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Character 8 Leg spine microstructure (0) scale-like projections (Fig. 9); (1) longitudinal grooves dor-sally (Fig. 2. 6-8). Electron micrographs of largespines on the legs of specimens suggest that scale-like projections on the surface of these spines are ple-siomorphic. The longitudinal grooves dorsally on thespines of Hersiliola and Tyrotama are derived (clade 2,Fig. 1a).Character 9 Legs spination, ventral surface (0)pair of parallel rows of strong spines along the entireleg; (1) pair of parallel rows of strong spines onlyalong femora; (2) strong spines absent. Rheims &Brescovit (2004) noted the presence of two rows ofstrong spines ventrally along the entire length ofUroctea legs. In the ground active representatives ofHersiliidae these rows are restricted to the femoraand they are absent in arboreal hersiliids and the rep-resentatives of Eresidae.

PROSOMA

Character 10 ALE/AME (0) ALE as large as orlarger than AME; (1) ALE < 0.7 × AME Rheims &Brescovit (2004: 194, character 2) identified theenlarged ALE, that is usually small and barely con-spicuous in other hersiliids as a reversal in the genusYabisi and it is supported by this phylogeny.Character 11 Interdistance AME-ALE (0) equalor smaller than 1.5 AME diameter; (1) more thantwice AME diameter (Fig. 19). Rheims & Brescovit(2004: 1945, character 3) observed the projection ofthe AME anteriorly on the carapace and postulatedit as a synapomorphy for all members of Yabisi. Inthis tree topology it is also an autapomorphy for Her-silia sigillata (Fig. 19).Character 12 Lateral eyes (0) on slightly conspic-uous tubercles (Figs 13, 15, 17); (1) on prominenttubercles (Figs 19, 21, 83). Baehr & Baehr (1993: 84,character 4) proposed that eyes on prominent tuber-cles is a synapomorphy for the genus Murricia.Rheims & Brescovit (2004: 194, character 1) clado-gram suggests that it is a synapomorphy for Yabisi.In this analysis it is an autapomorphy for H. sigillataand M. uva, respectively. Eyes on slight tubercles area synapomorphy for the family Hersiliidae (clade 1).Character 13 Cheliceral teeth (0) present (Fig. 4);(1) absent. If present, there are usually three largecheliceral teeth on the promargin. The loss is derivedfor all ground-dwelling hersiliids and eresids in thiscladogram with a reversal for the clade of all arbo-real hersiliids (clade 5).

OPISTHOSOMA

Character 14 Female opisthosoma with twopairs of tubercles laterally (0) absent; (1) present

(Fig. 16). The presence of two pairs of tubercles lat-erally on the abdomen is a putative synapomorphyfor Hersilia scrupulosa and H. incompta (clade 21). A smooth abdomen is considered plesiomorphic.Character 15 Sigilla (0) large, four or more(Fig. 16); (1) small, inconspicuous, less than four(Fig. 10). The inconspicuous sigilla is a putativesynapomorphy for the clade comprising Tyrotamaand Heriliola (clade 2). Character 16 V-shaped ventral muscular pits(0) absent; (1) present. The cladogram suggests thatthe presence of V-shaped ventral muscular pits on the opisthosoma is derived. This character is secondarily lost in Tyrotama and Hersiliola (clade 2).Its adaptive significance could be correlated with thatof the dorsal sigilla. The abdomens of those taxa that have the sigilla are dorso-ventrally flattened. Inthe case of arboreal hersiliids this could aid in theirability remain cryptic on the surface of bark.Character 17 Cribellum (0) present; (1) absent.The absence of a cribellum is a synapomorphy forthe clade comprising Uroctea and Hersiliidae.Character 18 – PLS: tS/bS (0) < 0.5; (1) close to one; (2) > 3. A very elongate terminal segment ofthe posterior lateral spinneret, at least three timesthe length of the basal segment, is derived and foundin all arboreal hersiliids (clade 5) (Baehr & Baehr1993a: 84, character 21), whereas an elongate terminal segment is derived for Uroctea and all hersiliids.

HABIT

Character 19 Habitat (0) ground-dwelling; (1) arboreal. Both Baehr & Baehr (1993a) andRheims & Brescovit (2004) suggested that groundactive behaviour is plesiomorphic in Hersiliidae. The latter authors also noted that Tama edwardsi haselongated spinnerets and modified metatarsi, sug-gesting that this species were pre-adapted for an arbo-real life style. Character 20 Prey capture (0) do not encircleprey; (1) encircle prey, fixing it to substrate with elon-gate PLS. This is a behavioural adaptation shared byUroctea and hersiliids.Character 21 Retreat (0) absent; (1) present. Alltaxa that are basal in this cladogram construct aretreat that is used in the capture of prey and that areoccupied for certain periods of time. The characteris absent in all arboreal hersiliids (clade 5).Character 22 Retreat shape (0) dome-shaped; (1) pholcid-like; (2) circular.The construction of a circular retreat is a synapo-morphy for the genus Tyrorama (clade 3) (Jocqué &Dippenaar-Schoeman 2006). Whereas a pholcid-likeretreat is derived in clade 2.


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Character 23 Incorporate bits of debris intoretreats and around egg sacs (0) absent; (1) pres-ent. This behaviour is shared by all hersiliids andUroctea (Smithers 1945).

MALE GENITALIA

Character 24 Palpal patella with spinose ridge(0) absent; (1) present.The possession of a spinose ridge on the palpal patella(Baehr & Baehr 1993:26, Fig. 20d) is derived inclade 24 that includes H. asiatica and H. savignyi.Character 25 Tibia/patella male palp (0) short,close to one; (1) elongate, longer than one; (2) veryelongate, twice as long as patella. An elongate palpaltibia evolves once in clade 4 and reverses back to a shorter tibia several times in this tree topology. A very elongate tibia is a synapomorphy for the genusNeotama (clade 7). Character 26 Male palpal tibia with angularprojection (0) absent, palpal tibia cylindrical(Figs 37-38, 41-42); (1) present, palpal tibia withangular dorsal projection (Figs 45-46, 61-62). Anangular projection on the palp is independentlyderived in clades 20 and 26 Character 27 Palpal tibia, dorsal spines(0) absent (Fig. 65); (1) present (Figs 46, 54). Most Hersiliidae have no dorsal spines on the pal-pal tibia and this character is only found in represen-tatives of Hersilia. These spines arise independently in clade 15, clade 20 and H madagascariensis(clade 27).Character 28 Cymbium and alveolus shape (0) short cymbium with circular alveolus (Rheims &Brescovit 2004:Fig. 32); (1) elongate cymbium withoval alveolus (Rheims & Brescovit 2004:Fig. 34). Anelongate cybium with oval alveolus is an autapomor-phy for the genus Tamopsis.Character 29 Cymbium distally (0) narrows grad-ually (Rheims & Brescovit 2004:Fig. 33); (1) narrowsabruptly (Rheims & Brescovit 2004:Fig. 31).Rheims & Brescovit (2004) found that this characterunites Neotama cunhabebe and two other Neotropi-cal Neotama species as well as Neotama variata fromSri Lanka but is uniformative in this cladogram asonly the latter species is included here.Character 30 Length of cymbium distally (0) short, < 0.1 × cymbium length (Figs 30, 45); (1) elongate, > 0.3 × cymbium length (Figs 29, 69,80). An elongate distal part of the cymbium is inde-pendently derived in Hersiliola and at clade 5, itreverses in H. sigillata and clade 21 (H. incompta andH. scrupulosa).Character 31 Tegulum with ridged modifica-tions (0) absent (Fig. 37); (1) present (Figs 45, 50).The bulbus forms a ridged modification that sur-

rounds the median apophysis (Figs 45, 50). Thecharacter is arises as a synapomorphy in clade 19.Character 32 Sperm duct (0) no basal modifica-tions (Rheims & Brescovit 2004:Fig. 35); (1) withbasal curvature (Fig. 22); (2) with single basal loop(Fig. 23); (3) with two basal loops (Rheims &Brescovit 2004:Fig. 38); (4) starting from the baseof the embolus, medially with one inward bend fol-lowed by a large outward bend, that in turn is fol-lowed by a medial inward bend and a basal loop(Fig. 25); (5) starting from the base of the embolus,basally with a hairpin outward bend followed by abend inwards medially (Fig. 26); (6) Starting fromthe base of the embolus, the sperm duct has a vari-able number of inward and outward bends across thebulbus (Fig. 24). In the ventral view of the left palp,the duct has to be followed in a clockwise direction,starting from the embolus base. Not all species allowthe entire course of the duct to be followed withoutclearing. A sperm duct with a basal curvature isderived in clade 1, whereas a single basal loop is a synapomorphy for clades 2 and 6, respectively. Two basal loops is an autapomorphy for the genusYabisi, a sperm duct that medially has one inwardbend followed by a large outward bend, that in turnis followed by a medial inward bend and a basal loopis a synapomorphy for the genus Murricia. A hairpinoutward bend followed by a bend inwards mediallyis an autapomorphy for Prima ansiae. The presence ofan unmodified sperm duct is a reversal in the genusYpypuera.Character 33 Bulbus with a basal swelling0 = absent; 1 = present (Figs 58, 62).The presence of a basally swollen bulbus unites alltaxa in clade 25.Character 34 Median apophysis (0) present(Figs 30, 37, 41, 45, 53, 65, 73, 86, 88); (1) absent(Figs 33, 49-50, 58, 62, 69). Rheims & Brescovit(2004) suggested that the presence of a medianapophysis is plesiomorphic based on Griswold et al.’s(1999) assessment of the secondary repeated loss ofthe median apophysis in several spider families e.g.Eresidae and Filistatidae. The character is very homo-plasious and is secondarily lost several times through-out the topology. The loss of a median apophysisunites a large number of species in clade 22.Character 35 Median apophysis, insertion(0) medially (Fig. 41); (1) basal (between five andseven o’clock) (Rheims & Brescovit 2004:Fig. 103);(2) distally (between eleven and one o’clock) (Figs 53,80). The phylogeny suggests that a medially insertedmedian apophysis is plesiomorphic. A distallyinserted median apophysis unites Yabisi habanensisand Prima ansiae. It is synapomorphy for genus Murri-cia, and a autapomorphy for H. sericea and H mada-gascariensis. A basally inserted median apophysis

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unites H. savignyi and H. asiatica, and is an autapo-morphy of Ivaraiva pachyura. The retrolateral inser-tion is an autapomorphy of Ypypuera crucifera. Character 36 Median apophysis, shape (0) t-shaped; (1) bifid (Fig. 73); (2) concave, with lamel-lae distally (Fig. 42); (3) hook-shaped, convex(Figs 29, 37, 41); (4) lamellae basally (Baehr &Baehr 1993, Figs 22c-d); (5) brush-like apophysis(Baehr & Baehr 1993, Figs 20c-d); (6) large,extremely intricate (Baehr & Baehr 1993, Figs 38c-d); (7) flap-like (Figs 53-54). A T-shaped medianapophysis is considered to be plesiomorphic in thistree topology. A bifid median apophysis is anautapomorphy for Prima ansiae. The presence of a hook-shaped median apophysis is derived in the clade that unites all hersiliids with Uroctea. A concave and hollowed median apophysis is asynapomorphy for clade 17. Character 37 Hook-shaped median apophysisform (0) smooth, without appendages, distallypointed (Fig. 37); (1) smooth, distally pointed withbasal projection (Figs 41-42); (2) smooth, withoutappendages, tip blunt (Fig. 65). The presence of a distally pointed median apophysis with a basal pro-jection is an autapomorphy of Hersilia aldabrensis, a characteristic shared with Neotropical representa-tives of Neotama that were not included in this analy-sis. A distally truncate median apophysis is an autapo-morphy of Hersilia sigillata.Character 38 Embolus in relation to medianapophysis (0) disposed around median apophysis(Fig. 29); (1) adjacent to median apophysis (Fig. 45).The phylogeny suggests that an embolus that is dis-posed around the median apophysis is plesiomorphic,whereas an adjacent embolus is a synapomorphy forclades 8 and 19, respectively.Character 39 Slide-like lateral projection (0)absent; (1) present. The presence of a slide-like lateralprojection is homoplasious and is derived in the genusHersiliola, Neotama and clade 15, respectively.Character 40 Embolus, position from whichembolus originates on bulbus (0) apical (betweeneleven and one o’clock) (Fig. 33); (1) prolateral(between eight and ten o’clock) (Fig. 45); (2) basal(between five and seven o’clock) (Fig. 29); (3) retro-lateral (between two o’clock and four o’clock)(Fig. 57). The tree topology suggests that an apicallyinserted embolus is plesiomorphic. A prolaterallyinserted embolus is derived for Uroctea durandti andin clade 19. A retrolaterally inserted embolus is asynapomorphy for clade 25 and a basally originatingembolus is derived in clades 6 and 23 with reversalsto an apically origin in the clade 9.Character 41 Segment of bulbus circumferencecircled by embolus (0) at most 270° (Fig. 37); (1)270°-360° (Figs 80-81); (2) more than 360° (Fig. 29);

(3) straight, do not circle bulbus circumference. A long embolus has independently evolved threetimes viz. Murricia, Tamopsis and Ypypuera. A veryelongate embolus is also homoplasious being inde-pendently derived in Dresserus colsoni, the genus Her-siliola and Hersilia flagilifera. A straight embolus thatdoes not encircle the bulbus is a synapomorphy forclade 25.Character 42 Embolus form (0) filiform, cylin-drical (Fig. 30); (1) medial projections (Fig. 87); (2) wide, hollowed, tip wide (Figs 45-46); (3) sinu-ate or forked (Figs 54, 57, 62); (4) corkscrew-shapedH. kinabaluensis (Baehr & Baehr 1993:Fig. 19c). Thecharacter is ambiguous in clade 14. A wide, hollowedembolus independently evolved in clades 10 and 19.An embolus with medial projections unites H. cau-data, H. albicomis and H. occidentalis. The presenceof a sinuate or forked embolus evolved in clade 22and a corkscrew embolus is an autapmorphy for H. kinabaluensis.Character 43 Basal embolar process (0) absent(Fig. 37); (1) present (Figs 57-58, 61-62). This is asynapomorhy for clade 23.Character 44 Sclerotised tegular projection(0) absent; (1) present (Figs 61-62). This straight pro-jection arises medially on the bulbus and it definesthe clade containing H. mowomogbe, H. alluaudi andH. sagitta.

FEMALE GENITALIA

Character 45 Epigynal plate (0) entire (Figs 55,59, 63); (1) divided, median plate with pair of lateralborders (Figs 31, 39). Rheims & Brescovit (2004)tree topology suggested that an entire epigynal plateis plesiomorphic whereas a divided epigynal plate isderived and confirmed by this analysis.Character 46 Oval lateral openings betweenmedian plate and lateral borders (0) absent (1) present (Fig 72). This is a synapomorphy for thegenus Neotama (clade 7).Character 47 Median plate (0) partially coveredby lateral borders (Figs 31, 39, 43); (1) free (Figs 47,67, 71). This character was also proposed by Rheims& Brescovit (2004). A free median plate is anautapomorphy for Hersiliola, Neotama corticola andHersilia sigillata, respectively, and a synapomorphyfor clade 15. Character 48 Median plate laterally (0) smooth(Fig. 31); (1) with lateral sclerotisations or fixingstructures (Figs 39, 43); (2) rippled (Fig. 48). Fixingstructures laterally on the median plate is a synapo-morphy for clade 15. A rippled lateral border of themedian plate is a syanpomorphy for clade 19.Character 49 Lateral borders with triangularpockets (0) absent; (1) present (Figs 39, 44). The


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presence of triangular pockets on the lateral bordersis derived in clade 15.Character 50 Epigynal atrium (0) absent; (1) present (Fig. 35). An epiginal atrium is a synapo-morphy for the genus Tyrotama (clade 3).Character 51 Copulatory openings (0) incon-spicuous, (1) visible as circular or oval openings thatare either sclerotised or not (Fig. 55). This characteris a synapomorphy for clade 22.Character 52 Epigynum with oval median plate(0) absent; (1) present (Figs 42e-f). Baehr & Baehr(1993) proposed an oval central field on the epigy-num as a synapomoprhy for the genus Murricia. It is confirmed by this cladogram.Character 53 Copulatory openings relative tospermathecae (0) posterior (Fig. 32); (1) median(Fig. 79); (2) anterior (Fig. 36). A copulatory open-ing that opens anteriorly on the epigynal plate is a synapomorphy for Tyrotama species (clade 3) andthe clade comprising the genera of Iviraiva and Yabisi(clade 11), whereas a copulatory opening that opensmesally is derived in clade 9.Character 54 Copulatory ducts (0) narrow(Fig. 32); (1) wide, exceeding diameter of fertilisa-tion ducts (Fig. 72). Wide copulatory ducts wereindependently derived in Neotama (clade 7) andYpypuera.Character 55 Length of copulatory duct (0) verylong, at least three times spermathecae length(Fig. 32); (1) median, twice as long as spermathecaelength (Fig. 48); (2) short, equal (Fig. 40). The lengthof the copulatory duct is very homoplasious. A shortcopulatory duct is derived in clade 4 but reverts backto an elongate duct several times. Ducts of medianlength independently evolved in Murricia, clade 10,clade 20 and clade 25.Character 56 Copulatory duct (0) simple, straight(Fig. 40); (1) circular, looped (Figs 85-86); (2) regu-larly spirally coiled (Fig. 32); corkscrew-shaped(Baehr & Baehr 1993:Fig. 19f). A looped copulatoryduct is a synapomorphy for the genus Murricia(clade 13), whereas a corkscrew-shaped duct is anautapomorphy for Hersilia kinabaluensis.Character 57 Spermathecae (0) visible; (1) incon-spicuous. Inconspicuous spermathecae is an autapo-morphy for the genus IviraivaCharacter 58 Spermathecae basally with bul-bous sac (0) absent; (1) present (Fig. 56). A bulboussac basally independently evolved in H. savignyi andH. madagascariensis.Character 59 Spermathecae with glands basally(0) absent; (1) present (Fig. 44).The presence of these glands is a synapomorphy forthe clade 16.Character 60 Number of spermathecae (0) sev-eral spermathecae, more than 30 (Fig. 86); (1) one

large spermathecae (Figs 32, 40). Forster (1980) suggested a possible evolutionary sequence startingwith Grandungula sorenseni Forster (Gradungulidae),where the invagination of bursal wall with severalsmall spermathecae forms a stable bilobed arrange-ment of the spermathecae and a seminal receptacle.The possible loss of these secondary spermathecaewould then give rise to the bilobed structure found inHersilia sigillata group. The presence of multiple –more than 30 – small spermathecae are most appar-ent in Murricia uva (Fig. 49). Character 61 Spermathecae shape (0) globose(Fig. 40); (1) cylindrical (Fig. 44); (2) kidney-shaped(Fig. 68). Kidney-shaped spermathecae is an autapo-morphy of the genus Yabisi. A cylindrical spermath-ecae is derived in clades 6, 16, 21 and 25 and reversesin clade 10.Character 62 Spermathecae stalk (0) absent(Figs 41, 65); (1) present (Fig. 64). Rheims &Brescovit (2004) proposed stalked spermathecae assynapomorphies for Hersilia and Hersiliola, respec-tively. This tree topology supports this proposal forHersiliola but not for all Hersilia. A stalked spermath-ecae is derived in clade 12 and reverses in clades 21and 25.Character 63 Spermathecae and seminal recep-tacles clearly separated from ducts (0) absent(Fig. 79); (1) present (Fig. 40). A spermathecae thatis clearly separated from both the copulatory and fertilisation ducts is a synapomorphy for the genusHersilia (clade 14).Character 64 Number of seminal receptacles(0) absent; (1) several; (2) two; (3) one (Fig. 40); (4) one, large hyaline (Fig. 36) (Rheims & Brescovit2004). The definition of a seminal receptacle is takenfrom (Sierwald 1999: in Rheims & Brescovit 2004)as opening directly into the spermathecae and itsstalk with no connection to the fertilisation duct orcopulatory duct. The presence of a seminal recepta-cle unites all hersiliids (clade 1). A large hyaline sem-inal receptacle is a synapomorphy for Tyrotamaspecies (clade 3). Some representatives of the genusNeotama have two, whereas it is secondarily lost inYabisi and the presence of several seminal recepta-cles is an autapomorphy for Iviraiva. Character 65 Seminal receptacle shape (0) glo-bose (Figs 32, 36, 40); (1) irregular with apohysis(Fig. 64); (2) kidney-shaped (Fig. 44) (3) tube-like,complexly coiled (Fig. 86); (4) cylindrical. A cylin-drical seminal receptacle is derived in clade 4. A tube-like, elongate and complexly coiled seminal recepta-cle is a synapomorphy for the genus Murricia(clade 13). A kidney-shaped seminal receptacle is asynapomorphy for clade 16 and an irregularly shapedseminal receptacle with apophyses is a synapomorphyfor clade 25.

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Character 66 Seminal receptacle borders(0) smooth (Figs 32, 36, 48, 72, 79, 86); (1) with dis-tal tubercle (Figs 39, 44, 55, 64, 66, 68).Baehr & Baehr (1993a) referred to this character asglandular parts covered by threads. This character issynapomorphy for the genus Hersilia (clade 14). Itreverses in H. vanmoli and H. furcata.Character 67 Fertilisation duct (0) short, shorterthan spermathecae length (Fig. 40); (1) long, longerthan spermathecae length (Figs 60, 64); two verylong, twice as long as spermathecae length (Fig. 48).A long fertilisation duct is a synapomorphy forclade 8 and reverses in clade 11 (Iviraiva and Yabisi).It is also derived in clade 25. Very long fertilisationducts is derived in clade 18.

TAXONOMY

Family Hersiliidae Thorell 1870 (Figs 4, 5)

Herséliensis Simon, 1864: 343 (Lycosiformes, inpart).Hersilioidae Thorell, 1870: 109, 114; Ausserer, 877: 110.Chalinuroidea Thorell, 1873: 605; Kaston, 1938: 640 (Chalinuroidae = Hersiliidae).Hersilidae Simon, 1874: 14, 15 (lapsus).Hersiliidae Simon, 1882: 255; Lucas, 1869: 1; Simon,1893: 440; Smithers, 1945: 1; Benoit, 1967: 1; Baehr& Baehr, 1987: 351; 1993a: 3; Dippenaar-Schoeman& Jocqué, 1997: 181; Levi, 2003: 1; Rheims &Brescovit, 2004: 202; Wunderlich 2004: 814.

Type genus: Hersilia Audouin, 1827.

Diagnosis Posterior lateral spinnerets long, terminalsegment tapering, longer than basal segment; longestleg at least 1.4 times total body length. Hersiliidaeresembles Oecobiidae by the presence of elongateposterior lateral spinnerets and the way in which preyis captured: the spider rapidly encircles the prey andenswathes it by holding its spinnerets over the preyand fixing it to the substrate. They are distinguishedby the reduced anal tubercle; plumose setae that coverprosoma and abdomen; modified claw tufts sur-rounding tarsal claws (Fig. 5); 3-5 trichobothria dis-tally on metatarsi. Phylogenetics Six unambiguous synapomorphiesare hypothesised for the family Hersiliidae: The pres-ence of plumose setae (character 1[1]), fourmetatarsal trichobothria (character 5[1]), the pres-ence of claw tufts (character 7[1]), ALE < 0.7× AME(character 10[1]), posterior lateral eyes on slight orprominent tubercles (character 12[1&2]); presenceof one seminal receptacles (character 63[3]).

Description Size: small to medium (3-13 mm).Colour: varies from golden brown to white to almostblack with mottled appearance.Carapace: ovoid and flattened, as wide as long;narrow longitudinal fovea with three pairs of radi-ating striae; densely covered with plumose setae.Cephalic region moderately elevated, thoracicregion inclined or raised, higher than thoracicregion and thoracic region flattened. Eyes: eight intwo strongly recurved rows (Fig. 10); anteriormedian eyes often largest; lateral eyes on slight orprominent tubercle (Figs 17, 19). Clypeus lengthvariable, short (Fig. 15) or very elongate (Fig. 13)Chelicerae: unarmed (Tyrotama Tama and Hersi-liola); armed with three large teeth on promarginand 5-11 minute teeth on retromargin (Hersilia,Murricia, Neotama and Prima); labium free withrounded tip; endites oblique often touching eachother. Sternum: heart-shaped, anterior edge straightor slightly convex.Abdomen: dorsoventrally flattened, with 4-5 distinctdorsal muscular pits (Hersilia, Neotama, Prima andTama) (Fig. 10) or convex with four or less indistinctdorsal muscular pits (Tyrotama, Hersiliola) (Fig. 12);abdomen oval to triangular in shape; densely coveredwith plumose setae (Fig. 2); respiratory system: twobooklungs; tracheal spiracle opening close to spin-nerets; anal tubercle small, reduced. Spinnerets: six spinnerets; posterior lateral spinnerets longer thanprosoma width (Hersilia, Murricia, Neotama andPrima), shorter than prosoma width (Tyrotama Tamaand Hersiliola); cylindrical with elongated and taper-ing terminal segment; inner surface with series oflong spinules (see spinneret terminology in Codding-ton 1999) producing the thin silk threads (Fig. 3). Legs: three claws with modified claw tufts surround-ing claws (Fig. 5); unpaired claw simple with twoteeth; paired claws with 5-12 minute teeth, legs verylong; leg III shortest; all legs except leg III withmetatarsi uniarticulate (Tyrotama, Hersiliola andYabisi), elongate flexible zone distad (Prima, Tama,Tamposis and Iviraiva), narrow flexible zone (Neo-tama), biarticulate (Hersilia and Murricia); 3-5 tri-chobothria distad on metatarsi; legs with few spinesof which scale-like projections covering spine surface(Figs 6, 8, 9) (Hersilia, Murricia, Neotama, Prima), orspines dense, longitudinal grooves dorsally (Fig. 7)(Tyrotama, Tama and Hersiliola); autospasy occurs atpatella-tibia joint.Epigyne: Epigynal plate entire (Fig. 60) or longitu-dinally divided into three parts, a median plate andtwo lateral borders (Fig. 28); median plate free(Fig. 44) or slightly covered by lateral borders(Figs 36, 40); median plate smooth (Fig. 28) or withsub-triangular or sickle-shaped fixing structures laterally (Figs 36, 40), atrium surrounded by a


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sclerotised rim in Tyrotama (Fig. 32) or ripples(Fig. 45 dorsal view of epigynum). Copulation open-ings posterior (Fig. 29), medial (Fig. 80) or anterior(Fig. 33); copulatory ducts simple short (Fig. 37),simple, elongate, as long as epigynal plate (Fig. 45)or coiled (Fig. 29). Spermathecae unique (Fig. 37),several and small (Fig. 41) several and large (Fig. 49);spermathecae shape varies from globose (Fig. 37),cylindrical (Fig. 41) to subtriangular (Fig. 69). Seminal receptacle large, hyaline (Fig. 33), smallsclerotised (Fig. 37); fertilisation ducts short, curved(Fig. 37), elongate, straight (Fig. 45), elongate withmedial curve (Fig. 65).Palps: tibiae short as long as patellae, apophysis andmodified structures absent; cylindrical (Fig. 34) orwith dorsal projection (Fig. 43), spines absent(Fig. 35) or 2-7 strong spines present on promargin(Fig. 43); ventrally with several setae that extends infront of bulbus (Fig. 34). Cymbium covered withshort setae; 2-7 strong spines apically; cymbiumcompact, not projecting much beyond bulbus(Fig. 30); apex narrowing abruptly beyond bulbus(Fig. 75), digitate, (Fig. 26); bulbus, globose (Fig. 30),flattened (Fig. 26), round (Fig. 35) or basally swollen(Figs 59, 63); subtegulum semicircular, sclerotisedand entire; tegulum with tegular projection (Fig. 63);conductor, if present, accompanies the embolus(Figs 58, 62) or slide like tegular projection of varying sizes that accompany embolus often form-ing a fulcrum in which the embolus lies (Figs 81-83);embolus stout, hook-shaped (Fig. 50), filiform(Fig. 38); ridged (Fig. 42); embolic projections usu-ally absent, when present subapical (Fig. 81); embo-lus arise apical on bulbus (Fig. 82), basal (Fig. 26),prolateral (Fig. 42), or retrolateral (Fig. 58); spermduct with basal curvature (Fig. 22); basal loop(Fig. 23), repeated curvature (Fig. 24); basal loopwith medial curvature (Fig. 25). Median apophysis,if present, hook-shaped, convex, apex acute (Fig. 34)or hook-shaped, hollowed medially, apex acute ortruncate with or without basal projection (Figs 38,83) or complex (Fig. 42) or bifid (Fig. 74); medianapophysis insertion, medial (Figs 34, 38) or distal(Fig. 74). Composition Hersilia Audouin, 1826 – 73 species;Hersiliola Thorell, 1870 – seven species; MurriciaSimon, 1882 – five species; Tama Simon, 1882 – onespecies; Tamopsis Baehr & Baehr, 1987 – 49 species;Neotama Baehr & Baehr, 1993 – one species; Ypy-puera Rheims & Brescovit, 2004 – three species;Ivivraiva Rheims & Brescovit, 2004 – two species;Yabisi Rheims & Brescovit, 2004 – two species; Tyro-tama – nine species; Prima – one species.Distribution Tropical and subtropical zones,worldwide. The genus Hersilia from the Palearctic,Afrotopical, Oriental and Australian Regions;

Hersiliola from the Palearctic and AfrotropicalRegions; Murricia from the Oriental and Afrotrop-ical Regions; Prima from the Afrotropical Region;Promurricia from the Oriental Region; Neotamafrom the Oriental, Neotropical and AfrotropicalRegions; Tama from the Palearctic Region; Tyrotamafrom the Afrotropical Region; Ypypuera and Iviraivafrom the Neotropical Region; Yabisi from theNearctic and Neotropical Regions.

Key to the genera of Hersiliidae from theAfrotropical Region 1. Chelicerae unarmed; posterior lateral spinnerets

< carapace width; leg IV longest; thoracic regionof carapace sloping (Figs 10-11) ........................... 2— Chelicerae armed; posterior lateral spinnerets> carapace width; leg I or II longest; thoracicregion of carapace dorso-ventrally flattened(Figs 12-21) ............................................................... 3

2. Seminal receptacle, small and sclerotised; bulbusflattened with regularly curved sperm duct; embo-lus coiled, elongate, complete more than one cir-cle; median apophysis hook-shaped; tegular projec-tion, absent (Figs 29-30) ............ Hersiliola (Thorell)— Seminal receptacle large, unsclerotised; bulbusglobose, sperm duct with one or more switchbacks; embolus originate distally on bulbus, shortdoes not complete a full circle; median apohysisabsent; tegular projection, if present, short stoutdistad on bulbus (Figs 33-34) ................... Tyrotama

3. Metatarsi distally with flexible zone on legs I, IIand IV (Fig. 28) ........................................................ 4— Metatarsi biarticulate in legs I, II and IV(Fig. 27) ...................................................................... 5

4. Distal third of metatarsus I, II, IV flexible (Fig. 28).................................................................... Prima gen. n.— Narrow flexible zone in distal third of meta-tarsi I, II and III ...................................................................................................... Neotama Baehr & Baehr

5. Lateral eyes on slight tubercles, abdomen longerthan wide (Figs 12, 14, 16, 18), copulatory ductstraight simple, seminal receptacles simple spherical to subquadrate with distal tubercle(Figs 40, 44) ...................................... Hersilia Audouin— Lateral eyes on prominent tubercles, abdomenwider than long, subtriangular to subquadrate(Fig 20), copulatory ducts looped, seminal recep-tacles cylindrical, very elongate and complexlycoiled (Figs 85-86) ................ Murricia Simon, 1882

Prima gen. n.

Type species: Prima ansiae spec. n.Etmology Prima was the daughter of Hersilia,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 = aciniformgland 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, lateralview. 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, epigyneventral 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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Diagnoses Species of the genus Prima resemblethose of Tama, Tamopsis and Iviraiva by the presenceof an elongate flexible zone on metatarsi I, II and IV.The males are distinguished by an apically inserted,bifid median apophysis, short stout embolus (Fig. 74)and a sperm duct, starting from the base of the embo-lus, basally with a hairpin outward bend followed bya bend inwards medially (Fig. 26). The femalesresemble that of Ypypuera by the median copulatoryopenings on the epigynal plate and that of Promurri-cia by the sub-triangular median plate and indistinctseparation of the spermathecae and seminal recep-tacles. The absence of a biarticulation on metatarsi I,II and IV distinguishes it from Promurricia (Fig. 80). Phylogenetics Three unambiguous synapomorphiesare hypothesised for the clade Prima: a sperm duct,starting from the base of the embolus, basally with ahairpin outward bend followed by a bend inwardsmedially (character 32[5]), a bifid median apophysis(character 36[1]) and an elongate cymbium narrow-ing abruptly at the distal margin of the alveolus (char-acter 29[1]).Description Female Size: small, range (4.5-6).Colour: Carapace pale yellow; clypeus pale yellowwith dark or white markings; eye area dark aroundmedian ocular quadrangle and around posterior eyerow, white marks posteriad of eye tubercle in somespecies; sternum, labium and endites pale grey.Abdomen: dorsum mottled white; heart mark darkbrown, lancet-shaped extends up to third pair of dor-

sal muscular pits; antero-lateral border dark brown;venter pale grey to mottled white; femora and tibiaepale yellow with faint to dark brown annulation andlateral stripes. Carapace: as wide as long; thoracic region widest;cephalic region narrow; dorso-ventrally flattened;fovea longitudinal with radial striae; covered withplumose setae; clypeus truncate in dorsal view,sloping, short, varies in length from 0.46-0.97 ×median ocular quadrangle length; eye areadepressed to slightly raised (Figs 85, 86); ALEsmallest; AME ≥ PME ≥ PLE >> PLE with ratiorange AME:ALE:PME:PLE = 1: 0.38-0.83: 0.77-1.25: 0.54-1; chelicerae stout, retromargin with onerow of minute teeth, promargin with three largeteeth; sternum heart-shaped; labium half the lengthof endite, triangular to crescent-shaped; enditeselongate, rectangular.Abdomen: as long as wide, circular (Fig. 77); dorso-ventrally flattened; dorsum with lancet-shaped heartmark; four small dorsal muscular pits that vary insize; second pair largest; ventrum with V-shaped mus-cular pits. Spinnerets: posterior lateral spinneretslong, at least 2 × longer than carapace width; termi-nal segment > 4 × length of basal segment; spinuleson posterior lateral spinnerets conical, tapering dis-tally, apex truncate or acute.Legs: either leg I or II longest; very long, leg I at least3.23 × longer than total length of body; leg formulaI:II:IV:III; leg III very short, < 0.4 × length of leg I;


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

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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, epigyneventral view (71) dorsal view (72).

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metatarsus I > 6 × longer than tarsus I; metatarsi I,II and IV with elongate, flexible zone distally. Spinesshort spines < leg diameter; spine formula rather sim-ilar between species, variations of the following: I-Fe 1r1p-1d-1d-1r1p1d, Pat, Tib 1p, Mt 1p1r; II- Fe1r1p-1d-1d-1r1p1d-1r-1p-1d, Pat, Tib, Mt 1p1r; III-Fe 1d-1d-1d, Pat 1d, Tib, Mt; IV- Fe 1d-1d-1d-1r-1p1d, Pat, Tib 1p, Mt 1r1p. Spine microstruture withirregular patterns of lancet-shaped scales; tarsal clawswith seven tarsal teeth.Epigyne (Figs 78-79): Epygynum entire, white; cop-ulatory openings medially of spermathecae; copula-tory ducts with medial curve; spermathecae and sem-inal receptacles indistinctly separated; fertilisationduct medially curved.Male Size: Small (4.5). Resembles female in shapeand colour but differs structurally as follows: smallerin size; abdomen more slender.Palps (Figs 73, 74): tibia elongate; cymbium elongate,narrows abruptly, digitate, one spine apically; alveo-lus oval; bulbus flattened distally; sperm duct withmedial curve; median tegular apophysis apicallyinserted; embolus short, stout.Distribution Madagascar.

Prima ansieae new species Figs 73-79

Holotype: Ω: MADAGASCAR, FianarantsoaProvince: Parc Nationale, Ranomafana: Talatakely(21°14'S 47°25'E), 5-18.iv.1998, C.E. Griswold, D.H.Kavanaugh, N.D. Penny, M.J. Raherilalao, J.S. Rano-rianarisoa, J. Scweikert, D. Ubick, (CAS).Paratypes: 1 æ, 8 juveniles: same data; 1 æ: ParcNationale Ranomafana: 23 km N. Vohiparona village, (21°12'S 47°23'E), c.a. 1100 m, 10-11.iv.1998, C.E. Griswold, D.H. Kavanaugh, N.D.Penny, M.J. Raherilalao, J.S. Ranorianarisoa, J.Scweikert, D. Ubick, (CAS); 1 æ: AntsirananaProvince; PN Montagne d’Ambre, 2.79 km NE ofpark entrance (12°32'S 49°10'E), 21-30.xi.1993, J. Coddington, C. Griswold, N. Scharff, S. Larcher,R. Adrianmazinama, (CAS); 1 æ: Androrona(15º50'S 49º31E), x.1970, A. Lambillon, (MRAC42940).Etymology The specific epithet is a patronym inhonour of Dr Ansie Dippenaar-Schoeman for hercontribution to the knowledge of African spiders.Diagnosis Males are distinguished by the medialcurve of the sperm duct; median tegular apophysis,apically attached, bifid with concave flap laterally,medially with a hook-shaped process excised atapex (Fig. 73); females with copulatory duct withmedial bend, fertilisation ducts with basal loop(Fig. 79).

Description Male Carapace and clypeus palebrown; eye area dark with tridentate white spot pos-teriad on eye tubercle; abdomen white with antero-lateral dark border; cardiac impression linear broad-ening posteriorly; posterior half of dorsum with darkchevron markings (Fig. 77), distinct transverse lines;ventrum white; posterior lateral spinnerets withannulation; legs pale brown; femora and palps annu-late. Total length 5.36; carapace: 2.4 long, 2.4 wide;clypeus: 0.39 high, short, 4.88 × OAL; cheliceraeshort (l/w 0.39), cheliceral groove with eight dentic-ules on the retromargin; eye measurements: diame-ters: AME 0.26; ALE 0.07; PME 0.13; PLE 0.2; eyeratio: 1: 0.25: 0.5: 0.75; interdistances: AME-AME0.08; PME-PME 0.08; PME-PLE 0.07; AME-PME0.47; legs: I – Fe 7.2; II – 8; III – 3.12, 2.4, 2.88, 0.8,total 9.2; IV – 7.6; Palp – 1.33, 1.61, 0.91, total 3.85;opisthosoma length: 3.72, width: 2.4; DMP round;ventral muscular pits V-shaped; PLS, bS: 0.8, tS: 4,tS/bS 5, 2 × CW.Palps (Figs 73, 74): Tibia short, 0.53 × patella, sim-ple, dorsal tibial spines absent; cymbium digitate, 2.8× longer than wide, one apical spine; row of longwhite setae on prolateral border of cymbium extendsacross bulbus; bulbus distally truncate, sperm ductcircular with basal and medial curvature (Fig. 73);embolus short, slightly curved, apex acute; mediantegular apophysis apically attached; hollowed, bifid,laterally with concave flap and medially with curvedprocess excised at apex; embolus short, stout, apexacute. Female Size (n = 3). Colouration and pattern as in males. Total length 5.62 (5.25-6); carapace: 2.06 (1.84-2.16) long, 2.14 (2-2.16) wide; clypeus:0.28 (0.26-0.3) high, clypeus short, 2.39 × OAL;chelicerae: (l/w 1.67); eye measurements: diame-ters: AME 0.14; ALE 0.11; PME 0.2; PLE 0.15; eyeratio: 1: 0.68: 1.04: 0.82; interdistances: AME-AME0.21; PME-PME 0.13; PME-PLE 0.14; AME-PME0.11; legs: I – Fe 4.73, Pat + Tib 5.23, Mt I 6.54, Ta0.81, total 16.88; II – 4.86, 6.12, 7.69, 0.85, total18.79; III – 1.66, 1.96, 1.73, 0.7, total 5.71; IV –4.69, 3.78, 5.4, 0.9, total 14.31; Palp – 1.09, 0.98,0.91, total 2.98; legs long, 3.23 × body length; legratio: 1: 1.11: 0.34: 0.96; opisthosoma length: 3.16,width: 3.16; DMP round; ventral muscular pits V-shaped; PLS bS: 0.91, tS: 3.93, tS/bS 4.37, 2.26× CW.Epigynum (Figs 79, 80): wide (el/ew 0.5); epigynewith trapezoid central unpigmented field, copulatoryducts with medial, fertilisation ducts with basal loopduct, medially curved distally.Distribution Madagascar (Fig. 94).Natural history Specimens were collected at1100 m above sea level; females collected in Novem-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

Murricia Simon, 1882: 255; 1893: 446; Pocock,1900: 242; Baehr & Baehr 1993: 64.Type species: Murricia indica (Lucas, 1836).Diagnosis Murricia resembles Hersilia by the pres-ence of a biarticulation on metatarsi of legs I, IIand IV. The males are distinguished by a longembolus, completing one circle, which is positionedapically on the bulbus. The females are distin-guished by the presence of wide, elliptical poste-rior-median field, very long and looped copulatoryducts, and the presence of one tube-like, elongateand complexly coiled seminal receptacle. Theopisthosoma is often subtriangular to subquadrate.Phylogenetics Six unambiguous synapomorphiesare proposed for the genus Murricia: a sperm ductthat, starting from the base of the embolus, has aninward bend medially followed by a large outwardbend, which in turn is followed by a medial inwardbend and a basal loop (character 32[4]), an distallyinserted median apophysis (character 35[3]), a longembolus that completes at least one circle (charac-ter 42[1]), a copulatory duct that is twice the lengthof the spermathecae (character 54[1]), a looped,circular copulatory duct (character 55[1]) andtube-like, complexly coiled seminal receptacles(character 64[3]). Description Female Size: small to medium, range(4.52-6.38). Colour: Carapace varies from pale yellow to darkbrown, with dark rim of varying thickness aroundborder; clypeus pale with dark or white markings;eye area dark, white mark posteriad on eye tuber-cle; sternum pale with mottled white markings;labium and endites pale. Abdomen: dorsum withmottled white background; heart mark narrow,elongate, along entire length of abdomen; antero-lateral border dark brown; venter pale to mottledwhite. Legs: femora and tibiae pale with faint todark annulation and lateral striae. Carapace: varying between longer than wide to aswide as long; thoracic region widest, cephalicregion narrow; dorso-ventrally flattened; fovea lon-gitudinal with radial striae; clypeus truncate in dor-sal view, not projecting much beyond eye area,varies in length from 0.88-0.94 × median ocularquadrangle length; eye tubercle slightly raised; eyes:PER recurved seen from above and in front; AERrecurved seen from above and in front; ALE small-est, translucent; PLE > PME <> AME >> PLE withratio range AME:ALE:PME:PLE = 1: 0.27-0.42:0.92-1.3: 1.28-2; median ocular quadrangle widerthan long; MOQ-AW = MOQ-PW; cheliceraerather stout, 1.61 × longer than wide, retromargin

with row of four minute teeth, promargin withthree large teeth. Sternum: heart-shaped; labium:sub-triangular, two-thirds the length of endites;endites stout, subquadrate, broadening anteriorly. Abdomen: wider than long, sub-triangular; dorso-ventrally flattened; four to five pairs of distinct dor-sal muscular pits that vary in size (Fig. 20); venterwith V-shaped muscular pits. Spinnerets: posteriorlateral spinnerets long > 1.5 × carapace width; ter-minal segment ≥ 4 × length of basal segment. Legs: rather short ≤ 2.22 × total body length; eitherleg I or II longest; leg formula I:II:IV:III; leg III veryshort, < 0.4 × length of leg I; metatarsus I > 6 ×longer than tarsus I; distal segment of metatarsus> 0.7 × length of proximal segment; femur, patella,and metatarsus with spines, spine formula tends tobe rather similar between species and were varia-tions of the following: I- Fe 1p1d1r-1p1d1r-1p1d1r-1p1r, Pat 1d1d, Tib 1d-1p-1d1r-1p1d1r,Mt 1p1r-1d; II- Fe 1p1d1r-1p1d-1r-1p1d-1p1r, Pat1p1r-1d, Tib 1d-1p1r-1d-1p1d1r, Mt 1p1r-1p-1d;III- Fe 1d-1d-1d-1r, Pat 1d-1d, Tib 1d-1v-1d, Mt1d-1d; IV- Fe 1p1d-1d1r-1d-1p1r, Pat 1d-1r-1d,Tib 1d1r1p-1d-1p1r1d, Mt 1r1p-1r1p-1d; spinemicrostrucutes with randomly arranged conicalscales covering spine surface. Epigynum: ovate depressions posterior of copula-tory openings (Fig. 85); copulatory ducts elongate,looped; seminal receptacles cylindrical, complexlycoiled (Fig. 86); fertilisation duct curved medially.Remark Forster (1980) suggested a possible evo-lutionary sequence starting with Grandungulasorenseni (Grandungulidae) where the invaginationof bursal wall with several small spermathecae formstable bilobed spermathecae. The numerous sec-ondary spermathecae found in Afrotropical femalesof Murricia are hypothesised to be homologous tothese small spermathecae.Male Size: small (4.88-5.03). Resemble female inshape and colour; differs structurally as follows:smaller in size; abdomen more slender, sub-quadrate, widest posteriorly; distal segment ofmetatarsus > 0.6 × length of proximal; metatarsusI > 10 × tarsus I, legs relatively longer, 3.45 × totalbody length.Palps (Figs 80-81): tibia stout, dorsal spines absent;cymbium compact; bulbus large, round, mediantegular apophysis, if present small, triangular, hook-shaped, distally attached; embolus circular, apexacute, originate distally on bulbus.Composition Murricia uva sp. n. (Kenya, Uganda,Rwanda, Cameroon), M. cornuta Baehr & Baehr1993 (Singapore), M. crinifera Baehr & Baehr 1993(Sri Lanka), M triangularis Baehr & Baehr 1993(India).

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Murricia uva new speciesFigs 80-86, 95

Holotype: 1 Ω: KENYA, from Rift Valley Province,Marich Pass, Field Studies Centre (1º32'S, 35º27'E),7.vi.1999, W.J. Pulawski, J.S. Schweikert (CAS). Paratypes: CAMEROON: 1 Ω: Chabal Mbabo, SWslope (07º25'N, 12º49'E), 1300 m a.s.l., grassland,12.iii.1981, Bosmans & van Stalle, (MRAC162559); CONGO D.R.: 1 Ω: Avakubi (01º19'N,27º33'E), vii.1932, B. Randoux (MRAC 137384);UGANDA: 1 æ: Semliki Forest (0º48'N, 30º8'E),5.-12.ii.1997, Wagner, deposited in MNHU;RWANDA: 1 æ: PN Akagera, 6 km S de la pêcherie,foret seche a Sansevieria (15°5'S, 30°45'E), 14.xi.-3.xii.1985, R. Jocqué, Nsengimana, Michiels(MRAC 165429); 1 æ: PN Akagera, 50 km Nde lapêcherie Ihema, pres du lac Mihindi (13°2'S,30°43'E), 6.xii.1985, R. Jocqué, Nsengimana,Michiels, (MRAC 165058).

Etymology The specific epithet is from latin (uva-um= ‘bunch of grapes’) and refers to the large number ofspermathecae abutting a central stalk, adjective.Diagnosis The males of M. uva sp. n. resemble thatof H. jajat Rheims & Brescovit (2004:2852, Figs 1-4 (figures of palp was used for this comparison)) inthe elongate and apical embolus and truncated cym-bium (Fig. 80). They are distinguished by the pres-ence of a median apophysis (Fig. 81). Females of M. uva sp. n. resemble that of H. jajat sp. n. Rheims& Brescovit (2004b:2852, Figs 5-6) by the long andcircular copulation duct and the long tube-like seminal receptacles. They are distinguished by theseveral, spherical spermathecae abutting a centralstalk (Fig. 86).Description Male Size (n = 2). Carapace red-brown to pale yellow, dark broad posterior border,distinct longitudinal fovea; clypeus with dark medialline; eye area dark; opisthosoma: dorsum white withdark markings, dark antero-lateral border; cardiacimpression lancet-shaped, as long as opisthosoma(Fig. 84); dorsum pale; posterior lateral spinneretswith strong annulation; legs pale with striations onmargins, patellae dark; palps with tarsi dark. Totallength 4.95 (4.88-5.03); carapace: 2.25 long, 2.03(1.95-2.1) wide; clypeus: 0.32 high, short, 2.47 ×OAL; chelicerae long (l/w 1.85), cheliceral groovewith four denticules on the retromargin; eye meas-urements: diameters: AME 0.2; ALE 0.08; PME0.17; PLE 0.18; eye ratio: 1: 0.41: 0.85: 0.9; interdis-tances: AME-AME 0.16; AME-ALE 0.16; PME-PME 0.12; PME-PLE 0.25; AME-PME 0.5; legs: I– Fe 4.28, Pat + Tib 4.8, Mt I 3.98, II 2.4, Ta 0.6,total 16.29; II – 4.35, 5.33, 4.13, 2.48, 0.6, total16.51; III – 1.65, 1.76, 0.9, 0.38, total 4.65; IV –

3.68, 4.2, 3.75, 2.1, 0.6, total 14.25; Palp – 0.9, 0.83,0.75, total 2.55; legs long, 3.33 × body length; leg ratio: 1: 1.03: 0.32: 0.88; opisthosoma length:2.78, width: 2.85; five dorsal muscular pits; DMPelliptic, second pair by far the largest, third pair thesmallest; ventral muscular pits form two converg-ing lines; bS: 0.75, tS: 3, tS/bS 4, 1.92 × CW.Palp (Figs 80-81): Tibia short, as long as patella,tibia simple; cymbium with three apical spines;embolus circular, filiform, apex acute; medianapophysis distally inserted (Fig. 80), apex acute, pro-laterally directed with rugous surface (Fig. 81). Female Size (n = 4). Carapace pale yellow, sparselycovered with white setae; clypeus pale yellow;median ocular quadrangle dark; opisthosoma later-ally covered with dense long white setae; mottledwhite with dark markings postero-laterally; dor-sum with narrow lancet-shaped cardiac impression(Fig. 20); posterior half of opisthosoma with tri-angular darkly mottled area; legs and palps pale yel-low, without annulations. Total length 5.56 (4.52-6.38); carapace: 2.13 (2.03-2.25) long, 2.22(2.03-2.48) wide; clypeus: 0.39 high, short, 3.11 ×OAL; chelicerae long: (l/w 1.61), cheliceral groovewith ten retromarginal denticules; eye measure-ments: diameters: AME 0.14; ALE 0.04; PME 0.16;PLE 0.23; eye ratio: 1: 0.36: 1.1: 1.63; interdis-tances: AME-AME 0.16; PME-PME 0.13; PME-PLE 0.2; AME-PME 0.13; legs: I – Fe 3.9, Pat + Tib3.86, Mt I 2.81, II 2.06, Ta 0.64, total 13.13; II –4.01, 4.14, 2.81, 2.06, 0.6, total 13.35; III – 1.53,1.57, 1.08, 0.55, total 4.77; IV – 3.57, 3.39, 2.89,1.8, 0.56, total 12.19; Palp – 0.89, 1.11, 0.75, total2.63; legs short, 2.22 × body length; leg ratio: 1:1.02: 0.37: 0.93; opisthosoma length: 3.26, width:4.43; five dorsal muscular pits; DMP round; ventralmuscular pits V-shaped; bS: 0.78, tS: 3.23, tS/bS4.1, 1.71 × CW.Epigynum (Figs 85-86): Elongate, (el/ew 0.9);externally with two longitudinal oval depressions;fertilisation duct short, medially curved.

Distribution Cameroon, Kenya, DemocraticRepublic of the Congo, Rwanda, Uganda (Fig. 95).

Natural history The male paratype was caught atan altitude of 1300 m a.s.l., in grassland. Some ofthe type specimens were collected from trees in theforests biome.

ACKNOWLEDGEMENTS

I wish to thank the curators for loaning material for this study; the researchers and technicians of the Biosys-tematic Division/Arachnida Unit of the ARC Plant Pro-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 malefrom 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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Dippenaar-Schoeman (NCA), Prof. Clarke Scholtz (Uni-versity of Pretoria), Rudy Jocqué (MRAC), CristinaRheims (Instituto Butantan, Brazil), Barbara Baehr (QM)and the two anonymous reviewers who made helpful sug-gestions on this manuscript, which was presented as partof a PhD thesis at the University of Pretoria in February

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

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