Model mimics: antlike jumping spiders of the genus Myrmarachne from Sri Lanka

Model mimics: antlike jumping spiders of the genus Myrmarachne from Sri Lanka Suresh P. Benjamin a,b * a Department of Ecology and Environmental Biology, Institute of Fundamental Studies, Kandy, Sri Lanka; b Arthropoda Department, Alexander Koenig Research Museum of Zoology, Bonn, Germany (Received 22 April 2014; accepted 17 March 2015; first published online 12 May 2015) All nominal species of ant-mimicking jumping spiders of the genus Myrmarachne from Sri Lanka are redescribed, based on type and newly collected material. Three new species are described: Myrmarachne aurantiaca sp. nov., M. dishani sp.nov.and M. morningside sp. nov. Panachraesta Simon, 1900 is shown to be a junior synonym of Myrmarachne MacLeay, 1839, syn. nov. The following species are synonymized: Myrmarachne orientales Tikader, 1973 = Myrmarachne melanocephala MacLeay, 1839 syn. nov., Myrmarachne paivae Narayen, 1915 and Myrmarachne bengalensis Tikader, 1973 = Myrmarachne prava (Karsch, 1880) syn. nov., Myrmarachne hanoii Zabka, 1985 = Myrmarachne pumilio (Karsch, 1880) syn. nov., Myrmarachne maratha Tikader, 1973 = Myrmarachne robusta (Peckham & Peckham, 1892). One new combination is proposed: Myrmarachne paludosa (Simon, 1900) comb. nov. Myrmarachne ramunni Narayan, 1915 is recorded for the first time in Sri Lanka. A total of 12 valid species are now known from the island; six of them are endemic. http://zoobank.org/urn:lsid:zoobank.org:pub:B723C180-996B-471D-B9204D08E7A8CD53 Keywords: taxonomy; India; Pakistan; Burma; Ceylon; biodiversity Introduction The ant-mimicking genus Myrmarachne MacLeay, 1839, is one of the most speciesrich genera of Salticidae, containing over 217 nominal species naturally distributed over five continents and numerous tropical islands (Edwards and Benjamin 2009; Edwards 2013; Platnick 2013; Prószyński 2013). They are specialized ant-mimicking salticids formerly assigned to the pluridentati group (Ceccarelli 2010). Results of the most recent molecular study place Myrmarachne into a new group called the Astioida, which include pluridents, unidents and fissidents (Maddison et al. 2008). They are primarily characterized by their ant-like appearance, an issue appropriately raised by Prószyński and Deeleman-Reinchold (2010). As ant mimicry is a rather common behavioural/morphological phenomenon throughout many Arthropod taxa (Mclver and Stonedahl 1993; Cushing 1997,2012), it alone cannot be used to define a large group of spiders (Prószyński and Deeleman-Reinchold 2010; Edwards 2013). The argument is then about the monophyly of Myrmarachne and its species groups. However, this issue could only be satisfactorily settled with the analysis of molecular phylogenetic data, independent of behaviour and morphology. *Email: [email protected] Journal of Natural History, 2015 Vol. 49, Nos. 43–44, 2609–2666, http://dx.doi.org/10.1080/00222933.2015.1034209 © 2015 Taylor & Francis Downloaded by [University of Cambridge] at 05:14 08 April 2016 Given the wide distribution and the large number of species involved, a stable taxonomy is highly desirable. The recent description of the type species of the genus, Myrmarachne melanocephala MacLeay, 1839, and phylogenetic analysis provide a firm basis for revisionary taxonomic studies of the genus (Edwards and Benjamin 2009). Nevertheless, its composition is still poorly understood, leading to disagreement of the placements of some species (Edwards and Benjamin 2009; Prószyński and DeelemanReinchold 2010;Edwards2013). Some regional Myrmarachne faunas have recently been fairly well documented: Africa (Wanless 1978), Australia (Ceccarelli 2010), India (Narayan 1915; Sherriffs 1931; Tikader 1973; Bradoo 1980; Sadana and Gupta 1998; Bastawade 2002), Pacific Islands (Berry et al. 1996), South East Asia (Edmunds and Prószyński 2003; Prószyński and Deeleman-Reinchold 2010;Yamasaki2012; Yamasaki and Ahmad 2013; Yamasaki and Edwards 2013), Seychelles (Saaristo 1978; Wanless 1983), Taiwan (Huang 2004) and Vietnam (Zabka 1985). Apart from a few papers, generally very little is known of the behaviour and life history of Myrmarachne. Although they generally tend to be found close to ants, they are known to actively avoid contact with them (Ceccarelli 2007). The behaviours of a handful of species have been observed (Mathew 1931,1940,1944;Wanless1978; Cushing 1997, 2012; Edmunds 2006;Borgesetal.2007;Sharma2012; Yamasaki and Edwards 2013). However, most observations centre on the common oriental species M. plataleoides. The standard work on Sri Lanka’sinvertebrate fauna was published as a part of the monographic series ‘Fauna of British India, including Ceylon and Burma’. However, it did not treat many smaller arachnids (Pocock 1900). This paper is part of an ongoing island-wide study to collect and record Sri Lanka’s invertebrate biodiversity. The end of the 30-year long civil conflict in Sri Lanka provides us an unparalleled opportunity to conduct island-wide biodiversity surveys and inventorying. Here I aim to survey and inventory the Myrmarachne fauna of Sri Lanka, describing all species of Myrmarachne from the island based on museum as well as newly collected material. Further, a key to all known species is provided. Material and methods Morphology Digital images were taken with a Nikon DXM1200F camera (Nikon, Tokyo, Japan). Images were edited using an AutoMontage software package (Syncroscopy, Cambridge, UK). Left structures are depicted unless otherwise stated. Setae are usually not depicted in the final palp drawings. All measurements are in millimetres and were made with a stereo microscope equipped with a 10× ocular and an ocular micrometer scale. An Amray 1810 housed at the Smithsonian Institution’s National Museum of Natural History Scanning Electron Microscope (SEM) facility was used to study and photograph morphological features. Targeted parts were cleaned ultrasonically for 1–3 min and dehydrated with 100% ethanol (transferred from 70% ethanol to absolute ethanol and left overnight), then critical point dried. After critical point drying, the specimens were glued to rounded aluminium rivets using an acetone solution of polyvinyl resin (Paraloid B72) and then Au/Pd coated for examination in the SEM. Female genitalia were excised using sharpened needles. Abdominal tissue was digested with SIGMA Pancreatin LP 1750 enzyme complex (Álvarez-Padilla and Hormiga 2007), in a solution of sodium borate prepared following methods described in Dingerkus and Uhler (1977). The specimen was then transferred to methyl 2610 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 salicylate (Holm 1979) and temporarily mounted as described in Grandjean (1949) and Coddington (1983) for examination and illustration under microscope. I have placed more weight on the use of illustrations (drawings, colour photos, SEMs) to convey the unique nature of each species, instead of the traditional method of words, as I think that a single well-composed illustration can convey more information on a species. Chelicerae dentition has been traditionally used in the identification of Myrmarachne species. However, my examination of specimens of Sri Lanka species shows that dentition patterns are variable and are thus, not used in their characterization. Abbreviations CD: copulatory duct; CO: copulatory opening(s); E: embolus; ED: ejaculatory duct (s); S: spermatheca; T: tegulum. Institutions CAS: California Academy of Sciences, San Francisco, CA, USA; MCZ: Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA; MHNG: Muséum d’Histoire Naturelle, Geneva, Switzerland; MNHN: Muséum National d’Histoire Naturelle, Paris, France; MZT: Zoological Museum, Department of Biology, University of Turku, Turku, Finland; NMSL: National Museum, Colombo, Sri Lanka; OUMNH: Hope Entomological Collections, Oxford University Museum, Oxford, UK; USNM: National Museum of Natural History, Smithsonian Institution, Washington, DC, USA; ZMB: Museum für Naturkunde der Humboldt-Universität, Berlin, Germany. Key to the Sri Lankan species of Myrmarachne 1. Tegulum of the male palp cylindrical, embolus is positioned on distal end of the tegulum, spermatheca kidney-shaped, connected by a CD to CO. Proximal half of chelicerae constricted. Generally not found in primary forest (Figures 30A–D,31A,35B,36C,37B, D) .................................................................... 2 –Tegulum spherical or disk-shaped, embolus is positioned on outer edges of the tegulum, spermatheca elongated tubes or oval. Generally found in primary forest (Figures 3A, F,4B,22A) ...................................................................... 3 2(1). Spiders brown in colour, large (7.2–30.0), Figure 29A, B ........ M. plataleoides –Spiders black in colour, much smaller (6.4–3.6), Figure 33A–D........ M. spissa 3(1). Spermatheca oval, connected by a CD to CO ................................................ 4 –Spermatheca an elongated tube, no clear distinction of CD ........................... 5 4(3). Adult spiders brownish with thoracic part black .................. M. melanocephala –Adult spiders black with thoracic part red/brown (Figures 23A–E,24A–D) ............................................................................................................. M. prava 5(3). Opisthosoma cylindrical, as much as 3× longer than wide, no constriction (Figures 5A–C,8G, H,9A–D)....................................................................... 6 –Opisthosoma oval or round, less than 2× longer than wide, constriction present in the centre of the anterior half of opisthosoma ........................................... 7 Journal of Natural History 2611 Downloaded by [University of Cambridge] at 05:14 08 April 2016 6(5). Adult spiders brownish with conspicuous white bands on prosoma and chelicerae (Figures 5A–C,8G, H) ..................................... M. dishani sp. n. –Adult spiders dark brown with no white bands on prosoma or chelicerae (Figure 9A–D) .............................................................................. M. imbellis 7(5). male palp longitudinally oval, embolus is positioned away from the borders of the tegulum (Figure 3A, F) .................................................................... 8 –Male palp circular, disk-shaped, embolus is positioned on the borders of the tegulum, small spiders (Figures 7B,22A) ................................................... 9 8(7). Chelicerae with parallel lateral sides, distal half slightly wider than the proximal half, RTA hook-shaped with a constriction at the base (Figures 2A, B, 3B,4D, E) ...................................................................................... M. bicurvata –Distal half of the chelicerae wider than the proximal half, The inner and outer sides of the proximal half are parallel, whereas the distal half has convex outer sides and parallel inner sides, RTA of the male palp is broad based and tapered to a point in one corner (Figures 2E, F,3E, F) ............. M. ramunni 9(7). Chelicerae fangs with a bulbous base, thickest at the centre (Figure 38A, B) ................................................................................................... M. aff. tristis –Chelicerae without a bulbous base, if base constricted, the entire proximal half is constricted ...................................................................................... 10 10(9). Chelicerae outer sides more or less parallel, dorsum curved, body covered with fine white hairs (Figures 14A–G,15A, B) ........ M. paludosa comb. nov. –Entire proximal or distal half of chelicerae constricted or dorsal surface flat (Figure 21A, B) ......................................................................................... 11 11(10). Proximal half of chelicerae constricted, distal half laterally rounded, dorsal and ventral surfaces well rounded forming a bulb (Figure 21A, B) ............... .................................................................................. M. morningside sp. nov. –Chelicerae proximal half not constricted, dorsal surface flat (Figure 1A, B) . .................................................................................... M. aurantiaca sp. nov. Taxonomy Family Salticidae Genus Myrmarachne MacLeay, 1839 Panachraesta Simon, 1900: 405. Type species by original designation P. paludosa Simon,1900: 405, examined. New synonymy. Remarks The type species of Myrmarachne, M. melanocephala was redescribed and a neotype designated by Edwards and Benjamin (2009). The type species of Panachraesta is here considered a typical member of the genus Myrmarachne; see redescription of the species below. Diagnosis and description for the genus is provided by Wanless (1978), 2612 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Edwards and Benjamin (2009), Prószyński and Deeleman-Reinchold (2010) and Yamasaki and Ahmad (2013). Myrmarachne aurantiaca sp. nov. (Figure 1A–D) Type material Holotype, male from Sri Lanka, North Western Province, Kurunegala District, Kurunegala, Ethagala range, 07°28ʹ17″N, 80°22ʹ30″E, 190 m, secondary forest, 20 September 2009, leg. Ziyard Jaleel et al. MHNG. Etymology The specific name refers to the red colouring of the prosoma. Diagnosis M. aurantiaca sp. nov. could be separated from other Sri Lankan Myrmarachne species by the body size, dorsally flat surface of the male chelicerae (Figure 1A, B) and reddish orange-coloured carapace. M. aurantiaca sp. nov. is very similar in size, habitat preference and genital morphology to M. paludosa comb. nov. and M. morningside sp. nov. However, individuals of both these species are dark black in colour (Figures 14A–G, 21A, B). Further, M. aurantiaca sp. nov. can be distinguished by the shape of the RTA which is strongly curved at the base: M. paludosa comb. nov. has a upward pointing, tapering RTA and M. morningside sp. nov. has a longer RTA with a curved tip. Description Male holotype. Total length: 3.5; prosoma length: 1.6, width: 0.8. Leg I: femur 0.9, patella 0.3, tibia 0.9, metatarsus 0.5, tarsus 0.3. Prosoma orange, opisthosoma anteriorly brownish, posterior half, black colour (Figure 1A, B). The cephalic part of the prosoma elevated, viewed dorsally lateral sides slightly rounded. The thoracic part is lower than the cephalic, the highest point being the centre (Figure 1A, B). Opisthosoma oval, slightly longer than wide, with no clear constriction, except for a white line (Figure 1); dorsum sclerotized, venter softer. Chelicerae are shorter than the carapace, with almost parallel lateral sides; distal half is a bit wider than the proximal half (Figure 1A, B). Chelicerae dentition not examined. Leg formula 4132. Leg spination not examined. Palps as in Figure 1C, D), with an RTA which is characteristically curved at its base. Female remains unknown. Distribution Known only from the type locality. Myrmarachne bicurvata (O. Pickard-Cambridge, 1869) (Figures 2A–D,3A–D,4A–F) Journal of Natural History 2613 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 1. Myrmarachne aurantiaca sp. nov., male holotype from Ethagala range (MHNG). (A,B) male dorsal view; (C,D) male palp (C, ventral D, retrolateral). Scale bars = 0.2 mm. 2614 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Salticus bicurvatus O. Pickard-Cambridge, 1869: 67, plate 6, figures 57–60. Myrmarachne bicrivata: Sherriffs, 1931: 539 (lapsus). Type material Holotype of Salticus bicurvatus: male from Sri Lanka (OUMNH 1735.2), no more data given, examined. Pickard-Cambridge (1869) says that the single specimen in OUMNH was captured in Sri Lanka by Mr G.H.K. Thwaits. Exact locality unknown. Diagnosis Very similar to M. ramunni; however, males are separated by the shape of the male chelicerae: with chelicerae in dorsal view, outer margins of chelicerae are weakly Figure 2. (A–D) Myrmarachne bicurvata. (A,B) male from Lahugala; (C,D) holotype, Sri Lanka, OUMNH. (E,F) Myrmarachne ramunni from Inginiyagala. (A,C,F) dorsal view; (B,D,E) retrolateral view. Journal of Natural History 2615 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 3. (A–D) Myrmarachne bicurvata. (A,B) male from Lahugala; (C,D) holotype, Sri Lanka (OUMNH); (E,F) Myrmarachne ramunni from Inginiyagala; (A,C,F) ventral view; (B,D,E) retrolateral view. Scale bars = 0.2 mm. 2616 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 convex in M. bicurvata, but distinctly convex in M. ramunni (Figure 2A, B). The RTA of the male palp in M. bicurvata is apically hooked and strongly curved at the base (Figure 3B–D). Description Male from Lahugala: total length: 5.6; prosoma length: 3.6, width: 1.2. Leg I: femur 1.6, patella 0.4, tibia 1.2, metatarsus 0.8, tarsus 0.4. The cephalic part of the prosoma is elevated and rounded on all sides. The thoracic part is lower than the cephalic, the highest point being the centre (Figure 2A–D). Opisthosoma oval, slightly longer than wide, with no clear constriction (Figure 2A–D). Opisthosoma dorsally sclerotized, Figure 4. Scanning electron micrographs of Myrmarachne bicurvata (USNM). (A–F) right male palp. (A) prolateral view; (B,F) ventral view; (C,D,E) retrolateral view. Scale bars = 10 µm (D,E) 20 µm (A,C) 30 µm (F) 40 µm (B). Journal of Natural History 2617 Downloaded by [University of Cambridge] at 05:14 08 April 2016 length: 2.8, width: 1.2. Leg 1: femur 1.2, patella 0.6, tibia 1.2, metatarsus 0.8 tarsus 0.4. Morphology as above. Leg formula 4132. Leg spination not examined.EpigynumandvulvaasinFigure 11A–C. CO rounded and S relatively longer (Figure 11A–C). Distribution Known only from Sri Lanka. Figure 9. Myrmarachne imbellis, male in life from Horton Plains, Sri Lanka (MHNG). 2624 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Other material examined Sri Lanka, Central province, 1 male, Kandaela Reservoir, 5.6 mi SW of Nuwara Eliya, 1889 m, 10–21 February 1970, Davis & Rowe, USNM. 1 male, Horton plains NP, Worlds-end, 2072 m, 21 February 1970, Davis & Rowe, USNM. 1 female, Nuwara Eliaya District, Path to Pitdurutalagalla, in light jungle, 17 November 1972–18 January 1973, P. Lehtinen, A. Withanage, & N. Wickremasinghe, MZT. 2 males, Horton Plains NP, 9 March 2000, SP. Benjamin, MHNG. 2 males 1 female, Nuwara Eliaya District, Horton Plains NP, 20–21 February 2007, SP. Benjamin and Z. Jaleel, MHNG. Myrmarachne melanocephala MacLeay, 1839 (Figures 17A–D,18A–D,19A–D) Myrmarachne orientales Tikader, 1973:60–62, figures 3–6. New synonymy. Synonymy M. orientales is well illustrated and is here compared with the neotype of M. melanocephala. I can find no characters that would suggest that the specimens Figure 10. Myrmarachne imbellis. Male palp. (A,B) from Kandaela (USNM); (C) Horton Plains (MHNG). (A) ventral view; (B,C) retrolateral view. Scale bars = 0.2 mm. Journal of Natural History 2625 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 11. Myrmarachne imbellis.(A–C) female from Piduruthalagala. (A) epigynum, ventral view; (B) vulva, ventral view; (C) vulva, dorsal view. Scale bars = 0.1 mm (B,C); 0.2 mm (A). 2626 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 from India illustrated by Tikader (1973) constitute a species different from M. melanocephala. Diagnosis Separated from other Sri Lankan Myrmarachne species by the dorsally flattened chelicerae, elongated prosoma with a constriction at the 50% point, anterior half rectangular and black in colour, posterior half oval and red/brown in colour. Figure 12. Scanning electron micrographs of Myrmarachne imbellis from Kandaela (USNM). (A–F) male palp. (A–C) retrolateral view; (D) prolateral view; (E,F) retrolateral view. Scale bars = 10 µm (B,C,F); 20 µm (A,D,E). Journal of Natural History 2627 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Opisthosoma elongated with a constriction at the posterior 25% margin, anterior 25% brown/black, posterior 75% black (Figure 17A–D). Further, the flattened hookshaped RTA and the elongated petioles are diagnostic (Figures 17A–D,19A–D). Similar to M. prava. However, can be easily separated from M. prava by the overall black colour of the male, very much shorter pedicel, slanted dorsal surface of the chelicerae, oval opisthosoma, and by the tapering, straight RTA in M. prava. Description See Edwards and Benjamin (2009). Natural history Edwards and Benjamin (2009) and Tikader (1973) report that this species mimics the ant Tetraponera allaborans (Walker, 1859). Distribution Sri Lanka, India, Pakistan to Indonesia (Edwards and Benjamin 2009). Material examined See Edwards and Benjamin (2009). Myrmarachne morningside sp. nov. (Figures 20A–D,21A–D,22A–D) Type material Holotype. male from Sri Lanka, Sabaragamuuwa Province, Ratnapura District, Eastern Sinharaja, Morningside section, 23 February 2007, SP. Benjamin and Z. Jaleel, MHNG. Paratype Male from Sri Lanka, same data as holotype. Etymology The specific name is a noun in apposition taken from the type locality. Diagnosis Very similar in outward appearance to M. spissa. Separated from it by the diskshaped tegulum and shiny appearance, mostly of the dorsal parts of the prosoma. In addition M. spissa has a pair of long whitish setae at about the centre of its prosoma, which are absent in M. morningside. Very similar in size, habitat 2628 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 preference and genital morphology to M. paludosa comb. nov. and M. aurantiaca sp. nov. M. paludosa comb. nov. has an upward pointing, tapering RTA and M. aurantiaca sp. nov. has an RTA with a curved base. It is predicted that females of this species would have an elongated S similar to other endemic Myrmarachne of the island. Description Male holotype from Morningside: total length: 5.6; prosoma length: 3.5, width: 1.0. Leg I: femur 1.1, patella 0.4, tibia 1.1, metatarsus 1.0, tarsus 0.4. Prosoma oval, 1.5× long than wide. Opisthosoma oval, 2× longer than wide, constricted at the centre of the anterior half. Uniformly black in colour, legs a lighter yellow-brown. Prosoma and opisthosoma are both covered with fine whitish hairs. Chelicerae, somewhat longer than carapace, anteriorly bulb-like, tapering towards the base, dentition not examined. Leg formula 4132. Leg spination not examined. Male palps as in Figures 21C, D,22A–D. Female unknown. Distribution Endemic to Sri Lanka. Natural history Collected together with ants of the genus Camponotus –probably the same species collected with M. paludosa comb. nov. However, it is not the Camponotus species found with M. prava. Other material examined Sri Lanka, Sabaragamuwa province, 1 male, Ratnapura District, Gilimale Forest Reserve, 23 February 2007, SP. Benjamin and Z. Jaleel, MHNG. Myrmarachne paludosa (Simon, 1900) comb. nov. (Figures 13A, B,14A–G,15A–H,16A–D) Panachraesta paludosa Simon, 1900: 405 Type material Syntypes of Panachraesta paludosa: 6 females and 4 juveniles, Sri Lanka, Colombo, no more data given (MNHN 20511/2306). Examined. Diagnosis Myrmarachne paludosa (Simon, 1900) comb. nov. could be separated from other Sri Lankan Myrmarachne species by the shape of the chelicerae: proximal half of Journal of Natural History 2629 Downloaded by [University of Cambridge] at 05:14 08 April 2016 chelicerae not constricted, outer sides more or less parallel, curved dorsum. Further, the black coloured body covered with fine white hairs (Figures 14A–G,15A, B) and the upward pointing, tapering RTA is diagnostic. Females may be separated by the elongated S that partly surrounds CO, connecting to it at the bottom margins (Figure 15E–H). Figure 13. Myrmarachne paludosa (Simon, 1900) comb. nov. Female syntype (MNHN 20511/ 2306). (A) dorsal view; (B) epigynum, dorsal view. Scale bars = 0.2 mm (B); 2.0 mm (A). 2630 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 14. Myrmarachne paludosa (Simon, 1900) comb. nov. In life. (A,D,E) male; (B,C,F,G) female. Journal of Natural History 2631 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 15. Myrmarachne paludosa (Simon, 1900) comb. nov. (A,B) male from Kandy. (C,D) (E–G) female from Kitulgala. (H) female from Corbetts Gap. (A,C) dorsal view; (B,D) lateral view; (E,H) epigynum, ventral view; (F) vulva, dorsal view; (G) vulva, ventral view. Scale bars = 0.1 mm (F,G); 0.2 mm (E,H). 2632 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 16. Myrmarachne paludosa (Simon, 1900) comb. nov. (A,B) male from Kandy. (C,D) male from Gilimale. (A) ventral view; (B,C) retrolateral view; (D) dorsal view. Scale bars = 0.2 mm. Journal of Natural History 2633 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 22. Myrmarachne morningside sp. nov. (A–D) male palp. (A) ventral view; (B–D) retrolateral view. Scale bars = 10 µm (D); 100 µm (A,B,C). 2640 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 23. Myrmarachne prava in life. (A–D) male from Nikaravatiya; (E) juvenile. Journal of Natural History 2641 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Diagnosis Separated from other Sri Lankan Myrmarachne species by the shape of the dorsal surface of the chelicerae, which slant towards the median axis. Further, the oval opisthosoma which lack any constrictions and the tapering, upward pointed RTA. Females could be separated by the oval CO tipped towards the median axis (the anterior ends are closer to the median axis than the posterior ends) and the elongated S which are parallel to the median axis. Description Male (from Anuradapura): total length: 8.0; prosoma length: 5.2, width: 1.4. Leg I: femur 2.0, patella 0.8, tibia 1.9, metatarsus 1.3, tarsus 0.6. Coloration as in Figures 23A–D,25A–D. Prosoma black with evenly distributed white hair, chelicerae reddish. Opisthosoma grey with white hair. The cephalic part of the prosoma is elevated, sides somewhat curved. Prosoma constricted in the centre. The thoracic part is lower than the cephalic part, oval in shape (dorsal view) and slopes to towards the base. Figure 24. Myrmarachne prava in life. (A–D) female from Nikaravatiya. 2642 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Opisthosoma oval, 1.5× longer than wide. Chelicerae shorter than prosoma, dorsum slants towards the median axis, lateral sides rounded (Figure 25A–D). Chelicerae dentition not examined. Leg formula 4132. Leg spination not examined. Palps as in Figure 26A–D. Female: total length: 6.4–7.2; prosoma length: 2.8, width: 1.2. Leg 1: femur 1.6, patella 0.6, tibia 1.2, metatarsus 0.8 tarsus 0.4. Morphology as above, except for the unmodified chelicerae. Coloration as in Figures 23E,24A–D. Leg formula 4132. Leg spination not examined. Epigynum and vulva as in Figure 27A–C. The oval CO tipped towards the median axis (the anterior ends are closer to the median axis than the posterior ends) and the elongated S which are parallel to the median axis are characteristic for this species. Natural history The specimens from Nikaravatiya were collected along with a species of ants of the genus Camponotus. Both the spider and its ant host are illustrated in two beautiful hand drawings by G.M. Henry (2000). Borges et al. (2007) associated the ant Camponotus compressus (Fabricius) with this species (identified as Myrmarachne morpho species 2). However, Tikader (1973) reports that Myrmarachne bengalensis was collected along with the ant Leptogenys processionalis (Jerdon, 1851), which could be a misidentification. Distribution India, Sri Lanka. Figure 25. Myrmarachne prava. (A,B) male holotype (ZMB 1536). (C,D) male from Padaviya. (A,C) dorsal view; (B,D) lateral view. Journal of Natural History 2643 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Other material examined Sri Lanka, North Central Province, Anuradapura District, Padaviya, 1 female 2 juv, 18 March 1976, label gives coll. ‘KV KROMBEIN?’, USNM. 2 males and 1 juvenile, Same data, 19 March 1976, M. Trad. USNM. 24 October 1997, no other Figure 26. Myrmarachne prava, male palp. (A,B) holotype (ZMB 1536). (C,D) male from Padaviya. (A,C) ventral view; (B,D) retrolateral view. Scale bars = 0.2 mm. 2644 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 data, Saman, USNM. Western Province, 2 males, Colombo District, Nawala, 13 March 1998, M.M. Bahir and S.V. Nanayakkara, MHNG. 1 male, Maharagama, 24 July 2009, S.P. Benjamin, MHNG. North Western province, 1 male 1 female, Figure 27. Myrmarachne prava, female from Padaviya. (A) epigynum, dorsal view; (B) vulva, dorsal view; (C) vulva, ventral view. Scale bars = 0.1 mm (B,C); 0.2 mm (A). Journal of Natural History 2645 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Kurunagala District, Nikaravatiya, 1–3. February 2007, Z. Jaleel, USNM (photo voucher). Myrmarachne plataleoides (O. Pickard-Cambridge, 1869) (Figures 29A–D,30A–D,31A–E) Figure 28. Scanning electron micrographs of Myrmarachne prava.(A–E) male palp. (A) prolateral view; (B,E) ventral view; (C,D) retrolateral view. Scale bars = 10 µm (D,E); 30 µm (C); 100 µm (A,B). 2646 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Salticus plataleoides O. Pickard-Cambridge, 1869: 68, plate 6, figures 61–65a. See Platnick (2013) for a detailed taxonomic history. Type material The type specimen was not found, presumed lost. Pickard-Cambridge (1869) says that the single specimen that he described was found in the collections of OUMNH. Type locality was unknown at the time of description. Material examined Sri Lanka, Central Province, 1 male, Kandy District, Udawattakelle sanctuary, 510–580 m, 8–10 September 1977, K.V. Krombein, P.B. Karunaratne, T. Wijesinhe, M. Jayaweere, USNM. Sabaragamuwa Province, 1 male, Kegalle District, Kelani Ganga, 152 m, Kitulgala, 12 March 1973, Bauman and Cross, USNM. Western Province, 1 male 1 female, Colombo District, Godagama, 25 October 1973, M. Robinson, USNM. 1 male, Ratmalana, 6 June 1976, KV. Krombein, USNM. 1 female, Nawala, Nandimitra place, 13 March 1998, M.M. Bahir and S.V. Nanayakara. 1 female, same locality, 1 March 1998, MHNG. 2 males, Kalutara District, Ingiriya, Bodinagala Forest Reserve, 10 February 2007, Hand collecting, SP. Benjamin and Z. Jaleel. North Central Province, 1 female, Anuradapura District, Padaviya, 18 May 1976, M. Trap and K.V. Krombein, USNM. 1 male, same locality, 19 May 1976, K.V. Krombein (?), USNM. Figure 29. Myrmarachne plataleoides in life. (A,C) male; (B,D) female. Journal of Natural History 2647 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 30. Myrmarachne plataleoides. (A,B) male palp. (A) ventral view; (B) retrolateral view; (C) vulva, ventral view; (D) vulva, dorsal view. Scale bars = 0.1 mm (C,D); 0.2 mm (A,B). 2648 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 31. Scanning electron micrographs of Myrmarachne plataleoides.(A–E) male palp. (A,E) ventral view; (B–D) retrolateral view. Scale bars = 20 µm (B); 25 µm (C,E); 30 µm (A,D). Journal of Natural History 2649 Downloaded by [University of Cambridge] at 05:14 08 April 2016 examined. Male palps with a cylindrical tegulum, with an embolus positioned on the distal end of it as in Figure 35A–F. Female from Inginiagala: total length: 3.6; prosoma length: 1.6, width: 0.9. Leg 1: femur 0.8, patella 0.3, tibia 0.6, metatarsus 0.4 tarsus 0.2. Morphology as above, except for the lighter coloured prosoma and opisthosoma (Figure 32A–J). Some females seem to have reddish-brown chelicerae (Figure 34). Chelicerae dentition not examined. Leg formula 4132. Leg spination not examined. Epigynum and vulva as in Figure 36A–C. Distribution Sri Lanka. Variation The Monaragala District specimen is a bit larger than the specimen from Padaviya. Some recently collected specimens are darker, the light colour of the types might be due to preservation. Natural history Specimens from Atidiya live in association with a species of ants of the genus Technomyremex (vouchers deposited in MHNG). Specimens from Kurunagala, Nikaravatiya were collected with ants of the genus Cataulacus. Other material examined Sri Lanka, Uva Province, 1 male, Monaragala District, Angunakolaplesse 100 m, 21–23 January 1979, K.V. Krombein, P.B. Karunaratne, T. Wijesinhe, S. Figure 36. Myrmarachne spissa.(A–C) female from Bellanwila-Attidiya. (A) epigynum, ventral view; (B) vulva, dorsal view; (C) vulva, ventral view. Scale bars = 0.1 mm. 2656 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Siriwardane, T. Gunawardane, USNM. 1 male 1 female, Monaragala District (probably Ampara District), Inginiagala, 3 June 1995, D.H. Messersmith, G.L. Williams and P.B. Karunaratne, USNM. North Central Province, 1 male. Anuradapura District, Padaviya, 19 May 1976. Leg K.V. KROMBEIN(?), USNM. Eastern Province, 1 male, Ampara District, Inginiyagala, Gal Oya NP, Figure 37. Scanning electron micrographs of Myrmarachne spissa from Padaviya. (A–F) right male palp. (A) prolateral view; (B,D–F) ventral view; (C) retrolateral view. Scale bars = 10 µm (F); 20 µm (A–E). Journal of Natural History 2657 Downloaded by [University of Cambridge] at 05:14 08 April 2016 c.80 m, 07°13ʹ22″N, 81°31ʹ56″E,10February2010,leg.S.P.Benjamin, S. Batuwita, MHNG. Western Province, 1 male 1 female, Colombo District, Bellanwila-Attidiya, 21 February 2000, S.P. Benjamin. 5 males, Panadura, Mahabellana, along Bolgoda South Lake, July 2008, S.P. Benjamin, S.K. Dayananda and Z. Jaleel, MHNG. North Western Province, 11 males, 5 females, Kurunagala District, Kurunagala, Ethagala Mountains, c.300 m, 1–28 February 2007, Z. Jaleel, MHNG. 2 males, 11 October 2008, Z. Jaleel, MHNG. 6 males, 1–30 March 2008, leg. Z. Jaleel, MHNG. 2 females, Ethagala range, c.190 m, 07° 28ʹ17″N, 80°22ʹ30″E, 24 November 2009, Hand collecting, leg. S.P. Benjamin, S. Batuwita, Z. Jaleel, MHNG. Central Province, 1 male, 2 females, Kandy District, Dunumadalawa, c.600 m, 07°17ʹ00″N, 80°37ʹ49″E, 24 September 2009, leg. S.P. Benjamin et al., MHNG. 1 male, 1 female, Matale District, IFS Arboretum, c.180 m, 07°51ʹ34″N, 80°40ʹ28″E, 24 August 2010, Hand collecting, leg. S.P. Benjamin, S. Batuwita. 1 male ‘no label’USNM Sri Lanka collection (Sri Lanka?), MHNG. Myrmarachne aff. tristis (Simon, 1882) (Figure 38A–D) Salticus tristis Simon, 1882: 212. Remarks Provisional identification is based on the description of the tristis-group by Wanless (1978). Diagnosis Small spiders, separated from all other Sri Lankan species of the genus by the chelicerae: shorter than the prosoma, fangs with a bulbous base (Figure 38A, B). Further, the disk-shaped male palp, with the embolus positioned on the borders of the tegulum (Figure 38C, D) is characteristic. Male. total length: 5.6; prosoma length: 3.2, width: 1.2. Leg I: femur 1.2, patella 0.4, tibia 1.2, metatarsus 0.6, tarsus 0.4. Prosoma oval, almost as wide as long. The cephalic part of the prosoma is elevated and sides parallel. Posterior half rounded. Prosoma laterally with dark markings (Figure 38A, B). All eyes surrounded by dark rings. Opisthosoma oval, longer than wide, with a clear constriction in the centre of the anterior half (Figure 38A, B). Dorsally, sclerotized, ventrally softer. Chelicerae constricted at the base, thickest at the centre (Figure 38A, B). Male palps disk-shaped, with the embolus positioned on the borders of the tegulum (Figure 38C, D). Chelicerae dentition not examined. Leg formula 4132. Leg spination not examined. Females have not been found from Sri Lanka. 2658 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Distribution Libya to India. New record for Sri Lanka. Material examined Sri Lanka: Kimbissa: 1 male, 0.8 km west of Sigiriya Jungle, 28 June–4 July 1978, Coll. K.V. Krombein, P.B. Karunaratne, J. Wijesinghe, V. Kulasekara, USNM. Figure 38. Myrmarachne aff. tristis (Simon, 1882). (A,B) Photographs of male from Kimbissa; (C,D) left male palp. (A) dorsal view; (B) lateral view; (C) ventral view; (D) retrolateral view. Scale bars = 0.2 mm. Journal of Natural History 2659 Downloaded by [University of Cambridge] at 05:14 08 April 2016 List of comparative material examined Myrmarachne nemorensis (Peckham and Peckham, 1892) (Figure 39A, B) Holotype of Salticus nemorensis from Burma (Myanmar, MCZ 22264). The holotype is illustrated here (Figure 39A, B). Myrmarachne opaca (Karsch, 1880) (Figure 39C–F) Material examined Syntype of Synemosyna opaca from Philippines, Samar Province, 1 male (ZMB 1530), leg. Jager. Type specimen is illustrated here (Figure 39C–F). A description of the female, diagnosis and natural history is provide by Yamasaki and Ahmad (2013). Distribution Borneo, Philippines. Myrmarachne pumilio (Karsch, 1880) (Figure 40A–E) Material examined Syntype of Synemosyna pumilio from Bengal (West Bengal State of India & Bangladesh), 1 male (ZMB 1533), leg. Nietner. Examined. Figure 40A–E. Myrmarachne hanoii Zabka, 1985: 419, figures 364–367. New synonymy. Distribution Bangladesh, Borneo, China, India, Indonesia, Vietnam. Remarks This species has been redescribed with several detailed illustrations by Yamasaki and Ahmad (2013). The general shape of the chelicerae and palp are very characteristic. The papal tibia with a multi-pronged RTA is diagnostic. I thus consider Myrmarachne hanoii Zabka, 1985 a junior subjective synonym of Myrmarachne pumilio. A description of the female, diagnosis and natural history is given in Yamasaki and Ahmad (2013). Myrmarachne robusta (Peckham and Peckham, 1892) (Figure 41A–B) 2660 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 39. (A,B) Myrmarachne nemorensis, male holotype (MCZ 22264). (C–F) Myrmarachne opaca, male syntype (ZMB 1530). (A,D) retrolateral view; (B,C) ventral view; (E) dorsal view; (F) lateral view. Scale bars = 0.2 mm. Journal of Natural History 2661 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Figure 40. Myrmarachne pumilio.(A–E) male syntype (ZMB 1533). (A) ventral view; (B) retrolateral view; (C,D) dorsal view; (E) lateral view. Scale bars = 0.2 mm. 2662 S.P. Benjamin Downloaded by [University of Cambridge] at 05:14 08 April 2016 Material examined Holotype of Salticus robustus from Burma (Myanmar, MCZ 22915), 1 male, Figure 41A, B.Myrmarachne maratha Tikader, 1973 New synonymy. Described from Myanmar; distribution now extended to India. Figure 41. Myrmarachne robusta. (A,B) right male palp of holotype (MCZ 22915). (A) ventral view; (B) retrolateral view. Scale bars = 0.2 mm. Journal of Natural History 2663 Downloaded by [University of Cambridge] at 05:14 08 April 2016 Distribution India, Myanmar. Remarks The description of M. maratha fits the holotype of M. robusta. I thus consider the former a junior subjective synonym of the latter. Acknowledgements This study was funded by the Institute of Fundamental Studies with additional funding coming from a fellowship from the Alexander von Humboldt Foundation. I wish to thank the Department of Wildlife Conservation and the Department of Forest Conservation of Sri Lanka for facilitating fieldwork. Thanks to Ted R. Schultz (USNM) for identifying the associated ants. Thanks to Jonathan Coddington and Dana De Roche (USNM), James E. Hogan and Darren J. Mann (OUMNH), Chistine Rollard and Elise-Anne Leguin (MNHN), Jason A. Dunlop and Shahin Nawai (MNHU), Peter Schwendinger (MHNG), Gonzalo Giribet and Laura Leibensperger (MCZ) and Seppo Koponen (MZT) for providing access to collections under their care. Thanks to Namal Atukorala, Sudesh Batuwita, S.K. Dayananda, Ziyard Jaleel and Harshini Sandamali (all of IFS) for assistance in the field. G.B. Edwards and Takeshi Yamasaki provided literature. Bernhard Huber acted as my host and helped in various ways during my stay in Bonn as a Humboldt fellow. I thank G.B. Edwards and an anonymous reviewer for helpful comments. ORCID Suresh P. Benjamin http://orcid.org/0000-0003-4666-0330 References Álvarez-Padilla F, Hormiga G. 2007. A protocol for digesting internal soft tissues and mounting spiders for scanning electron microscopy. J Arachnol. 35:538–542. Bastawade DB. 2002. Three new species from the spider families Amaurobiidae. Thomisidae and Salticidae (Araneae: Arachnida) from India. J Bombay Nat Hist Soc. 92:274–281. Berry JW, Beatty JA, Proszynski J. 1996. 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Occas Pap Natur Hist Soc Wisconsin. 2:1–83. Pickard-Cambridge O. 1869. Descriptions and sketches of some new species of Araneidea, with characters of a new genus. J Natur Hist Ser 4. 3:52–74. Platnick NI 2013. The World Spiders Catalog, Version 13.5. New York: American Museum of Natural History. Available from: http://research.amnh.org/entomology/spiders/catalog8187/index.html Pocock RI. 1900. The Fauna of British India, including Ceylon and Burma. Arachnida. I–XII:1–279. Pollard SD. 1994. Consequences of sexual selection on feeding in male jumping spiders (Araneae: Salticidae). J Zool. 234:203–208. Prószyński J. 2013. Monograph of Salticidae (Araneae) of the World. 1995–2012 [Internet]. March 1st, 2006 Warszawa. Available from: http://www.peckhamia.com/salticidae/ Journal of Natural History 2665 Downloaded by [University of Cambridge] at 05:14 08 April 2016