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Zoological Journal of the Linnean Society, 2009, ••, ••–••. With 28 ﬁgures

1 Revision and cladistic analysis of the jumping spider

2 genus Onomastus (Araneae: Salticidae) 22 3 4 SURESH P. BENJAMIN* 5 6 Department of Entomology, National Museum of Natural History MRC 105, PO Box 37012, 7 Smithsonian Institution, Washington, DC 20013-7012, USA 8 9 Received 5 September 2008; accepted for publication 25 February 2009

11 The jumping spider genus Onomastus Simon, 1900 is revised. Four new species: Onomastus indra sp. nov., 12 Onomastus kaharian sp. nov., Onomastus pethiyagodai sp. nov., and Onomastus rattotensis sp. nov. are 13 described. Parsimony analysis of 26 morphological characters supported the monophyly of Onomastus. Lyssomanes 14 is sister to Onomastus. Onomastus separates into two clades: the widespread South-East Asia clade and the South 15 Asia clade. The South Asia clade is restricted to the Sri Lanka–Western Ghats biodiversity hotspot. Species of the 33 16 South Asia clade appear to be spot endemics, highly in danger of extinction because of habitat loss and climate 17 change. Male palps are complex and species-speciﬁc, suggesting rapid divergent evolution. © 2009 The Linnean 18 Society of London, Zoological Journal of the Linnean Society, 2009, ••, ••–••.

19 ADDITIONAL KEYWORDS: biodiversity – climate change – hotspots – India – South-East Asia – Sri Lanka.

21 INTRODUCTION Except for Onomastus complexipalpis sp. nov., female 45 genitalia are less complex and more similar in most 46 22 Our current understanding of the evolutionary rela- aspects. 47 23 tionships of jumping spiders, the largest spider Currently lyssomanines are defined by the presence 55 48 24 family, suggest at least four major lineages: the lys- of four rows of eyes and a tracheal system that is 66 49 25 somanines, the spartaeines, the Cocalodes group, and confined to the opisthosoma (Galiano, 1962; Wanless, 50 26 the much larger Salticoida clade (Wanless, 1980a, 1980a). Wanless (1980a) used the eye pattern to 51 27 1984; Maddison, 1996; Maddison & Hedin, 2003; define groupings of lyssomanine genera: Asemonea 52 28 Maddison & Needham, 2006). The three non- O. P.-Cambridge, 1869, Goleba Wanless, 1980, 53 29 Salticoida clades have recently been collectively Macopaeus Simon, 1900, and Pandisus Simon, 1900 54 30 termed ‘basal salticids’ (Maddison & Needham, 2006). was termed group III; group III was considered by 55 31 Jumping spiders of the genus Onomastus Simon, him to be related to group II, which consisted of the 56 32 1900 are lyssomanines (Wanless, 1980b). They are genera Lyssomanes Hentz, 1845 and Chinoscopus 57 33 small to medium sized spiders (2.0 to 5.2 mm total Simon, 1901. Onomastus was however, considered not 58 34 length) that live on foliage of tropical evergreen closely allied to either of them and was thus placed in 59 35 forests. They stalk for prey and do not build webs. group I (Wanless, 1980a). However, these hypotheses 60 36 They resemble other lyssomanines in coloration, have not been tested in a phylogenetic context. 61 37 general slender body shape, and by having the eyes The type species of Onomastus, Onomastus nigri- 62 38 arranged in four transverse rows (Wanless, 1980b). cauda Simon, 1900 is endemic to lowland evergreen 63 39 However, in contrast to Salticoida and other lyssoma- rainforests of western Sri Lanka. A further two 64 404 4 nines and spartaeines, Onomastus male palps are species, Onomastus quinquenotatus Simon, 1900 from 65 41 unusually complex and provide enough information to Sri Lanka and Onomastus patellaris Simon, 1900 66 42 clearly distinguish amongst closely related species. from the Western Ghats of southern India, were 67 43 described by Simon (1900). The first species outside 68 44 *E-mail: [email protected] the Indian subcontinent, O. complexipalpis was 69

2 S. P. BENJAMIN

1 described in a comprehensive treatment of the genus Shadab (1975). A complete synonymy of the species is 55 2 by Wanless (1980b). However, by 1980 only O. nigri- given in Platnick (2008). 56 3 cauda and O. patellaris were known by both sexes. 57 4 Recently several species from East Asia have been 5 described: Onomastus simoni Zabka, 1985 from ABBREVIATIONS 58 6 Vietnam (male unknown), Onomastus kanoi Ono, Institutional 59 7 1995 from Okinawa, Japan (female unknown), and BMNH, British Museum, Natural History, London; 60 8 Onomastus nigrimaculatus Zhang & Li from China CAS, California Academy of Sciences, San Francisco; 61 9 (Zabka, 1985; Ono, 1995; Zhang & Li, 2005). MHNG, Muséum d′Histoire Naturelle, Genève; 62 10 As part of an on-going survey of spider diversity in MNHN, Muséum National d′Histoire Naturelle, 63 11 Sri Lanka, a large number of Onomastus was col- Paris; RMNH, Naturalis, National Museum of 64 12 lected. They turned out to be of five species, two Natural History, Leiden; USNM, National Museum of 65 13 7 7 known and two new. All seem to be endemic and Natural History, Smithsonian Institution, Washing- 66 14 restricted to small patches of higher elevation rain ton, DC. 67 15 forest. Additionally, a collection of South-East Asian 68 16 Onomastus from the collections of Christa L. Morphological structures 69 17 Deeleman-Reinhold has been incorporated. A further AEB, anterior epigynal border; ALE, anterior lateral 70 18 undescribed species from the Western Ghats was also eye; AME, anterior median eye; AP, apical plates of 71 19 8 8 discovered in the collections of the CAS. Here, I epigynum; B, bulbus; BH, basal haematodocha; C, 72 20 provide comprehensive accounts of all Onomastus conductor; CD, copulatory duct; CO, copulatory 73 21 from Sri Lanka and the surrounding region with opening; CY, cymbium; E, embolus; EF, epigynal folds; 74 22 descriptions of new species and updated diagnosis. EP, embolic base; F, femur; FD, fertilization duct; MA, 75 23 Additionally, higher-level relationships of Salticidae median apophysis; MAP, mesal branch of MA; MP, 76 24 are discussed based on a cladistic analysis of Ono- mate plug; PA, patella apophysis; PLE, posterior 77 25 9 9 mastus plus exemplars of Wanless’s group I–III. lateral eye; PME, posterior median eye; PT, patella; 78 26 RTA, retrolateral tibial apophysis; S, spermatheca; 79 SP, spur mesal branch of C; ST, subtegulum; T, tibia; 80 27 MATERIAL AND METHODS TA1, tegular apophysis 1 (Figs 4A, 10A, 17A); TA2, 81 tegular apophysis 2 (Figs 4A, 17A); TA3, tegular apo- 82 28 Methodology and taxonomic descriptions follow the physis 3 (Figs 4A, 9B, 28D). 83 29 format of Benjamin (2004, 2006). Specimens used for 30 illustrations were placed on washed sand in 70% 84 31 ethanol and photographed using a dissecting micro- CLADISTIC ANALYSIS 11 85 32 scope (Leica MZAPO) with top illumination and a Outgroup choice 86 33 magnification of up to 50 ¥. Digital images were taken The primary aim of the cladistic analysis was to test 87 34 with a Nikon DXM1200F camera. Images were edited the monophyly and the internal structure of Onomas- 88 35 using an Auto-Montage software package. Either a tus. Unfortunately, relationships of basal salticids are 89 36 LEO 1430VP or an Amray 1810 scanning electron far from satisfactorily resolved, complicating the 90 37 microscope (SEM) was also used to study and photo- selection of the putative outgroup. The selection of 91 38 graph morphological features. Prior to SEM work outgroup taxa was based on Wijesinghe (1997), 92 39 specimens were prepared as mentioned in Benjamin Wanless (1980a), and Maddison & Needham (2006). 93 40 (2004) and mounted on to aluminium rivets with the Thus, I have included the genera Onomastus, Lysso- 94 41 help of conductive copper adhesive tape or wire. Such manes, Pandisus, Asemonea, and Goleba. Further, as 95 42 mounting, although rather intricate, enabled rotation suggested by Maddison & Needham (2006), I have 96 43 of specimens in the SEM chamber. Male palps were also included Hispo Simon, 1886, a possible sister to 97 44 cleared in methyl salicylate and mounted on tempo- Lyssomaninae. 98 45 rary slides to observe and illustrate their internal 46 structures. Left structures (e.g. palps, legs, etc.) are 99 47 depicted unless otherwise stated. Hairs and macrose- Taxa and characters scored 100 48 tae are usually not depicted in the final palp draw- The taxa and characters used served the main 101 49 ings. All measurements are given in millimetres and purpose of the study, which was to test the monophyly 102 50 were made with a compound microscope (Leica and the internal structure of Onomastus and not to 103 51 MZAPO) equipped with a 10 ¥ ocular and an ocular tackle higher-level relationships amongst salticids. As 104 52 10 10 micrometre scale. Abbreviations used in the text and in my previous work (Benjamin, 2004), I consider all 105 53 figures are given below. Leg spination is described character statements to be hypotheses of homology, 106 54 according to the system adopted by Platnick & unless putative homology of traits of two or more taxa 107

REVISION AND CLADISTICS OF ONOMASTUS 3

21 Hispo cingulata 1 Pandisus clade 4 84 Pandius sarae 1 12 24 1 2 3 1 0 1 23 25 Asemonea tenuipes 1 2 21 0 1 Goleba puella 1 1 22 26 1 Lyssomanes virides 1 1 South-east Asia clade Lyssomaniae 19 O. nigrimaculatus 0

8 10 20 O. kaharian 1 13 2 1 5 O. complexipalpis 1 95 11 1 4 7 9 15 16 O. kanoi 3 0 1 1 1 1 1 Onomastus O. pethiyagodai 81 14 18 19 O. quinquenotatus 1 1 0

6 10 11 17 O. patellaris

1 1 0 1 O. indra

13 O. nigricauda

1 O. rattotensis

South Asia clade

1 Figure 1. Preferred most parsimonious tree of Onomastus based on the character matrix presented in Table 1 2 (length = 32 steps, consistency index = 0.81, retention index = 0.89). Character state changes are mapped using Farris 3 optimization. Closed circles represent unambiguous character changes.

5 are rejected by the cladistic analysis (Fig. 1). All Ono- found in Asemonea and Pandisus (Fig. 26B; Wanless, 23 6 mastus species except for O. simoni are included in 1980b). 24 7 the phylogenetic analysis. Onomastus simoni is only 25 8 known from a single female specimen, additional 2. Position of FA: retrolateral/distal, 0; ventral/ 26 9 material was unavailable for study. All characters are median, 1. 27 10 scored directly from voucher specimens. However, as Refers to the position of the FA mentioned above. FA 28 11 a result of a lack of material in collections and to is more towards the ventral surface of the centre of 29 12 some extent because of the reluctance of curators to the femur in Goleba (Wanless, 1980a: figs 22c, d, 24b), 30 13 lend material for SEM work, outgroup taxa were whereas it is in a distal retrolateral position in Ase- 31 14 scored on the basis of published accounts. monea and Pandisus (Wanless, 1980a: figs 4e, 18c). 32 15 Wanless (1980a) described state 1 found in Goleba as 33 16 Male palp ‘ventral tubercle on the palpal femur’. 34 17 1. FA: absent/femora smooth, 0; present, 1. 35 18 FA is defined here as any structure projecting out- 3. Femoral groove: absent, 0; present, 1. 36 19 wards from the surface of the femur. Present in Ase- Femoral groove is defined as a cavity of the surface of 37 20 monea, Pandisus, and Goleba (Wanless, 1980a: the femur. It does not project outwards as is the case 38 21 figs 4e, 18c). In Onomastus femora are smooth and in the FA. The femoral groove is only present 39 22 unmodified, lacking any furrows or apophyses as in Asemonea (Wanless, 1980a: figs 10e, 13c). In 40

4 S. P. BENJAMIN

1 Table 1. Distribution of characters scored for ten Onomastus taxa and outgroups 2 3 01 4 5 1234567890123 6 7 Onomastus quinquenotatus 0–01001110110 8 Onomastus nigrimaculatus 0–01001112110 9 Onomastus nigricauda 0–01001110111 10 Onomastus patellaris 0–01011111010 11 Onomastus indra sp. nov. 0–01011111010 12 Onomastus pethiyagodai sp. nov. 0–01001110110 13 Onomastus rattotensis sp. nov. 0–01001110111 14 Onomastus complexipalpis 0–01101112111 15 Onomastus kanoi 0–01101112010 16 Onomastus kaharian sp. nov. 0–01001112110 17 Pandisus sarae* 10010000–––0– 18 Asemonea tenuipes* 1010–000–––0– 19 Goleba puella* 1100–000–––0– 20 Hispo cingulata* 0––0–000–––10 21 Lyssomanes viridis* 0––0–00100–10 22 23 20 24 25 4567890123456 26 27 O. quinquenotatus 1110100011001 28 O. nigrimaculatus 0110001011001 29 O. nigricauda 1110100011001 30 O. patellaris 1111100011001 31 O. indra sp. nov. 1111100011001 32 O. pethiyagodai sp. nov. 1110100011001 33 O. rattotensis sp. nov. 1110100011001 34 O. complexipalpis 0110011011001 35 O. kanoi 01100???11001 36 O. kaharian sp. nov. 0110011011001 37 P. sarae* –000010011101 38 A. tenuipes* –000010010111 39 G. puella* –000010110111 40 H. cingulata* 0000010101000 41 L. viridis* 0000010011001 42 43 Character states are scored 0 to 2, ? for unknown, – for inapplicable. Outgroup taxa are denoted with an asterisk. 44 45 Onomastus femora are smooth and unmodified, 5. PA shape: tapering, 0; blunt, 1. 59 46 lacking any furrows (Fig. 26B). PA is rather tapering with a smooth rounded tip in all 60 47 Onomastus, except O. complexipalpis and O. kanoi.In 61 48 4. PA: absent, 0; present, 1. O. complexipalpis and O. kanoi the PA broadens 62 49 Present in all Onomastus and Pandisus (Figs 9A, towards the end (Figs 3A–C, 3A, 7F; Ono, 1995). 63 50 10A, 14F, 17A; Wanless, 1980a; Wanless, 1980b). Ono- 64 51 mastus lack an RTA, instead the patella of the male 6. Relative size of patella: patella = tibia, 0; 65 52 palp is armed with an apophysis at the distal end that patella > tibia, 1. 66 53 probably functions as a replacement for the RTA. A The relatively elongated patella is unique to the two 67 54 PA is also present in Pandisus, as illustrated for Indian Onomastus, O. patellaris and Onomastus 68 55 Pandisus scalaris by Wanless (1980a: fig. 3). One indra (Figs 4A, B, 17A). Hence the origin of the spe- 69 56 would consider the PA to be homologous in both cific name ‘patellaris’, which was the first Onomastus 70 57 genera based on the criteria of Remane (1952). species with an elongated patella to be described from 71 58 However, it appears to be otherwise (Fig. 1). India; a second is described below. 72

REVISION AND CLADISTICS OF ONOMASTUS 5

1 7. RTA: present, 0; absent, 1. known species from Sri Lanka and India (Figs 4A, 56 2 A RTA is absent in Onomastus. As mentioned above, 11D, 19E, 27A). 57 3 the distal ends of the tibia of male palps of all Ono- 58 4 mastus are smooth. As a member of the RTA clade one 15. Origin of embolus: dorsal, 0; ventral, 1. 59 5 would expect to see a RTA in all Onomastus. However, The embolus originates from the tegulum within the 60 6 this is not the case and it is presumed lost in all alveolus in all Onomastus species (Fig. 9A, 15A, B; 61 7 species (Wanless, 1980b). Wanless, 1980b). The origin of the embolus is visible 62 8 when male palps are cleared in methyl salicylate. It is 63 9 8. Conductor: absent, 0; present, 1. also clearly visible in the expanded palp (Fig. 9A). 64 10 A very complex conductor is present in all known This character is diagnostic for Onomastus. 65 11 Onomastus species and is species-specific in size and 66 12 shape (Figs 2F, G, 5A, B, 9A; Wanless, 1980b). 16. TA1: absent, 0; present, 1. 67 13 Onomastus male palps possess three different 68 14 9. Type of conductor: hyaline, 0; sclerotized, 1. tegular structures. Each originates from different 69 15 A hyaline conductor also termed ‘membraneous sec- positions of the tegulum. I call them TA1, TA2, and 70 16 ondary conductor’ by Wanless (1980a: fig. 2g, h), is a TA3, to avoid mislabelling any one of them as 71 17 conductor that is transparent and very thin. A hyaline homologous to the tegular apophysis (or TA) found 72 18 conductor is present on Chinoscopus Simon, 1901 and in most spider male palps. TA1 (‘y’ in Wanless 73 19 Lyssomanes (Galiano, 1962, 1980; Wanless, 1980a: (1980b); ‘subtegular apophysis’ in Zhang & Li (2005) 74 20 fig. 2g, h). The presence of a hyaline conductor might is unique to Onomastus (Figs 4A, 10A, 14F, 17A, 75 21 turn out to be a synapomorphy that unites Chinosco- 25B, E, 28A). TA1 is not known to occur in any 76 22 pus and Lyssomanes, two exclusively New World other Salticidae (Wanless, 1980a, b). This character 77 23 genera. is diagnostic for Onomastus. 78 24 79 17. TA2: absent, 0; present, 1. 80 25 10. Shape of conductor: filiform, 0; stout, 1; broad, 2. The presence of TA2 (‘x’ in Wanless (1980b) is unique 81 26 Conductor shape varies from an elongated filiform- to the two Indian Onomastus, O. patellaris and O. 82 27 apophysis to a large disk-shaped structure. Scored as indra (Figs 4A, 17A). In addition to TA1 the male 83 28 filiform for Onomastus of Sri Lanka (Figs 11D, 19B, pulps of Indian Onomastus carry a second tegular 84 29 E), as stout for Onomastus of India (Figs 4A, 17A), apophysis termed TA2 in this study. It originates from 85 30 and as broad for the species of the South-East Asia the prolateral margin of the male palp, distally to the 86 31 clade (Figs 2B, 3A, 11D, E). TA1 (Figs 4A, 17A). 87 32 88 33 11. Number of hooks on the tip of C: one, 0; two, 1. 18. TA3: absent, 0; present, 1. 89 34 The conductor tip of O. indra, O. kano, and O. patel- TA3 is situated distally to TA1 and TA2, on the 90 35 laris is furnished with a single hook (Figs 4A, 7A, E, ventral/distal surface of the tegulum (Fig. 25H, 28D). 91 36 17A). The rest have a bifurcated tip ending with two TA3 is present in all known species from Sri Lanka 92 37 pointed hooks (Fig. 11A–C). and India (Figs 4A, 9B, 19E, 25E, H, 27A, 28D). 93 38 39 12. MA: absent, 0; present, 1. 94 40 All species of Onomastus have a heavily sclerotized Female genitalia 95 41 MA. They are branched and species-specific in shape 19. Female copulatory ducts: absent, 0; present, 1. 96 42 (Figs 9A, B, 11D, 14D, E, 17A, 25G, H; Wanless, Copulatory ducts are absent in Onomastus (Wanless, 97 43 1980b). MA is present in Hispo (Wanless, 1981) and 1980b). The exceptions are Onomastus kaharian, O. 98 44 Lyssomanes. However, it is absent in Chinoscopus complexipalpis, and O. simoni, all of the South-East 99 45 (Wanless, 1980a). Asia clade. It is present in all outgroup taxa (Wanless, 100 46 1980a).The female of O. kanoi is unknown; however, I 101 47 13. MA distal end: bifurcate, 0; smooth/tapering, 1. predict that it would have copulatory ducts. 102 48 In O. nigricauda, O. rattotensis, and O. complexipal- 103 49 pis, the MA is distally bifurcated, diverging into two 20. EF: absent, 0; present, 1. 104 50 branches (Figs 11D, 28A). Smooth or tapering in the EF, also called ‘septum’ in Zhang & Li (2005) are 105 51 rest (Figs 14D, E, 17A, 25G, H). absent in most Onomastus, instead the CO is a simple 106 52 opening in the cuticle (Wanless, 1980b). The excep- 107 53 14. MAP: absent, 0; present, 1. tions are species of the South-East Asia clade 108 54 The MA has a mesal branch that supports the con- (Fig. 5C–E; Zhang & Li, 2005). EF are also absent in 109 55 ductor embolus complex, which is present in all the outgroup taxa (Wanless, 1980a). 110

6 S. P. BENJAMIN

PA AA BB PA CC

SP MA

C E TA1 DD EE FF

MA C GG HH

1 Figure 2. Scanning electron micrographs of Onomastus complexipalpis from South Kalimantan, right male palp 2 (RMNH). A, H, prolateral view. B, F, ventral view. C, D, E, G, retrolateral view. Abbreviations: C, conductor; E, embolus; 3 MA, median apophysis; PA, patella apophysis; SP, spur mesal branch of C; TA1, tegular apophysis 1. Scale bars = 20 mm 4 (F, G, H), 30 mm (E), 100 mm (A, B, C, D).

REVISION AND CLADISTICS OF ONOMASTUS 7

1 21. Epigynal lip: absent, 0; present, 1. over convergence (Farris optimization or ACCTRAN). 13 56 2 Present only in Goleba puella (Wanless, 1980a). All multistate characters were treated as non- 57 3 additive (unordered or Fitch minimum mutation 58 4 Somatic morphology model; Fitch, 1971). Missing characters were coded as 59 5 22. Number of eye rows: three, 0; four, 1. ?, and inapplicable characters as -. Branches were 60 6 Three considered in Hispo cingulata (Wanless, 1981); collapsed if the minimum possible branch length was 61 7 four in all other taxa in the matrix (Galiano, 1962, zero (‘amb-’) or if there was no character that could be 62 8 1980; Wanless, 1980a, b). optimized to support a node. Tree statistics were only 63 9 calculated from phylogenetically informative charac- 64 1012 12 23. Relative size of PME: PME = PLE, 1; PME < PLE, ters. I also used PAUP (Swofford, 2002) to analyse the 65 11 0. data as carried out in Benjamin (2004); see Benjamin 66 12 PME = PLE in Asemonea and Goleba. PME < PLE in (2004) for justiﬁcation of the analytical parameters. 67 13 the rest. Outgroups are scored based on Wanless Below, characters and states are abbreviated and in 68 14 (1980a). This character might be informative in much bold, e.g. character 2 becomes 2, character 2, state 1, 69 15 broader cladistic studies as well. becomes 2-1. 70 16 71 17 24. Relative distance between PME–PME vs. ALE– 18 ALE: PME–PME subequal or longer than ALE–ALE, RESULTS 72 19 0; PME–PME < ALE–ALE, 1. ONOMASTUS RELATIONSHIPS 73 20 This character was proposed to deﬁne a grouping of 21 genera consisting of Asemonea, Goleba, Macopaeus, The phylogenetic analysis of the above-mentioned, 74 22 and Pandisus by Wanless (1980a), which he termed equally weighted characters performed in NONA 75 23 group III. Group III was considered by him to be (hold10000; hold/200; 1000 replicates of tree 76 24 related to group II, which consisted of the genera bisection-reconnection (TBR) + TBR (mult*1000) pro- 77 25 Lyssomanes and Chinoscopus. Onomastus was duced one tree of minimal length. Analysis of the 78 26 however, considered not closely allied to either of same data set in PAUP with heuristic search methods 79 27 them and was, thus placed in group I (Wanless, (100 replicates of random taxon addition subjected to 80 28 1980a). This character might be informative in much TBR branch swapping; ‘MulTrees’ set to 100000; 81 29 broader cladistic studies as well. branches collapsed if the minimum possible branch 82 30 length was zero (‘amb-’; ACCTRAN or Farris optimi- 83 31 25. Relative size of PLE to PME: much smaller, 0; zation) produced the same minimal length tree found 84 32 similar in size, 1. with NONA. This tree was selected as the preferred 85 33 PME are similar in size in Asemonea and Goleba. phylogenetic hypothesis for Onomastus (Fig. 1). Ono- 86 34 PME are much smaller in the rest. Outgroups are mastus is resolved at node 3 supported by four unam- 87 35 scored based on Wanless (1980a). biguous synapomorphies. Within Onomastus, species 88 36 are grouped into two clades, which are named here as 89 37 26. Prosoma coloration: red/brown, 0; light green, 1. the South Asia clade (SA clade) which includes species 90 38 Lyssomaninae are considered to be transparent green from Sri Lanka and India and the South-East 91 39 in colour (Wanless, 1980a; Maddison & Needham, Asian clade (SE clade), which includes the remaining 92 40 2006). However, available information in the litera- diversity. 93 41 ture on prosoma coloration is rather scanty. All Ono- 94 42 mastus are clearly green (Figs 8A–D, 18A–F) and so 43 are Asemonea tenuipes and Lyssomanes viridis (pers. DISCUSSION 95 44 observ.). The remaining taxa are scored based on the 45 their descriptions by Wanless (1980a, 1981). Onomastus is a relatively small genus of jumping 96 spiders mostly known from Sri Lanka. This study 97 46 adds four more species from Sri Lanka, India, and 98 47 ANALYSIS South-East Asia. Noteworthy is the micro endemism 99 48 The parsimony analyses were performed using the of species of the SA clade. In contrast, species of the 100 49 computer program NONA 2ed (Goloboff, 1999). Win- SE clade seem to be widespread. This study indicates 101 50 Clada version 1.00.08 (Nixon, 1999) and MESQUITE that Onomastus of Sri Lanka and India may have 102 51 version 1.12 (Maddison & Maddison, 2006) were used originated from a common ancestor and subsequently 103 52 to study character optimizations on the cladograms evolved independently of each other. This is not 104 53 and to build and edit the character matrix, respec- unusual and has been reported for several faunal 105 54 tively. Ambiguous character optimizations were groups from Sri Lanka (Bossuyt et al., 2004). Species 106 55 resolved so as to favour reversal or secondary loss of the SA clade are spot endemics found mostly in 107

8 S. P. BENJAMIN

1 higher elevation habitats. They may be highly in ment of Goleba. However, these hypotheses of rela- 56 2 danger of extinction because of habitat loss and tionships need to be further tested as the present 57 3 climate change. cladistic analysis was designed to test the monophyly 58 4 Wanless (1980b) considered Onomastus distinct and the internal structure of Onomastus and not that 59 5 from any other Lyssomaninae, although no clear argu- of Lyssomaninae. 60 6 ments were provided. However, this study suggests The tracheal system has been widely used in the 61 7 that it is a genus within a monophyletic Lyssomaninae classiﬁcation of spiders (Hormiga, 1994 and refer- 62 8 (node 1), which is supported by two unambiguous ences). The Lyssomaninae have been characterized as 63 9 synapomorphies (22-1, 26-1). The inclusion of Ono- a group that possess a tracheal system that terminates 64 10 mastus within Lyssomaninae is in agreement with in the opisthosoma (Wanless, 1980a and references) as 65 11 results of Wijesinghe (1997). However, although the is the case in Hispo (Wanless, 1981). Onomastus seem 66 12 14 14 monophyly of Lyssomaninae is widely accepted (Wijes- to be an exception (Fig. 27D, E). However, careful 67 13 inghe, 1997; Maddison & Hedin, 2003; Maddison & preparation of specimens seems to be critical to observ- 68 14 Needham, 2006), further assessments with the addi- ing the ﬁne tracheae that enter the prosoma. 69 15 tion of more outgroup taxa are considered necessary. 70 16 This study suggests that Onomastus is sister to the 17 New World genus Lyssomanes (Fig. 1). Interestingly SYSTEMATICS 71 18 Onomastus and Lyssomanes occupy similar ecological ONOMASTUS SIMON, 1900 72 19 niches. However, this result is in contrast to that of Type species: Onomastus nigricauda by original 73 20 Wijesinghe (1997) who suggested that Onomastus is designation. 74 21 sister to rest of the lyssomanines. Together, Onomas- 75 22 tus and Lyssomanes come out as sister to a grouping 23 of three Old World genera, Pandisus, Asemonea,and Remarks: Listed as O. nigricaudus in Platnick (2008). 76 24 Goleba informally termed here as the Pandisus clade However, nigri-cauda is a substantive and thus needs 77 25 (node 2), which is essentially ‘group lll’ proposed by no gender adjustment. 78 26 Wanless (1980a). The monophyly of the Pandisus 79 27 clade is supported by three unambiguous synapomor- Diagnosis: Onomastus is separated from all other 80 28 phies (1-1, 12-0, 24-1). This study sheds more light on Salticidae by the presence of TA1 (‘y’ in Wanless, 81 29 the phylogenetic position of Goleba, which was 1980b) in the male palp (16-1). The following addi- 82 30 unclear in Maddison & Needham (2006). They did not tional characters support the monophyly of Onomas- 83 31 include Onomastus, Pandisus,orAsemonea in their tus and may serve to separate it: loss of the RTA (7-1), 84 32 molecular study, which might have affected place- presence of a hyaline conductor (9-1), the dorsal 85

34 KEY TO MALE ONOMASTUS (O. SIMONI IS EXCLUDED FOR LACK OF MATERIAL) 35 1 Male palp with TA2 (Figs 4A, 17A); endemic to India...... 2 36 – Male palps without TA2...... 3 37 2(1) TA2 with upward hook, relatively longer (Fig. 17A)...... Onomastus patellaris 38 – TA2 otherwise, TA1 and TA2 shorter (Fig. 4A)...... Onomastus indra sp. nov. 39 3(1) Male palps with a broad, in some cases, disk-shaped C (Figs 2B, 6B, 7C), female epigynum with folds, vulva with 40 clearly defined CD (3B, 5D)...... 7 41 – Male palp with filiform C, female epigynum without folds, vulva without CD, all species endemic to Sri Lanka 42 ...... 4 43 4(3) MA deeply bifurcate, prolateral section with pleats (Figs 27A, 28B, C) ...... Onomastus rattotensis sp. nov. 44 – MA without pleats ...... 5 45 5(4) MA less strongly bifurcated (Fig. 10A) ...... Onomastus nigricauda 46 – MA tip bulb like (Figs 20D, 23A)...... 6 47 6(5) Tip of MA pointed, spermatheca oval (Figs 23A, 24C, D...... Onomastus quinquenotatus 48 – Tip of MA stout, spermatheca relatively round (Figs 20D, 21B, C)...... Onomastus pethiyagodai sp. nov. 49 7(3) C broad, partly covers the tegulum, conductor with SP (Figs 2G, 7D) ...... 8 50 – C filiform resting between MA and tegulum...... 9 51 8(7) C large, covers most of the tegulum, MA tapering with a bifurcated tip (Figs 2A, 3A)...... 52 ...... Onomastus complexipalpis 53 – C smaller, MA stout (Fig. 7C, D)...... Onomastus kanoi 54 9(7) MA chisel-shaped, CD absent (Figs 14E, 16D) ...... Onomastus nigrimaculatus 55 – MA scoop like, CD C-shaped (Figs 5D, 6D)...... Onomastus kaharian sp. nov.

REVISION AND CLADISTICS OF ONOMASTUS 9

1 KEY TO FEMALE ONOMASTUS (FEMALES OF O. KANOI REMAIN UNKNOWN) 2 1 CD absent or very short, endemic to Sri Lanka and India ...... 5 3 – CD clearly visible, epigynal folds present ...... 2 4 2(1) CD characteristically long, coiled (Fig. 3B)...... Onomastus complexipalpis 5 – CD relatively shorter or absent, epigynal folds present (Fig. 5D–E) ...... 3 6 3(2) CD absent, S round (Fig. 16C–E) ...... Onomastus nigrimaculatus 7 – CD present, epigynal folds present...... 4 8 4(3) CD inverted ‘C’-shaped (Fig. 5D, E)...... Onomastus kaharian sp. nov. 9 – CD inverted ‘L’-shaped, running partly parallel to each other...... Onomastus simoni 10 5(1) S round, endemic to South India ...... Onomastus patellaris, Onomastus indra sp. nov. 11 – S elongated endemic to Sri Lanka .....Onomastus nigricauda, Onomastus pethiyagodai sp. nov., Onomastus 12 quinquenotatus, Onomastus rattotensis sp. nov.

14 origin of the embolus (15-1). Further, Onomastus is A 15 separated from all other Salticidae except for Lysso- 16 manes and Chinoscopus by the presence of a conduc- 17 tor (8-1) and the lack of CD (19-0; gained in O. 18 kaharian, O. kanoi, and O. complexipalpis). The lack 19 of a clearly defined fovea might also serve to charac- E 20 terize Onomastus. See also Wanless (1980b) for a 21 detailed diagnosis. MA 22 23 Description: Male palp: complex and species-specific, C 24 suggesting that they have undergone rapid divergent 25 evolution. Femur lacks any form of modification such 26 as apophysis or furrows (Figs 3A, 26B). Patella with a PTA 27 retrolateral apophysis (Figs 3A, 4A, 6F). Tibia with a 28 reduced retrolateral apophysis, hardly visible in most 29 cases. Cymbium distally tapering (‘finger-like exten- 30 sion’ in Wanless, 1980b), proximal promargin sclero- 31 tized and fringed with setae (Figs 2H, 11C, 15C, 28G). 32 Embolus broad based (EP in Fig. 9A), tapering, 33 slender, moderate to long in length, originates from CD 34 the dorsal parts of the tegulum within the alveolus B 35 (Figs 5B, 9A). It may be used in some species as a CO 36 mating plug (Fig. 11F, G). Tegulum large, makes up to 37 half of the copulatory bulb. Distal tegular end tapers 38 to an apophysis unique to Onomastus (TA1; ‘Y’ in 39 Wanless, 1980b). The prolateral margin of O. patel- 4015 15 laris and O. indra holds apophysis unique to them 41 (TA2; ‘X’ in Wanless (1980b). A third tegular apophy- 42 sis is present in some species (Figs 4A, 10A, 19E, 4316 16 27A), but absent or reduced in others. The apical half FD 44 of the copulatory bulb constitutes the MA and 45 conductor–embolus complex. The conductor originates 46 from in between the tegulum, which is grooved on the 47 peripheral margin to form an embolic guide (e.g. in Figure 3. Onomastus complexipalpis from South Kali- 55 48 Wanless, 1980b). Its tip is species-specifically modi- mantan (RMNH). A, left male palp (BMNH 1979.7.4.1). B, 56 49 fied (spur in Wanless, 1980b). Except for O. rattoten- vulva, ventral view. Abbreviations: C, conductor; CD, copu- 57 50 sis, which has three distal branches, the MA is well latory duct; CO, copulatory opening; E, embolus; FD, fer- 58 tilization duct; MA, median apophysis; PTA, ••. Scale 59 51 sclerotized and distally bifurcate. The distal branches bars = 0.1 (B), 0.5 (A). 60 52 are species-specifically modified. In the South Asian 53 species the base of the MA possesses an apophysis 54 (MAP) that serves as a rest for the apical portion of

10 S. P. BENJAMIN

E MA E MAP MA C TA2 C TA2 TA3

TA1 CY

A B T PA PA

CO C

FD DE

1 Figure 4. Onomastus indra sp. nov. (CAS). A, left male palp, ventral view. B, ditto, retrolateral view. C, epigynum, 2 ventral view. D, vulva, ventral view. E, ditto, dorsal view. Abbreviations: C, conductor; CO, copulatory opening; CY, 3 cymbium; E, embolus; FD, fertilization duct; MA, median apophysis; MAP, mesal branch of MA; PA, patella apophysis; 4 S, spermatheca; T, tibia; TA1, tegular apophysis 1; TA2, tegular apophysis 2; TA3, tegular apophysis 3. Scale bars = 0.05 5 (E), 0.1 (C, D), 0.2 (A, B).

6 the embolus–conductor complex (Figs 4A, 11D, 19E). the Sri Lanka–Western Ghats biodiversity hotspot. 13 7 Epigynum: simple to absent. The CO leads directly to The Sri Lankan endemics are restricted to small 14 8 the spermathecae. CD lacking in most species. Sper- patches of rain forest and are highly endangered as a 15 9 mathecae are large and well sclerotized. See also result of habitat loss. 16 10 Wanless (1980b) for a detailed description. 17 11 Distribution: Oriental region, from Sri Lanka to Composition: Eleven species: Onomastus complexipal- 18 12 Japan. However, 55% of the species are endemic to pis Wanless, 1980, O. indra sp. nov., O. kaharian sp. 17 19

REVISION AND CLADISTICS OF ONOMASTUS 11

1 nov., O. kanoi Ono, 1995, O. nigricauda Simon, 1900, ONOMASTUS COMPLEXIPALPIS WANLESS, 1980 11 2 Onomastus nigrimaculatus Zhang & Li, 2005, O. (FIGS 2A–H, 3A–B) 12 3 patellaris Simon, 1900, O. pethiyagodai sp. nov., O. Holotype: 1¢ from Indonesia: Borneo, Santan, Kali- 18 13 4 quinquenotatus Simon, 1900, O. rattotensis sp. nov., mantan, 03.vii.1976. Leg. R. Thompson, BMNH 14 5 and O. simoni Zabka, 1985. 1979.7.4.1, examined. 15

MA E C

C E

A B EF EF

C E

7 Figure 5. Onomastus kaharian sp. nov. from Central Kalimantan (RMNH). A, left male palp, ventral view. B, ditto, 8 retrolateral view. C, epigynum, ventral view. D, vulva, ventral view. E, ditto, dorsal view. Abbreviations: C, conductor; CO, 9 copulatory opening; E, embolus; EF, epigynal folds; FD, fertilization duct; MA, median apophysis. Scale bars = 0.1 (C–E), 10 0.2 (A, B).

12 S. P. BENJAMIN

AA BB CC

MA E C PA C C PT

T DD EE FF

1 Figure 6. Onomastus kaharian sp. nov. from Central Kalimantan (RMNH). A, left male palp, prolateral view. B, D, 2 ditto, ventral view. C, E, F, Ditto, retrolateral view. Abbreviations: C, conductor; E, embolus; MA, median apophysis; PA, 3 patella apophysis; PT, patella; T, tibia. Scale bars = 20 mm (D, E), 30 mm (A, B, C, F). 4 5 Diagnosis: Onomastus complexipalpis is readily dis- patella 0.5, tibia 1.3, metatarsus 1.0, tarsus 0.5. 13 6 tinguished from all known species of Onomastus by Prosoma oval, almost as wide as long. Opisthosoma 14 7 the large palpal conductor in the males and the long oval, longer than wide. All legs laterally without dark 15 8 coiled copulatory ducts in the females (Figs 2A–H, markings. All eyes except AME surrounded by dark 16 9 3A–B). rings. Chelicerae with pro- and retromarginal teeth, 17 10 number not examined. Leg formula 4132. Leg spina- 20 18 11 19 19 Male from South Kalimantan: Total length: 3.9; tion: leg 1: metatarsus D 1-0-0, V 2-2-2; tibia D 1-1-0, 19 12 prosoma length: 1.8, width: 1.3. Leg I: femur 1.2, V 2-2-2; femur D 1-1-2, P 0-0-1, R 0-0-1. Palps as in 20

REVISION AND CLADISTICS OF ONOMASTUS 13

1 Figures 2A–H and 3A. A detailed description of the primary forest, in leaf litter, 2–16.ix.1985. Leg. 22 19 2 male is provided by Wanless (1980b). Suharto Djojosudharmo. 1¢, 1 ™; South Kalimantan, 23 20 3 40 km north-west of Palangkaraya, secondary forest, 21 4 Female from South Kalimantan: Total length: 3.3; 1¢, 2 ™ (right palp on a sputter-coated SEM stub). 22 5 prosoma length: 1.1, width: 0.9. Leg 1: femur 1.5, Deposited in RMNH. 23 6 patella 0.5, tibia 1.4, metatarsus 1.0 tarsus 0.4. Mor- 24 7 phology as above, except for the lighter coloured ONOMASTUS INDRA SP. NOV. (FIG. 4A–E) 25 8 prosoma and opisthosoma. Leg formula 4132. Leg 9 spination: leg 1: metatarsus V 2-1-2; tibia V 2-2-3; Holotype: ¢ from India, Kodaikanal, Grade, 1600 m, 26 10 femur D 1-0-1, P 0-0-1, R 0-0-1. Epigynum and vulva 30.iii.1962, leg. E. S. Ross and D. Q. Cavagnaro. 27 11 as in Figure 3B. Deposited in CAS. 28 12 29 13 Other material examined: Indonesia, Central Kali- Paratypes: 2 ™ from the same locality as the holotype. 30 1421 21 mantan, Kaharian, 20°2′S, 113°40′E. Swampy Label data as above. Deposited in CAS. 31 AA BB CC SP

PA MA C C

PT T T FF TA1 EE DD

16 Figure 7. Onomastus kanoi from Okinawa Prefecture (USNM). A, E, left male palp, ventral view. B, ditto, prolateral 17 view. C, D, ditto, retrolateral view. F, proﬁle of PA, retrolateral view. Abbreviations: C, conductor; MA, median apophysis; 18 PA, patella apophysis; PT, patella; SP, spur mesal branch of C; T, tibia; TA1, tegular apophysis 1. Scale bars = 100 mm.

14 S. P. BENJAMIN

AA BB

CC DD

1 Figure 8. Photographs of live Onomastus nigricauda from Bodinagala, Sri Lanka (MHNG). A, B, male. C, D, female. 2 3 Etymology: Indra is the deity of war in the Hindu Female paratype: Total length: 4.4; prosoma length: 26 4 religion. I use the species epithet as a noun in 2.0, width: 1.3. Leg 1: femur 1.6, patella 0.6, tibia 1.4, 27 5 apposition. metatarsus 1.2, tarsus 0.6. Morphology as above, 28 6 except for the lighter coloured prosoma and opistho- 29 soma and leg spination given below. Leg spination: leg 30 7 Diagnosis: Separated from all other species except for 1: metatarsus V 2-1-2; tibia V 3-2-2; femur D 1-1-1. 31 8 O. patellaris by the occurrence of TA2 in males. Sepa- Epigynum and vulva as in Figure 4C–E. 32 9 rated from O. patellaris by the shorter, stouter TA1 33 10 24 24 and TA2 (Fig. 4A, B); the further end of TA2 is in the Other material examined: None. 34 11 shape of an upward hook in O. patellaris (Fig. 17A). 35 12 13 Male holotype: Total length: 3.8; prosoma length: 2.0, ONOMASTUS KAHARIAN SP. NOV. 36 14 width: 1.3. Leg I: femur 1.6, patella 0.3, tibia 0.8, (FIGS 5A–E, 6A–F) 37 15 metatarsus 0.7, tarsus 0.3. All spiders are whitish to Holotype: ¢ from Indonesia, Central Kalimantan, 38 16 pale yellow in alcohol, no markings. Prosoma oval, Kaharian, 20°2′S, 113°40′E. Swampy primary forest, 39 17 almost as wide as long, laterals darker. Opisthosoma in leaf litter, 2–16.ix.1985. Leg. Suharto Djojosud- 40 18 oval, longer than wide, lighter in colour. Legs pale harmo. Deposited in RMNH. 41 19 yellow, laterally without dark markings; might be a 42 20 result of preservation. All eyes except AME sur- Paratypes: ™ from Indonesia, locality and label data 43 21 rounded by dark rings. Chelicerae with pro- and ret- as above. Deposited in RMNH. 44 22 romarginal teeth, number not examined. Leg formula 45 23 4132. Leg spination: leg 1: metatarsus V 2-1-2; tibia V 24 2-2-3; femur D 0-0-1, P 0-1-2. Palps as in Figure 4A Etymology: The species name is a noun in apposition, 46 25 and B. taken from the type locality. 47

REVISION AND CLADISTICS OF ONOMASTUS 15

Male holotype: Total length: 3.4; prosoma length: 1.5, 23 A width: 1.0. Leg I: femur 1.6, patella 0.5, tibia 1.7, 24 metatarsus 1.1, tarsus 0.5. All spiders are whitish to 25 pale yellow in alcohol, no markings. Prosoma oval, 26 almost as wide as long, laterals darker. Opisthosoma 27 BH oval, longer than wide, lighter in colour. Legs pale 28 yellow, laterally without dark markings; might be a 29 ST result of preservation. All eyes except AME 30 EP T surrounded by dark rings. Chelicerae with pro- and 31 retromarginal teeth, number not examined. Leg 32 E formula 4132. Leg spination: leg 1: metatarsus V 33 2-1-2; tibia V 2-2-3; femur D 0-0-1, P 0-1-2. Palp as in 34 C MA Figures 5A, B and 6A–F. 35 TA1 36 Female paratype: Total length: 3.5; prosoma length: 37 1.4, width: 1.0. Leg 1: femur 1.4, patella 0.4, tibia 1.5, 38 TA3 metatarsus 1.1, tarsus 0.4. Morphology as above, 39 except for the lighter coloured prosoma and opistho- 40 41 MA soma and leg spination given below. Leg spination: leg 1: metatarsus V 2-1-2; tibia V 3-2-2; femur D 1-1-1. 42 B Epigynum and vulva as in Figure 5C–E. 43 44

Distribution: Thailand and Indonesia. 45

Other material examined: Indonesia: Central Kali- 47 PA mantan, Kaharian, 20°2′S, 113°40′E. Swampy 48 primary forest, in leaf litter, 2–16.ix.85, 2¢ (a single 49 PT left palp on a sputter-coated SEM stub), 5™, leg. 50 Suharto Djojosudharmo, deposited in RMNH. Thai- 51 land: Nakhon Si Thammarat Prov. Khao Luang NP, 52 8°43′25.2″ N, 99°40′7.7″E, 355 m, 10–12.x.2003 53 1 Figure 9. Onomastus nigricauda from Bodinagala (ATOL expedition 2003), I¢ 1™, deposited in 54 2 (MHNG). A, expanded left male palp, prolateral view. B, USNM. 55 3 ditto, retrolateral view. Abbreviations: BH, basal hae- 4 matodocha; C, conductor; E, embolus; EP, embolic base; 56 5 MA, median apophysis; PA, patella apophysis; PT, patella; 6 ST, subtegulum; T, tibia; TA1, tegular apophysis 1; TA3, ONOMASTUS KANOI ONO, 1995 (FIG. 7A–E) 57 7 tegular apophysis 3. Scale bar = 0.2 mm. Diagnosis: Onomastus kanoi is readily distinguished 58 from all known species of Onomastus except for O. 59 8 complexipalpis by the large disk-shaped palpal con- 60 ductor in the males (Fig. 7A–D). The conductor of O. 61 9 Diagnosis: Males are readily separated from O. patel- kanoi is relatively smaller than that of O. complexi- 62 10 larisandO. indra by the absence of TA2. Separated palpis (Fig. 7D). The female of O. kanoi is unknown. 63 11 from O. rattotensis, O. nigricauda, O. quinquenotatus, However, I predict that the CD of O. kanoi females 64 12 and O. pethiyagodai by the presences of a broad would be longer than that of all other Onomastus,but 65 13 conductor (Fig. 6D). Separated from O. complexipal- shorter than that of O. complexipalpis. 66 14 pis, O. kanoi,andO. nigrimaculatus by the scoop-like 15 MA and shape of C (Fig. 6D). Females could be 67 16 separated from all other Onomastus by the presence Male: Total length: 5.2; prosoma length: 2.0, width: 68 17 of semicircular epigynal fold and C-shaped CD 1.5. Leg I: femur 2.0, patella 0.6, tibia 2.0, metatarsus 69 18 (Fig. 5D, E). 2.0, tarsus 0.6. For a detailed description see Ono 70 19 (1995). 71 20 Remarks: Onomastus simoni appears to be closely 72 21 related to O. kaharian. They may be separated from 22 each other by the parallel course of CD in O. simoni. Female: Unknown. 73

16 S. P. BENJAMIN

MA C B TA3

CY TA1

PA A C

FD S

1 Figure 10. Onomastus nigricauda. A, male lectotype (MNHN 20404/1-3), right palp, ventral view. B, female (found in the 2 same vial as the lectotype). C, female from Bodinagala, epigynum, ventral view. D, ditto, vulva, ventral view. E, ditto, 3 dorsal view. Abbreviations: C, conductor; CO, copulatory opening; CY, cymbium; FD, fertilization duct; MA, median 4 apophysis; PA, patella apophysis; S, spermatheca; TA1, tegular apophysis 1; TA3, tegular apophysis 3. Scale 5 bars = 0.1 mm (C), 0.2 mm (A, B, D, E).

REVISION AND CLADISTICS OF ONOMASTUS 17

AA BB CC

MA C

C MAP DD EE

CO MP

FF GG

1 Figure 11. Scanning electron micrographs of Onomastus nigricauda from Bodinagala (MHNG). A, left male palp, 2 prolateral view. B, D, ditto, ventral view. C, E, ditto, retrolateral view. F, G, epigynum, ventral view. Note the mating plug 3 in F and G. Abbreviations: C, conductor; CO, copulatory opening; MA, median apophysis; MAP, mesal branch of MA; MP, 4 mate plug. Scale bars = 10 mm (G), 100 mm (A–F).

18 S. P. BENJAMIN

ONOMASTUS NIGRICAUDA SIMON, 1900 (FIGS 8A–D, 12 9A–B, 10A–E, 11A–G, 12A–C, 13A–B) 13 Onomastus nigricauda Simon, 1900: 29. ¢ lectotype 14 and 2 ™ from Sri Lanka, Galle, in MNHN (20404/ 15 1-3), examined. 16 Onomastus nigricauda Simon, 1901: 400, fig. 398. 25 17 Onomastus nigricauda Wanless, 1980c: 181, fig. 26 18 1A–G. Initial description of the female and desig- 19 nation of types. 20 21 Diagnosis: Males are readily separated from O. patel- 22 laris and O. indra by the absence of TA2. Separated 23 AA from O. complexipalpis, O. kanoi, O. kaharian,andO. 24 nigrimaculatus by the filiform conductor. Separated 25 from O. pethiyagodai and O. quinquenotatus by the 26 shape of the MA. Onomastus nigricauda seems to be 27 most closely related to O. rattotensis, but may be 28 separated by the large partly pleated MA in O. rat- 29 totensis (Figs 27A, 28A–C). 30 31 Male (from Bodinagala): Total length: 3.0; prosoma 32 length: 1.5, width: 1.1. Leg I: femur 1.4, patella 0.4, 33 tibia 1.5, metatarsus 0.9, tarsus 0.5. The spiders are 34 green in nature. They turn whitish to pale yellow in 35 alcohol. Prosoma oval, longer than wide. Opistho- 36 soma longer than wide, lighter in colour. Leg I yel- 37 B lowish brown, leg II–IV light yellow. Legs are 38 B laterally without dark markings. All eyes except 39 AME surrounded by dark rings. Chelicerae are 40 yellow, with seven retromarginal and three promar- 41 ginal teeth. Leg formula 1432. Leg spination, meta- 42 tarsus D 1-0-0, V 2-2-2; tibia D 0-0-1, P 1-0-0, V 43 4-4-2, R 1-0-0; patella D 0-1-0; femur D 2-1-0, P 44 0-0-1, R 0-0-1. Palp: (Figs 9A–B, 10C, 11A–E, 45 12A–C, 13A–B). Tegulum as in Figure 9A, projecting 46 outwards, path of sperm duct with two loops as in 47 Figure 9A and B, embolus as in Figure 9A. A 48 detailed description of the male lectotype is also 49 provided by Wanless (1980b). 50 51 Female (from Bodinagala): Total length: 4.0; prosoma 52 CC length: 1.5, width: 1.2. Leg 1: femur 1.4, patella 0.5, 53 tibia 1.5, metatarsus 1.1, tarsus 0.5. Morphology as 54 above, except for the lighter coloured prosoma and 55 1 Figure 12. Scanning electron micrographs of Onomastus 2 nigricauda from Bodinagala (MHNG). A, profile of MA. B, opisthosoma. Leg spination, metatarsus V 4-2-2, P 56 3 ditto, detail. C, profile of TA1. Scale bars = 5 mm (B), 10 mm 1-0-0, R 1-0-0; tibia D 1-1-0, V 4-4-2; patella P 1-0-0, 57 4 (A, C). R 1-0-0; femur D 1-1-1, P 0-0-1, R 0-0-1. Epigynum 58 and vulva as in Figures 10B–E and 11F–G. A detailed 59 5 description of the female is provided by Wanless 60 6 Distribution: Endemic to Okinawa Island of Japan. (1980b). 61 62 7 Variation: The male specimen from Morningside has a 63 8 Material examined: Japan, Okinawa Prefecture, Oki- bifurcate MA in which both branches taper towards 64 9 nawajima Island, Kijoka. 01.iv.1997. 1¢; Japan, the end. Live male spiders of O. nigricauda from 65 10 Okinawa Prefecture, Okinawajima Island, Zutsun. Bodinagala tend to also have a much darker palp 66 11 01.iv.2000, 1¢. All leg. Akio Tanikawa. than that of Morningside. 67

REVISION AND CLADISTICS OF ONOMASTUS 19

A B

1 Figure 13. Onomastus nigricauda from Morningside (MHNG). A, left male palp, ventral view. B, ditto, retrolateral view. 2 Scale bars = 0.2 mm. 3 4 Distribution: Endemic to Sri Lanka; Bodinagala Diagnosis: Males of O. nigrimaculatus are readily 26 5 forest reserve is a remaining fragment of the once separated from those of O. patellaris and O. indra by 27 6 vast rain forest that covered the south-west of the the absence of TA2. Separated from O. rattotensis, O. 28 7 island. It is not far from the type locality, Galle. nigricauda, O. quinquenotatus, and O. pethiyagodai 29 8 by the presence of a broad conductor and chisel- 30 shaped MA. Separated from O. complexipalpis, O. 31 9 Other material examined: SRI LANKA: Western prov- kanoi, and O. kaharian by the chisel-shaped MA and 32 10 ince, Kalutara district, Horane, Bodinagala, details of C. Female O. nigrimaculatus could be sepa- 33 11 19.vii.1996, 1¢ 2™, leg. Suresh. P. Benjamin. Same rated from all other Onomastus by the combined 34 12 locality, 10.ii.2007, 5¢ 6™, leg. Suresh. P. Benjamin presence of C-shaped epigynal fold and lack of CD. 35 13 and Ziyard Jaleel. Sabaragamuwa province, Rat- 14 napura District, Morningside section. In primary 36 15 forest, 23.ii.2007, 1¢, leg. Suresh P. Benjamin and Male: Total length: 4.0; prosoma length: 1.8, width: 37 16 Ziyard Jaleel. All specimens deposited in MHNG. 1.3. Leg I: femur 1.4, patella 0.5, tibia 1.7, metatarsus 38 17 1.4, tarsus 0.8. Prosoma oval, almost as wide as long, 39 laterals with a faint black border. Opisthosoma oval, 40 18 ONOMASTUS NIGRIMACULATUS ZHANG &LI, 2005 longer than wide, lighter in colour, four to six dark 41 19 (FIGS 14A–F, 15A–B, 16A–E) black spots on dorsum. Legs light yellow, leg 1 later- 42 20 Holotype: Male from China, Mingfeng Valley, Jianfen- ally with dark black spots, legs II–IV do not have 43 21 gling National Park, Hainan Province. Deposited in similar spots. All eyes except AME surrounded by 44 22 the Raffles Museum of Biodiversity Research (ZRC- dark rings. Chelicerae with pro- and retromarginal 45 23 ARA-494). As this species was adequately illustrated teeth, number not examined. Leg formula 4132. Leg 46 24 in its original description, I have not examined any spination: leg 1: metatarsus D 1-0-0, V 2-2-2; tibia D 47 25 type material. 1-0-1, V 2-4-2, P 1-0-0, R 1-0-0; patella D 1-0-0, V 48

20 S. P. BENJAMIN AA BB CC

DD EE FF

MA C MA

C TA1 PA

1 Figure 14. Scanning electron micrographs of Onomastus nigrimaculatus from Thailand (RMNH). A, D, left male palp, 2 prolateral view. B, E, F, ditto, ventral view. C, ditto, retrolateral view. Abbreviations: C, conductor; MA, median apophysis; 3 PA, patella apophysis; TA1, tegular apophysis 1. Scale bars = 10 mm (E), 30 mm (D), 100 mm (A–C, F).

REVISION AND CLADISTICS OF ONOMASTUS 21

4132. Leg spination: leg 1: metatarsus V 2-2-2, P 14 AA 1-0-0, R 1-0-0; tibia D 1-0-0, V 2-2-2; Patella D 1-0-0, 15 P 0-1-0, R 0-1-0. Epigynum and vulva as in 16 E Figure 16C–E. 17 MA 18 Distribution: Hainan province, China (Zhang & Li, 19 C 2005) and primary rain forest of the Khao Yai 20 National Park, Thailand. 21 22 Other material examined: Thailand, Khao Yai N.P, 23 3–7.iii.1986, leg. CL and PR Deeleman, 4¢. Same 24 locality as above, 23.x.1985, leg. PR Deeleman, 4™. 25 Same locality, 5.x.1987, leg. CL and PR Deeleman, 26 1¢1™. Same locality, 19–24.x.1985, leg. CL and 27 PR Deelema, 4™. All specimens deposited in 28 BB RMNH. 29 E 30 ONOMASTUS PATELLARIS SIMON, 1900 (FIG. 17A–C) 31 Onomastus patellaris Simon, 1900: 29. 32 33 C Lectotype: Single male from India, Kodaikanal, 34 MNHN 14858, examined. 35 36 Diagnosis: Separated from all other species of Ono- 37 mastus except O. indra by the presence of TA2 in 38 male palps (Fig. 17A). TA1 and TA2 in O. patellaris 39 are filiform. The further end of TA2 is in the shape of 27 40 CC an upward hook in O. patellaris (Fig. 17A). 41 42 Description: See Wanless (1980b). 43 44 Distribution: India, Madras, Kodaikanal. Probably 45 endemic to India. 46 47 Other material examined: Four Ǩ from India, Trichi- 48 nopoly, leg. R. P. Malat, MNHN 14898. 49 50 ONOMASTUS PETHIYAGODAI SP. NOV. (FIGS 18A–F, 51 19A–F, 20A–E, 21A–D, 22A–B) 52 Holotype: 1¢, Sri Lanka: Central Province, Nuwara 53 1 Figure 15. Scanning electron micrographs of Onomastus Eliya District, Agrapathana, Agrabopath forest, 54 2 nigrimaculatus from Thailand (RMNH). A, profile of MA 1100 m, 08.iii.2000. Deposited in MHNG. 55 3 and C, retrolateral view. B, ditto, retrolateral-apical view. 4 C, serrated setae of the cymbium, retrolateral view. Abbre- 56 5 viations: C, conductor; E, embolus; MA, median apophysis. Paratype: 3™, label data as above. Deposited in 57 6 Scale bars = 10 mm. MHNG. 58 59 7 Diagnosis: Males of O. pethiyagodai are readily sepa- 60 8 1-0-0; femur D 1-1-2, P 0-0-1, R 0-0-1. Palp as in rated from that of O. patellarisandO. indra by the 61 9 Figures 14A–F, 15A–C, 16A–B. absence of TA2. Separated from O. complexipalpis, O. 62 10 kanoi, O. kaharian,andO. nigrimaculatus by the 63 11 Female: Total length: 5.0; prosoma length: 2.2, width: presence of a filiform conductor. O. pethiyagodai is 64 12 1.5. Leg 1: femur 2.0, patella 0.7, tibia 2.4, metatarsus most closely related to O. quinquenotatus, and may be 65 13 1.5, tarsus 0.6. Morphology as above. Leg formula separated by the shape of the conductor and MA 66

22 S. P. BENJAMIN

CDE

1 Figure 16. Onomastus nigrimaculatus from Thailand (RMNH). A, left male palp, ventral view. B, ditto, retrolateral view. 2 C, female from the same locality, epigynum, ventral view. D, ditto, vulva, ventral view. E, ditto, dorsal view. Scale 3 bars = 0.2 mm (C–E), 0.5 mm (A, B).

5 (Figs 19E, 20D, 22A). Females are difficult to diag- Prosoma oval, longer than wide. Opisthosoma longer 13 6 nose, but may be separated by details of internal than wide, lighter in colour. Leg I yellowish brown, 14 7 genitalia. leg II–IV light yellow. Legs are laterally marked 15 8 with dark black blobs. All eyes except AME sur- 16 9 Male holotype: Total length: 2.9; prosoma length: 1.5, rounded by dark rings. Chelicerae with pro- and ret- 17 10 width: 1.1. Leg I: femur 1.4, patella 0.2, tibia 1.2, romarginal teeth, number not examined. Leg 18 11 metatarsus 1.0, tarsus 0.4. The spiders are green in formula 4132. Leg spination: metatarsus V 4-2-2; 19 12 nature. They turn whitish to pale yellow in alcohol. tibia P 1-0-0, V 2-4-2, R 1-0-0; patella V 2-0-0; 20

REVISION AND CLADISTICS OF ONOMASTUS 23

E C TA2

TA1

A PA

1 Figure 17. Onomastus patellaris from Kodaikanal, India (MNHN 14858). A, left male palp, ventral view. B, MA, 2 ventral view. C, epigynum, ventral view. Abbreviations: C, conductor; E, embolus; MA, median apophysis; PA, patella 3 apophysis; PT, patella; TA1, tegular apophysis 1; TA2, tegular apophysis 2. Scale bars = 0.1 mm (B), 0.2 mm (C) 4 0.4 mm (A).

6 femur D 0-1-1, P 0-0-1, R 0-0-1. Palp as in Other material examined: SRI LANKA: Central Prov- 22 7 Figures 19A–F, 20A–E, 22A–B. ince, Nuwara Eliya District, Agrapathana, Agrabo- 23 path forest, 1100 m, 07.iii.2000, 1¢ 2™; 1–30.vi.2003, 24 8 7¢ 7™ (a single left palp on a sputter-coated SEM 25 9 Female paratype: Total length: 2.9; prosoma length: stub), all leg. Suresh P. Benjamin. Same locality, 26 10 1.2, width: 1.1. Leg 1: femur 1.0, patella 0.3, tibia 1.0, 08.iii.2000, 1¢ leg. Sudath Nanayakkara. Same local- 27 11 metatarsus 0.8, tarsus 0.4. Morphology similar to the ity, 18–21.ii.2007, 3¢ 2™, leg. Suresh P. Benjamin and 28 12 male except for the following. Prosoma and opistho- Ziyard Jaleel. Central Province, Nuwara Eliya Dis- 29 13 soma lighter in colour. Legs with lateral black blobs. trict, Horton Plains National Park, c. 2000 m, 30 14 Leg spination: metatarsus V 2-2-2; tibia P 1-0-0, V 20–21.ii.2007, 15¢ 5™, leg. Suresh P. Benjamin and 31 15 2-4-4, R 1-0-0; patella P 1-0-0, R 1-0-0; femur D 0-2-1, Ziyard Jaleel. Deposited in MHNG. 32 16 P 0-0-1, R 0-0-1. Epigynum and vulva as in 17 Figure 21A–D. 33

18 ONOMASTUS QUINQUENOTATUS SIMON, 1900 34 19 Distribution: Endemic to the central highlands of Sri (FIGS 23A–D, 24A–D, 25A–H) 35 20 Lanka. Known from Agrapathana, Agrabopath forest Onomastus quinquenotatus Simon, 1900: 29. 2™ from 36 21 and Horton Plains National Park. Sri Lanka in MNHN 20771, examined. 37

24 S. P. BENJAMIN

AA BB

CC DD

EE FF

1 Figure 18. Photographs of live Onomastus pethiyagodai sp. nov. from Agrapathana, Sri Lanka (MHNG). A–C, male. 2 D–F, female.

4 O. quinquenotatus Wanless, 1980: 183, ﬁg. 2a–e. Des- from O. pethiyagodai and O. nigricauda by the shape 11 5 ignation of lectotype and paralectotype. of the MA (Fig. 23B). Onomastus quinquenotatus is 12 6 most closely related to O. pethiyagodai and they may 13 7 Diagnosis: Males are readily separated from O. patel- be separated from each other by the shape of the 14 8 larisandO. indra by the absence of TA2. Separated conductor and MA (Fig. 23A–D). Females are difficult 15 9 from O. complexipalpis, O. kanoi, O. kaharian, and O. to diagnose, but may be separated by details of the 16 10 nigrimaculatus by the ﬁliform conductor. Separated internal genitalia. 17

REVISION AND CLADISTICS OF ONOMASTUS 25

PA AA BB CC

MA MA C MA C E C MAP TA3

TA1

DD EE FF

1 Figure 19. Scanning electron micrographs of Onomastus pethiyagodai sp. nov. from Agrapathana, Sri Lanka 2 (MHNG). A, D, left male palp, prolateral view. B, E, ditto, ventral view. C, F, ditto, retrolateral view. Abbreviations: C, 3 conductor; E, embolus; MA, median apophysis; MAP, mesal branch of MA; PA, patella apophysis; TA1, tegular apophysis 4 1; TA3, tegular apophysis 3. Scale bars = 20 mm (A, D, E, F), 30 mm (B, C).

26 S. P. BENJAMIN

AA BB CC

DD EE

1 Figure 20. Scanning electron micrographs of Onomastus pethiyagodai sp. nov. from Agrapathana, Sri Lanka 2 (MHNG). A, MA, prolateral view. B, PA, retrolateral view. C, ditto, ventral view. D, MA, ventral view. E, C and E 3 prolateral view. Scale bars = 10 mm (B, C, D), 20 mm (A, E).

5 Male from Kanda-ela: Total length: 3.1; prosoma Female from Kanda-ela: Total length: 2.2; prosoma 19 6 length: 1.6, width: 1.3. Legs I: femur 1.6, patella 0.5, length: 1.8, width: 1.3. Leg 1: femur 1.4, patella 0.5, 20 7 tibia 1.5, metatarsus 1.3, tarsus 0.6. Prosoma round tibia 1.6, metatarsus 1.3, tarsus 0.5. Morphology 21 8 to oval, longer than wide, pale amber to light yellow. similar to the male except for the following: prosoma 28 22 9 Eyes with black surrounds except for AME. Opistho- and opisthosoma lighter in colour; legs with lateral 23 10 soma oval, longer than wide, lighter in colour, no black blobs. Leg spination: metatarsus V 4-2-2; tibia 24 11 markings. Living specimens green in colour. Leg 1 V 2-2-2; femur D 0-1-2. Epigynum and vulva as in 25 12 posterior laterally black along its length, all other Figure 24A–D. 26 13 legs uniformly yellow. Chelicerae light yellow, with 27 14 seven retromarginal and three promarginal teeth. 15 Spider ventrally light yellow. Leg formula Distribution: Known from two localities, Kandaela 28 16 4 > 1 > 3 > 2. Leg spination: metatarsus V 4-2-2; tibia and Hakgala, in the central highlands of Sri Lanka. 29 17 D 2-0-0, V 2-4-4; patella D 1-0-0; femur D 0-2-2. Palp The type locality in the labels is given as Sri Lanka 30 18 as in Figures 23A–D, 25A–H. (Simon, 1900). The exact locality of the lectotypes in 31

REVISION AND CLADISTICS OF ONOMASTUS 27

CO CD

A B

D C

1 Figure 21. Onomastus pethiyagodai sp. nov. from Agrapathana, Sri Lanka. A, D, epigynum, ventral view. B, vulva, 2 ventral view. C, ditto, dorsal view. Abbreviations: CD, copulatory duct; CO, copulatory opening; FD, fertilization duct. 3 Scale bars = 0.2 mm.

5 Sri Lanka is unclear. Onomastus quinquenotatus is Etymology: Adjective, ‘from Rattota’, after the type 25 6 endemic to Sri Lanka. locality. 26 7 27 Diagnosis: Males of O. rattotensis are readily sepa- 28 8 Other material examined: 1™ 1¢, SRI LANKA: rated from those of O. patellaris and O. indra by the 29 929 29 Central province, north-east district, Kandaela reser- absence of TA2. Separated from O. complexipalpis, O. 30 1030 30 voir, 5.6 miles south-west of Nuwara Eliya, 6200 ft kanoi, O. kaharian,andO. nigrimaculatus by the 31 11 10–21.ii.1970, leg. Davis and Rowe, deposited in ﬁliform conductor. Separated from O. pethiyagodai 32 12 USNM; 1¢ Hakgala, Hakgala forest, 1600 m, leg. and O. quinquenotatus by the shape of the MA. Sepa- 33 13 Suresh P. Benjamin, 27.vii.1996 Deposited in rated from O. nigricauda by the presence of a well- 34 14 MHNG. developed, pleated MA. Females are difficult to 35 15 diagnose, but may be separated by details of the 36 37 16 ONOMASTUS RATTOTENSIS SP. NOV. internal genitalia. 17 (FIGS 26A–D, 27A–E, 28A–G) 38 Male holotype: Total length: 2.8; prosoma length: 1.4, 39 18 Holotype: ¢ from Sri Lanka. Central Province, Knuck- width: 1.1. Legs I: femur 1.1, patella 0.5, tibia 3.2, 40 19 les range, along Rattota-Ilukkumbura Road, 900 m, metatarsus 2.3, tarsus 1.3. The spiders are green in 41 20 03–04.ix.2003, leg. Suresh P. Benjamin. Deposited in nature. They turn whitish to pale yellow in alcohol. 42 21 MHNG. Prosoma oval, longer than wide. Opisthosoma longer 43 22 than wide, lighter in colour. Leg I yellowish brown, 44 23 Paratype: ™ from Sri Lanka. Data as above. Depos- leg II–IV light yellow. Legs are laterally without dark 45 24 ited in MHNG. markings. Eyes surrounded by dark rings. Chelicerae 46

28 S. P. BENJAMIN

A B

1 Figure 22. Onomastus pethiyagodai sp. nov. from Agrapathana, Sri Lanka (MHNG). A, left male palp, ventral view. 2 B, ditto, retrolateral view. Scale bar = 0.2 mm.

MA B E C

3 TA3 TA1 C

A D

4 Figure 23. Onomastus quinquenotatus, male from Hakgala, Sri Lanka (MHNG). A, left male palp, ventral view. B, proﬁle 5 of MA. C, proﬁle of C. D, left male palp, retrolateral view. Abbreviations: C, conductor; E, embolus; MA, median apophysis; 6 PA, patella apophysis; TA1, tegular apophysis 1; TA3, tegular apophysis 3. Scale bars = 0.1 mm (B, C), 0.2 mm (A, D).

REVISION AND CLADISTICS OF ONOMASTUS 29

A B

CD FD

1 Figure 24. Onomastus quinquenotatus. A, female lectotype (MNHN 20771), epigynum, ventral view. B, female from 2 Kandaela, Sri Lanka (USNM), epigynum, ventral view. C, vulva, ventral view. D, ditto, ventral view. Abbreviations: CO, 3 copulatory opening; FD, fertilization duct. Scale bars = 0.2 mm.

5 yellow, with seven retromarginal and three promar- ACKNOWLEDGEMENTS 26 6 ginal teeth. Leg formula 1432. Leg spination: meta- This study was supported by a Smithsonian Insti- 27 7 tarsus V 4-2-2; tibia D 1-0-0, V 4-2-4; patella D 1-0-0; tution postdoctoral fellowship and my personal 28 8 femur D 0-2-2. Palp: cymbium oval. Retrolateral tibial funds. I am indebted to Christa L. Deeleman- 29 9 apophysis stout, tapering blunt end (Figs 26A, 27B). Reinhold and E. J. van Nieukerken (RMNH) for 30 10 kindly providing a substantial part of the specimens 31 11 Female paratype: Total length: 3.2; prosoma length: examined in this study. Thanks to Christine Rollard 32 12 1.4, width: 1.2. Legs 1: femur 1.3, patella 0.8, tibia and Elise-Anne Leguin (MNHN) for providing E. 33 13 1.8, metatarsus 1.0, tarsus 0.4. Morphology similar to Simon’s types, to Jonathan Coddington and Dana 34 14 the male except for the following. Leg spination: De Roche (USNM) for providing access to additional 35 15 metatarsus V 2-2-2; tibia V 2-2-2; femur D 0-1-0, P material, to Dr Akio Tanikawa for providing speci- 36 16 2-3-1, R 2-3-1. Epigynum and vulva as in mens of O. kanoi, to Janet Beccaloni (BMNH) for 37 17 Figure 26C–E. providing the type of O. complexipalpis, to Charles 38 18 Grisworld (CAS) for providing me with an unde- 39 scribed species of Onomastus from India, and to Mr 31 40 19 Distribution: Sri Lanka, central province, known only A. H. Sumanasena (Department of Wild Life Con- 41 20 from the type locality. Probably endemic to Sri Lanka. servation, Colombo) for providing a research permit 42 21 to collect in Sri Lanka. I want to express my grati- 43 22 Other material examined: SRI LANKA: Central Prov- tude to my brother Donald Benjamin and my 44 23 ince, Knuckles range, along Rattota-Ilukkumbura student Ziyard Jaleel (Open University of Sri 45 24 Road, 900 m, 03–04.ix.2003, 9¢, leg. Suresh P. Ben- Lanka) and Sudath V. Nanayakkara for accompany- 46 25 jamin. Deposited in MHNG. ing me during ﬁeldwork in Sri Lanka. 47

30 S. P. BENJAMIN AA BB CC

TA1

PA DD EE FF E MA C E TA3 C

TA1

GG HH

MA MA

TA3

1 Figure 25. Scanning electron micrographs of Onomastus quinquenotatus male from Hakgala, Sri Lanka (MHNG). A, D, 2 G, prolateral view. B, E, H, ventral view. C, F, retrolateral view. Abbreviations: C, conductor; E, embolus; MA, median 3 apophysis; PA, patella apophysis; TA1, tegular apophysis 1; TA3, tegular apophysis 3. Scale bars = 10 mm (F, H), 30 mm 4 (B–E, G), 100 mm (A).

REVISION AND CLADISTICS OF ONOMASTUS 31

A B

D E

1 Figure 26. Onomastus rattotensis sp. nov. from Rattota, Sri Lanka (MHNG). A, left male palp, ventral view. B, ditto, 2 retrolateral view. C, epigynum, ventral view. D, vulva, ventral view. E, ditto, dorsal view. Scale bars = 0.1 mm (D, E), 3 0.2 mm (A–C).

32 S. P. BENJAMIN A B C

MA E

C MA C MAP TA3

TA1

DD EE

1 Figure 27. Scanning electron micrographs of Onomastus rattotensis sp. nov. from Rattota, Sri Lanka (MHNG). A, left 2 male palp, retrolateral view. B, ditto, retrolateral view. C, ditto, prolateral view. D, female tracheal system, dorsal view. 3 E, ditto, detail. Abbreviations: C, conductor; E, embolus; MA, median apophysis; MAP, mesal branch of MA; TA1, tegular 4 apophysis 1; TA3, tegular apophysis 3. Scale bars = 100 mm. The arrow points to the tracheae, which enter the prosoma.

REVISION AND CLADISTICS OF ONOMASTUS 33

E MA MA

TA1 C MAP

PA AA TA3 BB

TA3

CC C DD

EE FF GG

1 Figure 28. Scanning electron micrographs of Onomastus rattotensis sp. nov. from Rattota, Sri Lanka (MHNG). A, left 2 male palp, ventral view. B, MA and C, ventral view. C, F, MA tip, ventral view. D, proﬁle of TA3, ventral view. E, proﬁle 3 of C tip, ventral view. G, serrated setae of the cymbium. Abbreviations: C, conductor; CY, cymbium; E, embolus; MA, 4 median apophysis; MAP, mesal branch of MA; PA, patella apophysis; TA1, tegular apophysis 1; TA3, tegular apophysis 5 3. Scale bars = 10 mm (B, E, F, G), 20 mm (C, D), 100 mm (A).

34 S. P. BENJAMIN

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