Revision and cladistic analysis of the jumping spider genus Onomastus (Araneae: Salticidae)

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1 Zoological Journal of the Linnean Society, 200, 59, With 28 figures Revision and cladistic analysis of the jumping spider genus Onomastus (raneae: Salticidae) SURSH P. NJMIN* Department of ntomology, National Museum of Natural History MR 05, PO ox 3702, Smithsonian Institution, Washington, D , US Received 5 September 2008; accepted for publication 25 February 2009 The jumping spider genus Onomastus Simon, 900 is revised. Four new species: Onomastus indra sp. nov., Onomastus kaharian sp. nov., Onomastus pethiyagodai sp. nov., and Onomastus rattotensis sp. nov. are described. Parsimony analysis of 26 morphological characters supported the monophyly of Onomastus. Lyssomanes is sister to Onomastus. Onomastus separates into two clades: the widespread South-ast sia clade and the South sia clade. The South sia clade is restricted to the Sri Lanka Western Ghats biodiversity hotspot. Species of the South sia clade appear to be spot endemics, highly in danger of extinction because of habitat loss and climate change. Male palps are complex and species-specific, suggesting rapid divergent evolution.. doi: 0./j x DDITIONL KYWORDS: biodiversity climate change hotspots India South-ast sia Sri Lanka. INTRODUTION Our current understanding of the evolutionary relationships of jumping spiders, the largest spider family, suggest at least four major lineages: the lyssomanines, the spartaeines, the ocalodes group, and the much larger Salticoida clade (Wanless, 980a, 984; Maddison, 996; Maddison & Hedin, 2003; Maddison & Needham, 2006). The three non- Salticoida clades have recently been collectively termed basal salticids (Maddison & Needham, 2006). Jumping spiders of the genus Onomastus Simon, 900 are lyssomanines (Wanless, 980b). They are small to medium sized spiders (2.0 to 5.2 mm total length) that live on foliage of tropical evergreen forests. They stalk for prey and do not build capture webs. They resemble other lyssomanines in coloration, general slender body shape, and by having the eyes arranged in four transverse rows (Wanless, 980b). However, in contrast to Salticoida and other lyssomanines and spartaeines, Onomastus male palps are unusually complex and provide enough *-mail: suresh.benjamin@gmail.com. urrent address: Institute of Fundamental Studies, Hantana Road, Kandy, Sri Lanka. information to clearly distinguish amongst closely related species. xcept for Onomastus complexipalpis sp. nov., female genitalia are less complex and more similar in most aspects. urrently lyssomanines are defined by the presence of four rows of eyes and a tracheal system that is confined to the opisthosoma (Galiano, 962; Wanless, 980a). Wanless (980a) used the eye pattern to define groupings of lyssomanine genera: semonea O. P.-ambridge, 869, Goleba Wanless, 980, Macopaeus Simon, 900, and Pandisus Simon, 900 was termed group III; group III was considered by him to be related to group II, which consisted of the genera Lyssomanes Hentz, 845 and hinoscopus Simon, 90. Onomastus was however, considered not closely allied to either of them and was thus placed in group I (Wanless, 980a). However, these hypotheses have not been tested in a phylogenetic context. The type species of Onomastus, Onomastus nigricauda Simon, 900 is endemic to lowland evergreen rainforests of western Sri Lanka. further two species, Onomastus quinquenotatus Simon, 900 from Sri Lanka and Onomastus patellaris Simon, 900 from the Western Ghats of southern India, were described by Simon (900). The first species outside the Indian subcontinent, O. complexipalpis was 7

2 72 S. P. NJMIN described in a comprehensive treatment of the genus by Wanless (980b). However, by 980 only O. nigricauda and O. patellaris were known by both sexes. Recently several species from ast sia have been described: Onomastus simoni Zabka, 985 from Vietnam (male unknown), Onomastus kanoi Ono, 995 from Okinawa, Japan (female unknown), and Onomastus nigrimaculatus Zhang & Li from hina (Zabka, 985; Ono, 995; Zhang & Li, 2005). s part of an on-going survey of spider diversity in Sri Lanka, a large number of Onomastus was collected. They turned out to be of four species, two known and two new. ll seem to be endemic and restricted to small patches of higher elevation rain forest. dditionally, a collection of South-ast sian Onomastus from the collections of hrista L. Deeleman-Reinhold has been incorporated. further undescribed species from the Western Ghats was also discovered in the collections of the S. Here, I provide comprehensive accounts of all Onomastus from Sri Lanka and the surrounding region with descriptions of new species and updated diagnosis. dditionally, higher-level relationships of Salticidae are discussed based on a cladistic analysis of Onomastus plus exemplars of Wanless s group I III (Wanless, 980a). TRIL ND MTHODS Methodology and taxonomic descriptions follow the format of enjamin (2004, 2006). Specimens used for illustrations were placed on washed sand in 70% ethanol and photographed using a dissecting microscope (Leica MZPO) with top illumination and a magnification of up to 50. Digital images were taken with a Nikon DXM200F camera. Images were edited using an uto-montage software package. ither a LO 430VP or an mray 80 scanning electron microscope (SM) was also used to study and photograph morphological features. Prior to SM work specimens were prepared as mentioned in enjamin (2004) and mounted on to aluminium rivets with the help of conductive copper adhesive tape or wire. Such mounting, although rather intricate, enabled rotation of specimens in the SM chamber. Male palps were cleared in methyl salicylate and mounted on temporary slides to observe and illustrate their internal structures. Left structures (e.g. palps, legs, etc.) are depicted unless otherwise stated. Hairs and macrosetae are usually not depicted in the final palp drawings. ll measurements are given in millimetres and were made with a compound microscope (Leica MZPO) equipped with a 0 ocular and an ocular micrometre scale. bbreviations used in the text and figures are given below. Leg spination is described according to the system adopted by Platnick & Shadab (975). complete synonymy of the species is given in Platnick (2009). RVITIONS Institutional MNH, ritish Museum, Natural History, London; S, alifornia cademy of Sciences, San Francisco; MHNG, Muséum d Histoire Naturelle, Genève; MNHN, Muséum National d Histoire Naturelle, Paris; RMNH, Naturalis, National Museum of Natural History, Leiden; USNM, National Museum of Natural History, Smithsonian Institution, Washington, D. Morphological structures, anterior epigynal border; L, anterior lateral eye; M, anterior median eye; P, apical plates of epigynum;, bulbus; H, basal haematodocha;, conductor; D, copulatory duct; O, copulatory opening; Y, cymbium;, embolus; F, epigynal folds; P, embolic base; F, femur; FD, fertilization duct;, median apophysis; P, mesal branch of ; MP, mate plug; P, patella apophysis; PL, posterior lateral eye; PM, posterior median eye; PT, patella; RT, retrolateral tibial apophysis; S, spermatheca; SP, spur mesal branch of ; ST, subtegulum; T, tibia; T, tegular apophysis (Figs 4, 0, 7); T2, tegular apophysis 2 (Figs 4, 7); T3, tegular apophysis 3 (Figs 4, 9, 28D). LDISTI NLYSIS Outgroup choice The primary aim of the cladistic analysis was to test the monophyly and the internal structure of Onomastus. Unfortunately, relationships of basal salticids are far from satisfactorily resolved, complicating the selection of the putative outgroup. The selection of outgroup taxa was based on Wijesinghe (997), Wanless (980a), and Maddison & Needham (2006). Thus, I have included the genera Onomastus, Lyssomanes, Pandisus, semonea, and Goleba. Further, as suggested by Maddison & Needham (2006), I have also included Hispo Simon, 886, a possible sister to Lyssomaninae. Taxa and characters scored The taxa and characters used served the main purpose of the study, which was to test the monophyly and the internal structure of Onomastus and not to tackle higher-level relationships amongst salticids. s in my previous work (enjamin, 2004), I consider all character statements to be hypotheses of homology, unless putative homology of traits of two or more taxa

3 RVISION ND LDISTIS OF ONOSTUS Lyssomaniae Hispo cingulata are rejected by the cladistic analysis (Fig. ). ll Onomastus species except for O. simoni are included in the phylogenetic analysis (Table ). Onomastus simoni is only known from a single female specimen, additional material was unavailable for study. ll characters are scored directly from voucher specimens. However, as a result of a lack of material in collections and to some extent because of the reluctance of curators to lend material for SM work, Lyssomanes virides 95 Onomastus Pandius sarae semonea tenuipes 8 0 Pandisus clade Goleba puella O. nigrimaculatus O. kaharian O. pethiyagodai O. quinquenotatus 0 South-east sia clade O. complexipalpis O. kanoi O. nigricauda O. patellaris O. indra O. rattotensis South sia clade Figure. Preferred most parsimonious tree of Onomastus based on the character matrix presented in Table (length = 32 steps, consistency index = 0.8, retention index = 0.89). haracter state changes are mapped using Farris optimization. losed circles represent unambiguous character changes. outgroup taxa were scored on the basis of published accounts. Male palp. F: absent/femora smooth, 0; present,. F is defined here as any structure projecting outwards from the surface of the femur. Present in semonea, Pandisus, and Goleba (Wanless, 980a: figs 4e, 8c). In Onomastus femora are smooth and

4 74 S. P. NJMIN Table. Distribution of characters scored for ten Onomastus taxa and outgroups Onomastus quinquenotatus Onomastus nigrimaculatus Onomastus nigricauda Onomastus patellaris Onomastus indra sp. nov Onomastus pethiyagodai sp. nov Onomastus rattotensis sp. nov Onomastus complexipalpis Onomastus kanoi Onomastus kaharian sp. nov Pandisus sarae* semonea tenuipes* Goleba puella* Hispo cingulata* Lyssomanes viridis* unmodified, lacking any furrows or apophyses as found in semonea and Pandisus (Fig. 26; Wanless, 980b). 2. Position of F: retrolateral/distal, 0; ventral/ median,. Refers to the position of the F mentioned above. F is more towards the ventral surface of the centre of the femur in Goleba (Wanless, 980a: figs 22c, d, 24b), whereas it is in a distal retrolateral position in semonea and Pandisus (Wanless, 980a: figs 4e, 8c). Wanless (980a) described state found in Goleba as ventral tubercle on the palpal femur O. quinquenotatus O. nigrimaculatus O. nigricauda O. patellaris O. indra sp. nov O. pethiyagodai sp. nov O. rattotensis sp. nov O. complexipalpis O. kanoi 0 0 0??? 0 0 O. kaharian sp. nov P. sarae* tenuipes* G. puella* H. cingulata* L. viridis* haracter states are scored 0 to 2,? for unknown, for inapplicable. Outgroup taxa are denoted with an asterisk. 3. Femoral groove: absent, 0; present,. Femoral groove is defined as a cavity of the surface of the femur. It does not project outwards as is the case in the F. The femoral groove is only present in semonea (Wanless, 980a: figs 0e, 3c). In Onomastus femora are smooth and unmodified, lacking any furrows (Fig. 26). 4. P: absent, 0; present,. Present in all Onomastus and Pandisus (Figs 9, 0, 4F, 7; Wanless, 980a; Wanless, 980b). Onomastus lack an RT, instead the patella of the male palp is armed with an apophysis at the distal end that

5 RVISION ND LDISTIS OF ONOSTUS 75 probably functions as a replacement for the RT. P is also present in Pandisus, as illustrated for Pandisus scalaris by Wanless (980a: fig. 3). One would consider the P to be homologous in both genera based on the criteria of Remane (952). However, it appears to be otherwise (Fig. ). 5. P shape: tapering, 0; blunt,. P is rather tapering with a smooth rounded tip in all Onomastus, except O. complexipalpis and O. kanoi. In O. complexipalpis and O. kanoi the P broadens towards the end (Figs 3, 3, 7F; Ono, 995). 6. Relative size of patella: patella = tibia, 0; patella > tibia,. The relatively elongated patella is unique to the two Indian Onomastus, O. patellaris and Onomastus indra (Figs 4,, 7). Hence the origin of the specific name patellaris, which was the first Onomastus species with an elongated patella to be described from India; a second is described below. 7. RT: present, 0; absent,. RT is absent in Onomastus. s mentioned above, the distal ends of the tibia of male palps of all Onomastus are smooth. s a member of the RT clade one would expect to see a RT in all Onomastus. However, this is not the case and it is presumed lost in all species (Wanless, 980b). 8. onductor: absent, 0; present,. very complex conductor is present in all known Onomastus species and is species-specific in size and shape (Figs 2F, G, 5,, 9; Wanless, 980b). 9. Type of conductor: hyaline, 0; sclerotized,. hyaline conductor also termed membraneous secondary conductor by Wanless (980a: fig. 2g, h), is a conductor that is transparent and very thin. hyaline conductor is present on hinoscopus Simon, 90 and Lyssomanes (Galiano, 962, 980; Wanless, 980a: fig. 2g, h). The presence of a hyaline conductor might turn out to be a synapomorphy that unites hinoscopus and Lyssomanes, two exclusively New World genera. 0. Shape of conductor: filiform, 0; stout, ; broad, 2. onductor shape varies from an elongated filiformapophysis to a large disk-shaped structure. Scored as filiform for Onomastus of Sri Lanka (Figs D, 9, ), as stout for Onomastus of India (Figs 4, 7), and as broad for the species of the South-ast sia clade (Figs 2, 3, D, ).. Number of hooks on the tip of : one, 0; two,. The conductor tip of O. indra, O. kano, and O. patellaris is furnished with a single hook (Figs 4, 7,, 7). The rest have a bifurcated tip ending with two pointed hooks (Fig. ). 2. : absent, 0; present,. ll species of Onomastus have a heavily sclerotized. They are branched and species-specific in shape (Figs 9,, D, 4D,, 7, 25G, H; Wanless, 980b). is present in Hispo (Wanless, 98) and Lyssomanes. However, it is absent in hinoscopus (Wanless, 980a). 3. distal end: bifurcate, 0; smooth/tapering,. In O. nigricauda, O. rattotensis, and O. complexipalpis, the is distally bifurcated, diverging into two branches (Figs D, 28). Smooth or tapering in the rest (Figs 4D,, 7, 25G, H). 4. P: absent, 0; present,. The has a mesal branch that supports the conductor embolus complex, which is present in all known species from Sri Lanka and India (Figs 4, D, 9, 27). 5. Origin of embolus: dorsal, 0; ventral,. The embolus originates from the tegulum within the alveolus in all Onomastus species (Fig. 9, 5, ; Wanless, 980b). The origin of the embolus is visible when male palps are cleared in methyl salicylate. It is also clearly visible in the expanded palp (Fig. 9). This character is diagnostic for Onomastus. 6. T: absent, 0; present,. Onomastus male palps possess three different tegular structures. ach originates from different positions of the tegulum. I call them T, T2, and T3, to avoid mislabelling any one of them as homologous to the tegular apophysis (or T) found in most spider male palps. T ( y in Wanless (980b); subtegular apophysis in Zhang & Li (2005) is unique to Onomastus (Figs 4, 0, 4F, 7, 25,, 28). T is not known to occur in any other Salticidae (Wanless, 980a, b). This character is diagnostic for Onomastus. 7. T2: absent, 0; present,. The presence of T2 ( x in Wanless (980b) is unique to the two Indian Onomastus, O. patellaris and O. indra (Figs 4, 7). In addition to T the male pulps of Indian Onomastus carry a second tegular apophysis termed T2 in this study. It originates from the prolateral margin of the male palp, distally to the T (Figs 4, 7). 8. T3: absent, 0; present,. T3 is situated distally to T and T2, on the ventral/distal surface of the tegulum (Fig. 25H, 28D).

6 76 S. P. NJMIN T3 is present in all known species from Sri Lanka and India (Figs 4, 9, 9, 25, H, 27, 28D). Female genitalia 9. Female copulatory ducts: absent, 0; present,. opulatory ducts are absent in Onomastus (Wanless, 980b). The exceptions are Onomastus kaharian, O. complexipalpis, and O. simoni, all of the South-ast sia clade. It is present in all outgroup taxa (Wanless, 980a).The female of O. kanoi is unknown; however, I predict that it would have copulatory ducts. 20. F: absent, 0; present,. F, also called septum in Zhang & Li (2005) are absent in most Onomastus, instead the O is a simple opening in the cuticle (Wanless, 980b). The exceptions are species of the South-ast sia clade (Fig. 5 ; Zhang & Li, 2005). F are also absent in the outgroup taxa (Wanless, 980a). 2. pigynal lip: absent, 0; present,. Present only in Goleba puella (Wanless, 980a). Somatic morphology 22. Number of eye rows: three, 0; four,. Three considered in Hispo cingulata (Wanless, 98); four in all other taxa in the matrix (Galiano, 962, 980; Wanless, 980a, b). 23. Relative size of PM: PM < PL, 0; PM = PL,. PM = PL in semonea and Goleba. PM < PL in the rest. Outgroups are scored based on Wanless (980a). This character might be informative in much broader cladistic studies as well. 24. Relative distance between PM PM vs. L L: PM PM subequal or longer than L L, 0; PM PM < L L,. This character was proposed to define a grouping of genera consisting of semonea, Goleba, Macopaeus, and Pandisus by Wanless (980a), which he termed group III. Group III was considered by him to be related to group II, which consisted of the genera Lyssomanes and hinoscopus. Onomastus was however, considered not closely allied to either of them and was, thus placed in group I (Wanless, 980a). This character might be informative in much broader cladistic studies as well. 25. Relative size of PL to PM: much smaller, 0; similar in size,. PM are similar in size in semonea and Goleba. PM are much smaller in the rest. Outgroups are scored based on Wanless (980a). 26. Prosoma coloration: red/brown, 0; light green,. Lyssomaninae are considered to be transparent green in colour (Wanless, 980a; Maddison & Needham, 2006). However, available information in the literature on prosoma coloration is rather scanty. ll Onomastus are clearly green (Figs 8 D, 8 F) and so are semonea tenuipes and Lyssomanes viridis (pers. observ.). The remaining taxa are scored based on the their descriptions by Wanless (980a, 98). NLYSIS The parsimony analyses were performed using the computer program NON 2ed (Goloboff, 999). Win- lada version (Nixon, 999) and MSQUIT version.2 (Maddison & Maddison, 2006) were used to study character optimizations on the cladograms and to build and edit the character matrix, respectively. mbiguous character optimizations were resolved so as to favour reversal or secondary loss over convergence [Farris optimization or accelerated transformation (TRN)]. ll multistate characters were treated as non-additive (unordered or Fitch minimum mutation model; Fitch, 97). Missing characters were coded as?, and inapplicable characters as -. ranches were collapsed if the minimum possible branch length was zero ( amb- ) or if there was no character that could be optimized to support a node. Tree statistics were only calculated from phylogenetically informative characters. I also used PUP (Swofford, 2002) to analyse the data as carried out in enjamin (2004); see enjamin (2004) for justification of the analytical parameters. elow, characters and states are abbreviated and in bold, e.g. character 2 becomes 2, character 2, state, becomes 2-. RSULTS ONOSTUS RLTIONSHIPS The phylogenetic analysis of the above-mentioned, equally weighted characters performed in NON (hold0000; hold/200; 000 replicates of tree bisection-reconnection (TR) + TR (mult*000) produced one tree of minimal length. nalysis of the same data set in PUP with heuristic search methods (00 replicates of random taxon addition subjected to TR branch swapping; MulTrees set to 00000; branches collapsed if the minimum possible branch length was zero ( amb- ; TRN or Farris optimization) produced the same minimal length tree found with NON. This tree was selected as the preferred phylogenetic hypothesis for Onomastus (Fig. ). Onomastus is resolved at node 3 supported by four unambiguous synapomorphies. Within Onomastus, species are grouped into two clades, which are named here as

7 RVISION ND LDISTIS OF ONOSTUS 77 P P SP T D F G H Figure 2. Scanning electron micrographs of Onomastus complexipalpis from South Kalimantan, right male palp (RMNH)., H, prolateral view., F, ventral view., D,, G, retrolateral view. bbreviations:, conductor;, embolus;, median apophysis; P, patella apophysis; SP, spur mesal branch of ; T, tegular apophysis. Scale bars = 20 mm (F, G, H), 30 mm (), 00 mm (,,, D).

8 78 S. P. NJMIN the South sia clade (S clade) which includes species from Sri Lanka and India and the South-ast sian clade (S clade), which includes the remaining diversity. DISUSSION Onomastus is a relatively small genus of jumping spiders mostly known from Sri Lanka. This study adds four more species from Sri Lanka, India, and South-ast sia. Noteworthy is the micro endemism of species of the S clade. In contrast, species of the S clade seem to be widespread. This study indicates that Onomastus of Sri Lanka and India may have originated from a common ancestor and subsequently evolved independently of each other. This is not unusual and has been reported for several faunal groups from Sri Lanka (ossuyt et al., 2004). Species of the S clade are spot endemics found mostly in higher elevation habitats. They may be highly in danger of extinction because of habitat loss and climate change. Wanless (980b) considered Onomastus distinct from any other Lyssomaninae, although no clear arguments were provided. However, this study suggests that it is a genus within a monophyletic Lyssomaninae (node ), which is supported by two unambiguous synapomorphies (22-, 26-). The inclusion of Onomastus within Lyssomaninae is in agreement with results of Wijesinghe (997). However, although the monophyly of Lyssomaninae is widely accepted (Wijesinghe, 997; Maddison & Hedin, 2003; Maddison & Needham, 2006), further assessments with the addition of more outgroup taxa are considered necessary. This study suggests that Onomastus is sister to the New World genus Lyssomanes (Fig. ). Interestingly Onomastus and Lyssomanes occupy similar ecological niches. However, this result is in contrast to that of Wijesinghe (997) who suggested that Onomastus is sister to rest of the lyssomanines. Together, Onomastus and Lyssomanes come out as sister to a grouping of three Old World genera, Pandisus, semonea, and Goleba informally termed here as the Pandisus clade (node 2), which is essentially group lll proposed by Wanless (980a). The monophyly of the Pandisus clade is supported by three unambiguous synapomorphies (-, 2-0, 24-). This study sheds more light on the phylogenetic position of Goleba, which was unclear in Maddison & Needham (2006). They did not include Onomastus, Pandisus, or semonea in their molecular study, which might have affected placement of Goleba. However, these hypotheses of relationships need to be further tested as the present cladistic analysis was designed to test the monophyly and the internal structure of Onomastus and not that of Lyssomaninae. The tracheal system has been widely used in the classification of spiders (Hormiga, 994 and references). The Lyssomaninae have been characterized as a group that possess a tracheal system that terminates in the opisthosoma (Wanless, 980a and references) as is the case in Hispo (Wanless, 98). Onomastus seem to be an exception (Fig. 27D, ). However, careful preparation of specimens seems to be critical to observing the fine tracheae that enter the prosoma. SYSTTIS ONOSTUS SIMON, 900 Type species: Onomastus nigricauda by original designation. D O Figure 3. Onomastus complexipalpis from South Kalimantan (RMNH)., left male palp (MNH )., vulva, ventral view. bbreviations:, conductor; D, copulatory duct; O, copulatory opening;, embolus; FD, fertilization duct;, median apophysis; P, patella apophysis. Scale bars = 0. (), 0.5 (). FD P

9 RVISION ND LDISTIS OF ONOSTUS 79 KY TO L ONOSTUS (O. SIMONI IS XLUDD FOR LK OF TRIL) Male palp with T2 (Figs 4, 7); endemic to India...2 Male palps without T () T2 with upward hook, relatively longer (Fig. 7)...Onomastus patellaris T2 otherwise, T and T2 shorter (Fig. 4)...Onomastus indra sp. nov. 3() Male palps with a broad, in some cases, disk-shaped (Figs 2, 6, 7), female epigynum with folds, vulva with clearly defined D (3, 5D)...7 Male palp with filiform, female epigynum without folds, vulva without D, all species endemic to Sri Lanka...4 4(3) deeply bifurcate, prolateral section with pleats (Figs 27, 28, )...Onomastus rattotensis sp. nov. without pleats...5 5(4) less strongly bifurcated (Fig. 0)...Onomastus nigricauda tip bulb like (Figs 20D, 23)...6 6(5) Tip of pointed, spermatheca oval (Figs 23, 24, D...Onomastus quinquenotatus Tip of stout, spermatheca relatively round (Figs 20D, 2, )...Onomastus pethiyagodai sp. nov. 7(3) broad, partly covers the tegulum, conductor with SP (Figs 2G, 7D)...8 filiform resting between and tegulum...9 8(7) large, covers most of the tegulum, tapering with a bifurcated tip (Figs 2, 3)......Onomastus complexipalpis smaller, stout (Fig. 7, D)...Onomastus kanoi 9(7) chisel-shaped, D absent (Figs 4, 6D)...Onomastus nigrimaculatus scoop like, D -shaped (Figs 5D, 6D)...Onomastus kaharian sp. nov. KY TO FL ONOSTUS (FLS OF O. KNOI RIN UNKNOWN) D absent or very short, endemic to Sri Lanka and India...5 D clearly visible, epigynal folds present...2 2() D characteristically long, coiled (Fig. 3)...Onomastus complexipalpis D relatively shorter or absent, epigynal folds present (Fig. 5D )...3 3(2) D absent, S round (Fig. 6 )...Onomastus nigrimaculatus D present, epigynal folds present...4 4(3) D inverted -shaped (Fig. 5D, )...Onomastus kaharian sp. nov. D inverted L -shaped, running partly parallel to each other...onomastus simoni 5() S round, endemic to South India...Onomastus patellaris, Onomastus indra sp. nov. S elongated endemic to Sri Lanka...Onomastus nigricauda, Onomastus pethiyagodai sp. nov., Onomastus quinquenotatus, Onomastus rattotensis sp. nov. Remarks: Listed as O. nigricaudus in Platnick (2009). However, nigri-cauda is a substantive and thus needs no gender adjustment. Diagnosis: Onomastus is separated from all other Salticidae by the presence of T ( y in Wanless, 980b) in the male palp (6-). The following additional characters support the monophyly of Onomastus and may serve to separate it: loss of the RT (7-), presence of a hyaline conductor (9-), the dorsal origin of the embolus (5-). Further, Onomastus is separated from all other Salticidae except for Lyssomanes and hinoscopus by the presence of a conductor (8-) and the lack of D (9-0; gained in O. kaharian, O. kanoi, and O. complexipalpis). The lack of a clearly defined fovea might also serve to characterize Onomastus. See also Wanless (980b) for a detailed diagnosis. Description: Male palp: complex and species-specific, suggesting that they have undergone rapid divergent evolution. Femur lacks any form of modification such as apophysis or furrows (Figs 3, 26). Patella with a retrolateral apophysis (Figs 3, 4, 6F). Tibia with a reduced retrolateral apophysis, hardly visible in most cases. ymbium distally tapering ( finger-like extension in Wanless, 980b), proximal promargin sclerotized and fringed with setae (Figs 2H,, 5, 28G). mbolus broad based (P in Fig. 9), tapering, slender, moderate to long in length, originates from the dorsal parts of the tegulum within the alveolus (Figs 5, 9). It may be used in some species as a mating plug (Fig. F, G). Tegulum large, makes up to half of the copulatory bulb. Distal tegular end tapers to an apophysis unique to Onomastus (T; Y in Wanless, 980b). The prolateral tegular margin of O. patellaris and O. indra holds an additional apophysis

10 720 S. P. NJMIN T2 T D P T3 P Figure 4. Onomastus indra sp. nov. (S)., left male palp, ventral view., ditto, retrolateral view., epigynum, ventral view. D, vulva, ventral view., ditto, dorsal view. bbreviations:, conductor; O, copulatory opening; Y, cymbium;, embolus; FD, fertilization duct;, median apophysis; P, mesal branch of ; P, patella apophysis; S, spermatheca; T, tibia; T, tegular apophysis ; T2, tegular apophysis 2; T3, tegular apophysis 3. Scale bars = 0.05 (), 0. (, D), 0.2 (, ). O T2 FD S P T Y unique to them [T2; X in Wanless (980b)]. third tegular apophysis is present in some species (Figs 4, 0, 9, 27), but absent or reduced in others. The apical half of the copulatory bulb constitutes the and conductor embolus complex. The conductor originates from in between the tegulum, which is grooved on the peripheral margin to form an embolic guide (e.g. in Wanless, 980b). Its tip is species-specifically modified (spur in Wanless, 980b). xcept for O. rattotensis, which has three distal branches, the is well sclerotized and distally bifurcate. The distal branches are species-specifically modified. In the South sian species the base of the possesses an apophysis (P) that serves as a rest for the apical

11 RVISION ND LDISTIS OF ONOSTUS 72 F portion of the embolus conductor complex (Figs 4, D, 9). pigynum: simple to absent. The O leads directly to the spermathecae. D lacking in most species. Spermathecae are large and well sclerotized. See also Wanless (980b) for a detailed description. Distribution: Oriental region, from Sri Lanka to Japan. However, 55% of the species are endemic to the Sri Lanka Western Ghats biodiversity hotspot. O Figure 5. Onomastus kaharian sp. nov. from entral Kalimantan (RMNH)., left male palp, ventral view., ditto, retrolateral view., epigynum, ventral view. D, vulva, ventral view., ditto, dorsal view. bbreviations:, conductor; O, copulatory opening;, embolus; F, epigynal folds; FD, fertilization duct;, median apophysis. Scale bars = 0. ( ), 0.2 (, ). D F The Sri Lankan endemics are restricted to small patches of rain forest and are highly endangered as a result of habitat loss. omposition: leven species: Onomastus complexipalpis Wanless, 980, O. indra sp. nov., O. kaharian sp. nov., O. kanoi Ono, 995, O. nigricauda Simon, 900, Onomastus nigrimaculatus Zhang & Li, 2005, O. patellaris Simon, 900, O. pethiyagodai sp. nov., O. FD

12 722 S. P. NJMIN T P D F quinquenotatus Simon, 900, O. rattotensis sp. nov., and O. simoni Zabka, 985. ONOSTUS OMPLXIPLPIS WNLSS, 980 (FIGS 2 H, 3 ) Holotype: from Indonesia: orneo, Santan, Kalimantan, 03.vii.976. Leg. R. Thompson, MNH , examined. PT Figure 6. Onomastus kaharian sp. nov. from entral Kalimantan (RMNH)., left male palp, prolateral view., D, ditto, ventral view.,, F, Ditto, retrolateral view. bbreviations:, conductor;, embolus;, median apophysis; P, patella apophysis; PT, patella; T, tibia. Scale bars = 20 mm (D, ), 30 mm (,,, F). Diagnosis: Onomastus complexipalpis is readily distinguished from all known species of Onomastus by the large palpal conductor in the males and the long coiled copulatory ducts in the females (Figs 2 H, 3 ). Male from South Kalimantan: Total length: 3.9 mm; prosoma length:.8, width:.3. Leg I: femur.2, patella 0.5, tibia.3, metatarsus.0, tarsus 0.5.

13 RVISION ND LDISTIS OF ONOSTUS 723 SP P PT F Prosoma oval, almost as wide as long. Opisthosoma oval, longer than wide. ll legs laterally without dark markings. ll eyes except M surrounded by dark rings. helicerae with pro- and retromarginal teeth, number not examined. Leg formula 432. Leg spination: leg : metatarsus D -0-0, V 2-2-2; tibia D --0, V 2-2-2; femur D --2, P 0-0-, R Palps as in Figures 2 H and 3. detailed description of the male is provided by Wanless (980b). T T Figure 7. Onomastus kanoi from Okinawa Prefecture (USNM).,, left male palp, ventral view., ditto, prolateral view., D, ditto, retrolateral view. F, profile of P, retrolateral view. bbreviations:, conductor;, median apophysis; P, patella apophysis; PT, patella; SP, spur mesal branch of ; T, tibia; T, tegular apophysis. Scale bars = 00 mm. SP D Female from South Kalimantan: Total length: 3.3; prosoma length:., width: 0.9. Leg : femur.5, patella 0.5, tibia.4, metatarsus.0 tarsus 0.4. Morphology as above, except for the lighter coloured prosoma and opisthosoma. Leg formula 432. Leg spination: leg : metatarsus V 2--2; tibia V 2-2-3; femur D -0-, P 0-0-, R pigynum and vulva as in Figure 3. Other material examined: Indonesia, entral Kalimantan, Kaharian, 20 2 S, Swampy primary forest, in leaf litter, 2 6.ix.985. Leg. Suharto Djojosudharmo., ; South Kalimantan, 40 km north-west of Palangkaraya, secondary forest,, 2 (right palp on a sputter-coated SM stub). Deposited in RMNH. T

14 724 S. P. NJMIN Figure 8. Photographs of live Onomastus nigricauda from odinagala, Sri Lanka (MHNG).,, male., D, female. ONOSTUS INDR SP. NOV. (FIG. 4 ) Holotype: from India, Kodaikanal, Grade, 600 m, 30.iii.962, leg.. S. Ross and D. Q. avagnaro. Deposited in S. Paratypes: 2 from the same locality as the holotype. Label data as above. Deposited in S. tymology: Indra is the deity of war in the Hindu religion. I use the species epithet as a noun in apposition. Diagnosis: Separated from all other species except for O. patellaris by the occurrence of T2 in males. Separated from O. patellaris by the shorter, stouter T and T2 (Fig. 4, ); further, the end of T2 is in the shape of an upward hook in O. patellaris (Fig. 7). oval, longer than wide, lighter in colour. Legs pale yellow, laterally without dark markings; might be a result of preservation. ll eyes except M surrounded by dark rings. helicerae with pro- and retromarginal teeth, number not examined. Leg formula 432. Leg spination: leg : metatarsus V 2--2; tibia V 2-2-3; femur D 0-0-, P Palps as in Figure 4 and. Female paratype: Total length: 4.4; prosoma length: 2.0, width:.3. Leg : femur.6, patella 0.6, tibia.4, metatarsus.2, tarsus 0.6. Morphology as above, except for the lighter coloured prosoma and opisthosoma and leg spination given below. Leg spination: leg : metatarsus V 2--2; tibia V 3-2-2; femur D --. pigynum and vulva as in Figure 4. Other material examined: None. D Male holotype: Total length: 3.8; prosoma length: 2.0, width:.3. Leg I: femur.6, patella 0.3, tibia 0.8, metatarsus 0.7, tarsus 0.3. ll spiders are whitish to pale yellow in alcohol, no markings. Prosoma oval, almost as wide as long, laterals darker. Opisthosoma ONOSTUS KHRIN SP. NOV. (FIGS 5, 6 F) Holotype: from Indonesia, entral Kalimantan, Kaharian, 20 2 S, Swampy primary forest,

15 RVISION ND LDISTIS OF ONOSTUS 725 H ST T P T3 P T PT Figure 9. Onomastus nigricauda from odinagala (MHNG)., expanded left male palp, prolateral view., ditto, retrolateral view. bbreviations: H, basal haematodocha;, conductor;, embolus; P, embolic base;, median apophysis; P, patella apophysis; PT, patella; ST, subtegulum; T, tibia; T, tegular apophysis ; T3, tegular apophysis 3. Scale bar = 0.2 mm.

16 726 S. P. NJMIN Y O S T3 P T FD D Figure 0. Onomastus nigricauda., male lectotype (MNHN 20404/-3), right palp, ventral view., female (found in the same vial as the lectotype)., female from odinagala, epigynum, ventral view. D, ditto, vulva, ventral view., ditto, dorsal view. bbreviations:, conductor; O, copulatory opening; Y, cymbium; FD, fertilization duct;, median apophysis; P, patella apophysis; S, spermatheca; T, tegular apophysis ; T3, tegular apophysis 3. Scale bars = 0. mm (), 0.2 mm (,, D, ).

17 RVISION ND LDISTIS OF ONOSTUS 727 D P O MP F G Figure. Scanning electron micrographs of Onomastus nigricauda from odinagala (MHNG)., left male palp, prolateral view., D, ditto, ventral view.,, ditto, retrolateral view. F, G, epigynum, ventral view. Note the mating plug in F and G. bbreviations:, conductor; O, copulatory opening;, median apophysis; P, mesal branch of ; MP, mate plug. Scale bars = 0 mm (G), 00 mm ( F).

18 728 S. P. NJMIN Figure 2. Scanning electron micrographs of Onomastus nigricauda from odinagala (MHNG)., profile of., ditto, detail., profile of T. Scale bars = 5 mm (), 0 mm (, ). in leaf litter, 2 6.ix.985. Leg. Suharto Djojosudharmo. Deposited in RMNH. Paratype: from Indonesia, locality and label data as above. Deposited in RMNH. tymology: The species name is a noun in apposition, taken from the type locality. Diagnosis: Males are readily separated from O. patellaris and O. indra by the absence of T2. Separated from O. rattotensis, O. nigricauda, O. quinquenotatus, and O. pethiyagodai by the presences of a broad conductor (Fig. 6D). Separated from O. complexipalpis, O. kanoi, and O. nigrimaculatus by the scoop-like and shape of (Fig. 6D). Females could be separated from all other Onomastus by the presence of semicircular epigynal fold and -shaped D (Fig. 5D, ). Remarks: Onomastus simoni appears to be closely related to O. kaharian. They may be separated from each other by the parallel course of D in O. simoni. Male holotype: Total length: 3.4; prosoma length:.5, width:.0. Leg I: femur.6, patella 0.5, tibia.7, metatarsus., tarsus 0.5. ll spiders are whitish to pale yellow in alcohol, no markings. Prosoma oval, almost as wide as long, laterals darker. Opisthosoma oval, longer than wide, lighter in colour. Legs pale yellow, laterally without dark markings; might be a result of preservation. ll eyes except M surrounded by dark rings. helicerae with pro- and retromarginal teeth, number not examined. Leg formula 432. Leg spination: leg : metatarsus V 2--2; tibia V 2-2-3; femur D 0-0-, P Palp as in Figures 5, and 6 F. Female paratype: Total length: 3.5; prosoma length:.4, width:.0. Leg : femur.4, patella 0.4, tibia.5, metatarsus., tarsus 0.4. Morphology as above, except for the lighter coloured prosoma, opisthosoma and leg spination given below. Leg spination: leg : metatarsus V 2--2; tibia V 3-2-2; femur D --. pigynum and vulva as in Figure 5. Distribution: Thailand and Indonesia. Other material examined: Indonesia: entral Kalimantan, Kaharian, 20 2 S, Swampy primary forest, in leaf litter, 2 6.ix.85, 2 (a single left palp on a sputter-coated SM stub), 5, leg. Suharto Djojosudharmo, deposited in RMNH. Thailand: Nakhon Si Thammarat Prov. Khao Luang NP, N, , 355 m, 0 2.x.2003 (TOL expedition 2003),, deposited in USNM. ONOSTUS KNOI ONO, 995 (FIG. 7 ) Diagnosis: Onomastus kanoi is readily distinguished from all known species of Onomastus except for O. complexipalpis by the large disk-shaped palpal conductor in the males (Fig. 7 D). The conductor of O. kanoi is relatively smaller than that of O. complexi-

19 RVISION ND LDISTIS OF ONOSTUS 729 palpis (Fig. 7D). The female of O. kanoi is unknown. However, I predict that the D of O. kanoi females would be longer than that of all other Onomastus, but shorter than that of O. complexipalpis. Male: Total length: 5.2; prosoma length: 2.0, width:.5. Leg I: femur 2.0, patella 0.6, tibia 2.0, metatarsus 2.0, tarsus 0.6. For a detailed description see Ono (995). Female: Unknown. Figure 3. Onomastus nigricauda from Morningside (MHNG)., left male palp, ventral view., ditto, retrolateral view. Scale bars = 0.2 mm. Distribution: ndemic to Okinawa Island of Japan. Material examined: Japan, Okinawa Prefecture, Okinawajima Island, Kijoka. 0.iv.997. ; Japan, Okinawa Prefecture, Okinawajima Island, Zutsun. 0.iv.2000,. ll leg. kio Tanikawa. ONOSTUS NIGRIUD SIMON, 900 (FIGS 8 D, 9, 0, G, 2, 3 ) Onomastus nigricauda Simon, 900: 29. lectotype and 2 from Sri Lanka, Galle, in MNHN (20404/ -3), examined. Onomastus nigricauda Simon, 90: 400, fig Onomastus nigricauda Wanless, 980: 8, fig. G. Initial description of the female and designation of types. Diagnosis: Males are readily separated from O. patellaris and O. indra by the absence of T2. Separated from O. complexipalpis, O. kanoi, O. kaharian, and O. nigrimaculatus by the filiform conductor. Separated from O. pethiyagodai and O. quinquenotatus by the shape of the. Onomastus nigricauda seems to be most closely related to O. rattotensis, but may be separated by the large partly pleated in O. rattotensis (Figs 27, 28 ). Male (from odinagala): Total length: 3.0; prosoma length:.5, width:.. Leg I: femur.4, patella 0.4, tibia.5, metatarsus 0.9, tarsus 0.5. The spiders are green in nature. They turn whitish to pale yellow in alcohol. Prosoma oval, longer than wide. Opisthosoma longer than wide, lighter in colour. Leg I yellowish brown, leg II IV light yellow. Legs are laterally without dark markings. ll eyes except M surrounded by dark rings. helicerae are

20 730 S. P. NJMIN D F T P Figure 4. Scanning electron micrographs of Onomastus nigrimaculatus from Thailand (RMNH)., D, left male palp, prolateral view.,, F, ditto, ventral view., ditto, retrolateral view. bbreviations:, conductor;, median apophysis; P, patella apophysis; T, tegular apophysis. Scale bars = 0 mm (), 30 mm (D), 00 mm (, F).

21 RVISION ND LDISTIS OF ONOSTUS 73 Figure 5. Scanning electron micrographs of Onomastus nigrimaculatus from Thailand (RMNH)., profile of and, retrolateral view., ditto, retrolateral-apical view., serrated setae of the cymbium, retrolateral view. bbreviations:, conductor;, embolus;, median apophysis. Scale bars = 0 mm. yellow, with seven retromarginal and three promarginal teeth. Leg formula 432. Leg spination, metatarsus D -0-0, V 2-2-2; tibia D 0-0-, P -0-0, V 4-4-2, R -0-0; patella D 0--0; femur D 2--0, P 0-0-, R Palp: (Figs 9, 0,, 2, 3 ). Tegulum as in Figure 9, projecting outwards, path of sperm duct with two loops as in Figure 9 and, embolus as in Figure 9. detailed description of the male lectotype is also provided by Wanless (980b). Female (from odinagala): Total length: 4.0; prosoma length:.5, width:.2. Leg : femur.4, patella 0.5, tibia.5, metatarsus., tarsus 0.5. Morphology as above, except for the lighter coloured prosoma and opisthosoma. Leg spination, metatarsus V 4-2-2, P -0-0, R -0-0; tibia D --0, V 4-4-2; patella P -0-0, R -0-0; femur D --, P 0-0-, R pigynum and vulva as in Figures 0 and F G. detailed description of the female is provided by Wanless (980b). Variation: The male specimen from Morningside has a bifurcate in which both branches taper towards the end. Live male spiders of O. nigricauda from odinagala tend to also have a much darker palp than that of Morningside. Distribution: ndemic to Sri Lanka; odinagala forest reserve is a remaining fragment of the once vast rain forest that covered the south-west of the island. It is not far from the type locality, Galle. Other material examined: SRI LNK: Western province, Kalutara district, Horane, odinagala, 9.vii.996, 2, leg. Suresh. P. enjamin. Same locality, 0.ii.2007, 5 6, leg. Suresh. P. enjamin and Ziyard Jaleel. Sabaragamuwa province, Ratnapura District, Morningside section. In primary forest, 23.ii.2007,, leg. Suresh P. enjamin and Ziyard Jaleel. ll specimens deposited in MHNG. ONOSTUS NIGRIULTUS ZHNG & LI, 2005 (FIGS 4 F, 5, 6 ) Holotype: Male from hina, Mingfeng Valley, Jianfengling National Park, Hainan Province. Deposited in the Raffles Museum of iodiversity Research (ZR- R-494). s this species was adequately illustrated in its original description, I have not examined any type material. Diagnosis: Males of O. nigrimaculatus are readily separated from those of O. patellaris and O. indra by the absence of T2. Separated from O. rattotensis, O. nigricauda, O. quinquenotatus, and O. pethiyagodai by the presence of a broad conductor and chiselshaped. Separated from O. complexipalpis, O. kanoi, and O. kaharian by the chisel-shaped and details of. Female O. nigrimaculatus could be separated from all other Onomastus by the combined presence of -shaped epigynal fold and lack of D.

22 732 S. P. NJMIN Male: Total length: 4.0; prosoma length:.8, width:.3. Leg I: femur.4, patella 0.5, tibia.7, metatarsus.4, tarsus 0.8. Prosoma oval, almost as wide as long, laterals with a faint black border. Opisthosoma oval, longer than wide, lighter in colour, four to six dark black spots on dorsum. Legs light yellow, leg laterally with dark black spots, legs II IV do not have similar spots. ll eyes except M surrounded by dark rings. helicerae with pro- and retromarginal teeth, number not examined. Leg formula 432. Leg D Figure 6. Onomastus nigrimaculatus from Thailand (RMNH)., left male palp, ventral view., ditto, retrolateral view., female from the same locality, epigynum, ventral view. D, ditto, vulva, ventral view., ditto, dorsal view. Scale bars = 0.2 mm ( ), 0.5 mm (, ). spination: leg : metatarsus D -0-0, V 2-2-2; tibia D -0-, V 2-4-2, P -0-0, R -0-0; patella D -0-0, V -0-0; femur D --2, P 0-0-, R Palp as in Figures 4 F, 5, 6. Female: Total length: 5.0; prosoma length: 2.2, width:.5. Leg : femur 2.0, patella 0.7, tibia 2.4, metatarsus.5, tarsus 0.6. Morphology as above. Leg formula 432. Leg spination: leg : metatarsus V 2-2-2, P

23 RVISION ND LDISTIS OF ONOSTUS 733 T2 T PT -0-0, R -0-0; tibia D -0-0, V 2-2-2; Patella D -0-0, P 0--0, R pigynum and vulva as in Figure 6. Distribution: Hainan province, hina (Zhang & Li, 2005) and primary rain forest of the Khao Yai National Park, Thailand. Other material examined: Thailand, Khao Yai N.P, 3 7.iii.986, leg. L and PR Deeleman, 4. Same locality as above, 23.x.985, leg. PR Deeleman, 4. Same locality, 5.x.987, leg. L and PR Deeleman,. Same locality, 9 24.x.985, leg. L and PR Deelema, 4. ll specimens deposited in RMNH. ONOSTUS PTLLRIS SIMON, 900 (FIG. 7 ) Onomastus patellaris Simon, 900: 29. P Figure 7. Onomastus patellaris from Kodaikanal, India (MNHN 4858)., left male palp, ventral view.,, ventral view., epigynum, ventral view. bbreviations:, conductor;, embolus;, median apophysis; P, patella apophysis; PT, patella; T, tegular apophysis ; T2, tegular apophysis 2. Scale bars = 0. mm (), 0.2 mm () 0.4 mm (). Lectotype: Single male from India, Kodaikanal, MNHN 4858, examined. Diagnosis: Separated from all other species of Onomastus except O. indra by the presence of T2 in male palps (Fig. 7). T and T2 in O. patellaris are filiform. Further, the end of T2 is in the shape of an upward hook in O. patellaris (Fig. 7). Description: See Wanless (980b). Distribution: India, Madras, Kodaikanal. Probably endemic to India. Other material examined: Four from India, Trichinopoly, leg. R. P. Malat, MNHN 4898.

24 734 S. P. NJMIN Figure 8. Photographs of live Onomastus pethiyagodai sp. nov. from grapathana, Sri Lanka (MHNG)., male. D F, female. ONOSTUS PTHIYGODI SP. NOV. (FIGS 8 F, 9 F, 20, 2 D, 22 ) Holotype:, Sri Lanka: entral Province, Nuwara liya District, grapathana, grabopath forest, 00 m, 08.iii Deposited in MHNG. Paratype: 3, label data as above. Deposited in MHNG. Diagnosis: Males of O. pethiyagodai are readily separated from that of O. patellaris and O. indra by the absence of T2. Separated from O. complexipalpis, O. kanoi, O. kaharian, and O. nigrimaculatus by the presence of a filiform conductor. O. pethiyagodai is most closely related to O. quinquenotatus, and may be separated by the shape of the conductor and (Figs 9, 20D, 22). Females are difficult to diagnose, but may be separated by details of internal genitalia. D F

25 RVISION ND LDISTIS OF ONOSTUS 735 P T3 T D F P Figure 9. Scanning electron micrographs of Onomastus pethiyagodai sp. nov. from grapathana, Sri Lanka (MHNG)., D, left male palp, prolateral view.,, ditto, ventral view., F, ditto, retrolateral view. bbreviations:, conductor;, embolus;, median apophysis; P, mesal branch of ; P, patella apophysis; T, tegular apophysis ; T3, tegular apophysis 3. Scale bars = 20 mm (, D,, F), 30 mm (, ).

26 736 S. P. NJMIN D Figure 20. Scanning electron micrographs of Onomastus pethiyagodai sp. nov. from grapathana, Sri Lanka (MHNG).,, prolateral view., P, retrolateral view., ditto, ventral view. D,, ventral view., and prolateral view. Scale bars = 0 mm (,, D), 20 mm (, ). Male holotype: Total length: 2.9; prosoma length:.5, width:.. Leg I: femur.4, patella 0.2, tibia.2, metatarsus.0, tarsus 0.4. The spiders are green in nature. They turn whitish to pale yellow in alcohol. Prosoma oval, longer than wide. Opisthosoma longer than wide, lighter in colour. Leg I yellowish brown, leg II IV light yellow. Legs are laterally marked with dark black blobs. ll eyes except M surrounded by dark rings. helicerae with pro- and retromarginal teeth, number not examined. Leg formula 432. Leg spination: metatarsus V 4-2-2; tibia P -0-0, V 2-4-2, R -0-0; patella V 2-0-0; femur D 0--, P 0-0-, R Palp as in Figures 9 F, 20, 22. Female paratype: Total length: 2.9; prosoma length:.2, width:.. Leg : femur.0, patella 0.3, tibia.0, metatarsus 0.8, tarsus 0.4. Morphology similar to the male except for the following. Prosoma and opisthosoma lighter in colour. Legs with lateral black blobs. Leg spination: metatarsus V 2-2-2; tibia P -0-0, V 2-4-4, R -0-0; patella P -0-0, R -0-0; femur D 0-2-, P 0-0-, R pigynum and vulva as in Figure 2 D. Distribution: ndemic to the central highlands of Sri Lanka. Known from grapathana, grabopath forest and Horton Plains National Park. Other material examined: SRI LNK: entral Province, Nuwara liya District, grapathana, grabopath forest, 00 m, 07.iii.2000, 2 ; 30.vi.2003, 7 7 (a single left palp on a sputter-coated SM stub), all leg. Suresh P. enjamin. Same locality, 08.iii.2000, leg. Sudath Nanayakkara. Same local-

27 RVISION ND LDISTIS OF ONOSTUS 737 O D D Figure 2. Onomastus pethiyagodai sp. nov. from grapathana, Sri Lanka., D, epigynum, ventral view., vulva, ventral view., ditto, dorsal view. bbreviations: D, copulatory duct; O, copulatory opening; FD, fertilization duct. Scale bars = 0.2 mm. ity, 8 2.ii.2007, 3 2, leg. Suresh P. enjamin and Ziyard Jaleel. entral Province, Nuwara liya District, Horton Plains National Park, c m, 20 2.ii.2007, 5 5, leg. Suresh P. enjamin and Ziyard Jaleel. Deposited in MHNG. ONOSTUS QUINQUNOTTUS SIMON, 900 (FIGS 23 D, 24 D, 25 H) Onomastus quinquenotatus Simon, 900: from Sri Lanka in MNHN 2077, examined. O. quinquenotatus Wanless, 980: 83, fig. 2a e. Designation of lectotype and paralectotype. Diagnosis: Males are readily separated from O. patellaris and O. indra by the absence of T2. Separated from O. complexipalpis, O. kanoi, O. kaharian, and O. nigrimaculatus by the filiform conductor. Separated from O. pethiyagodai and O. nigricauda by the shape of the (Fig. 23). Onomastus quinquenotatus is most closely related to O. pethiyagodai and they may be separated from each other by the shape of the FD conductor and (Fig. 23 D). Females are difficult to diagnose, but may be separated by details of the internal genitalia. Male from Kandaela: Total length: 3.; prosoma length:.6, width:.3. Legs I: femur.6, patella 0.5, tibia.5, metatarsus.3, tarsus 0.6. Prosoma round to oval, longer than wide, pale amber to light yellow. yes with black surrounds except for M. Opisthosoma oval, longer than wide, lighter in colour, no markings. Living specimens green in colour. Leg posterior laterally black along its length, all other legs uniformly yellow. helicerae light yellow, with seven retromarginal and three promarginal teeth. Spider ventrally light yellow. Leg formula 432. Leg spination: metatarsus V 4-2-2; tibia D 2-0-0, V 2-4-4; patella D -0-0; femur D Palp as in Figures 23 D, 25 H. Female from Kandaela: Total length: 2.2; prosoma length:.8, width:.3. Leg : femur.4, patella 0.5,

28 738 S. P. NJMIN Figure 22. Onomastus pethiyagodai sp. nov. from grapathana, Sri Lanka (MHNG)., left male palp, ventral view., ditto, retrolateral view. Scale bar = 0.2 mm. T T3 P D Figure 23. Onomastus quinquenotatus, male from Hakgala, Sri Lanka (MHNG)., left male palp, ventral view., profile of., profile of. D, left male palp, retrolateral view. bbreviations:, conductor;, embolus;, median apophysis; P, patella apophysis; T, tegular apophysis ; T3, tegular apophysis 3. Scale bars = 0. mm (, ), 0.2 mm (, D).

29 RVISION ND LDISTIS OF ONOSTUS 739 O Figure 24. Onomastus quinquenotatus., female lectotype (MNHN 2077), epigynum, ventral view., female from Kandaela, Sri Lanka (USNM), epigynum, ventral view., vulva, ventral view. D, ditto, ventral view. bbreviations: O, copulatory opening; FD, fertilization duct. Scale bars = 0.2 mm. tibia.6, metatarsus.3, tarsus 0.5. Morphology similar to the male except for the following: prosoma and opisthosoma lighter in colour; legs with lateral black blobs. Leg spination: metatarsus V 4-2-2; tibia V 2-2-2; femur D pigynum and vulva as in Figure 24 D. Distribution: Known from two localities, Kandaela and Hakgala, in the central highlands of Sri Lanka. The type locality in the labels is given as Sri Lanka (Simon, 900). The exact locality of the lectotypes in Sri Lanka is unclear. Onomastus quinquenotatus is endemic to Sri Lanka. FD D ONOSTUS RTTOTNSIS SP. NOV. (FIGS 26 D, 27, 28 G) Holotype: from Sri Lanka. entral Province, Knuckles range, along Rattota-Ilukkumbura Road, 900 m, ix.2003, leg. Suresh P. enjamin. Deposited in MHNG. Paratype: from Sri Lanka. Data as above. Deposited in MHNG. tymology: djective, from Rattota, after the type locality. Other material examined: SRI LNK: entral province, north-east district, Kandaela reservoir, 5.6 miles south-west of Nuwara liya, 2000 m 0 2.ii.970,, leg. Davis and Rowe, deposited in USNM; Hakgala, Hakgala forest, 27.vii.996, 600 m,, leg. Suresh P. enjamin. Deposited in MHNG. Diagnosis: Males of O. rattotensis are readily separated from those of O. patellaris and O. indra by the absence of T2. Separated from O. complexipalpis, O. kanoi, O. kaharian, and O. nigrimaculatus by the filiform conductor. Separated from O. pethiyagodai and O. quinquenotatus by the shape of the. Separated from O. nigricauda by the presence of a

30 740 S. P. NJMIN D F G H T P T3 T Figure 25. Scanning electron micrographs of Onomastus quinquenotatus male from Hakgala, Sri Lanka (MHNG)., D, G, prolateral view.,, H, ventral view., F, retrolateral view. bbreviations:, conductor;, embolus;, median apophysis; P, patella apophysis; T, tegular apophysis ; T3, tegular apophysis 3. Scale bars = 0 mm (F, H), 30 mm (, G), 00 mm (). T3

31 RVISION ND LDISTIS OF ONOSTUS 74 D Figure 26. Onomastus rattotensis sp. nov. from Rattota, Sri Lanka (MHNG)., left male palp, ventral view., ditto, retrolateral view., epigynum, ventral view. D, vulva, ventral view., ditto, dorsal view. Scale bars = 0. mm (D, ), 0.2 mm ( ).

32 742 S. P. NJMIN D P T3 T Figure 27. Scanning electron micrographs of Onomastus rattotensis sp. nov. from Rattota, Sri Lanka (MHNG)., left male palp, retrolateral view., ditto, retrolateral view., ditto, prolateral view. D, female tracheal system, dorsal view., ditto, detail. bbreviations:, conductor;, embolus;, median apophysis; P, mesal branch of ; T, tegular apophysis ; T3, tegular apophysis 3. Scale bars = 00 mm. The arrow points to the tracheae, which enter the prosoma.

33 RVISION ND LDISTIS OF ONOSTUS 743 T P P T3 Y T3 D F G Figure 28. Scanning electron micrographs of Onomastus rattotensis sp. nov. from Rattota, Sri Lanka (MHNG)., left male palp, ventral view., and, ventral view., F, tip, ventral view. D, profile of T3, ventral view., profile of tip, ventral view. G, serrated setae of the cymbium. bbreviations:, conductor; Y, cymbium;, embolus;, median apophysis; P, mesal branch of ; P, patella apophysis; T, tegular apophysis ; T3, tegular apophysis 3. Scale bars = 0 mm (,, F, G), 20 mm (, D), 00 mm ().

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