Bavia aericeps Simon, 1877

Bavia capistrata (C. L. Koch, 1846), combination restored, removed from synonymy with Evarcha flavocincta (C. L. Koch, 1846)

Bavia fedor Berry, Beatty & Prószyński, 1997

Bavia nessagyna Maddison, sp. nov.

Bavia gabrieli Barrion, 2000

Bavia intermedia (Karsch, 1880)

Bavia maurerae (Freudenschuss & Seiter, 2016), comb. nov., transferred from Epidelaxia

Bavia planiceps (Karsch, 1880)

Bavia sexpunctata (Doleschall, 1859)

Bavia valida (Keyserling, 1882)

Larger-bodied than most other baviines. Carapace relatively broad and having hexagonal shape, widest at or just behind the PLEs (Fig. 13). Chelicerae lack the sharp lateral ridge (Fig. 4) of Indopadilla. Embolus shorter than length of tegulum, arising in all known species on bulb’s distal prolateral corner. ECP on a prominent medial bulge. Male endite with small thumb-like lobe laterally (Fig. 6), as in the Padillothorax badut group.

Illustrations are given here of some of the well-known species of Bavia, including B. aericeps (Figs 36–41) and B. sexpunctata (Figs 54–63). B. capistrata was synonymized without explanation by Prószyński (2017) with the extremely different Evarcha flavocincta, possibly because of superficial similarities in the palp. C. L. Koch’s (1846) illustration of the male of Maevia capistrata is clearly a Bavia by body form and markings. The only doubt about the status of C. L. Koch’s species is which species of Bavia is it precisely. Candidates include the one figured by Cao, Li, and Żabka (2016), the one figured here as B. cf. capistrata (Figs 42–53), and B. nessagyna (Figs 64–75). The one figured here as B. cf. capistrata could be different from that figured by Cao, Li, and Żabka. The former shows a slightly wider embolus and more delicate ECP, and possibly more contrasting markings. Regardless, C. L. Koch’s species is removed from synonymy with Evarcha flavocincta and returned to Bavia.

A video of the living female B. cf. capistrata (specimen AS19.1128) is available in Maddison (2020).

In addition to the species below, we have seen an undescribed species near B. nessagyna from Mulu National Park (single female) and a species near B. intermedia (single male, here represented as specimen d079 in the Sanger data).

All from Malaysia: Sarawak: Lambir Hills Nat. Pk., and in UBCZ. Holotype: male IDWM.20005 from Bukit Pantu Trail, 4.2032°N, 114.0305°E to 4.2035°N, 114.0304°E, 210 m el. 5 April 2012 Maddison/Piascik/Ang WPM#12-135. Paratypes: Male SWK12-4726 from Inoue Trail, 4.2002°N, 114.0346°N, to 4.2004°N, 114.0342°E, 200 m el. 4 April 2012 Maddison/Piascik/Ang WPM#12-130; female SWK12-4087 from Inoue Trail, 4.1990°N, 114.0375°E to 4.1988°N, 114.0370°E, 120 m el. 1 April 2012 Maddison/Piascik WPM#12-113; female IDWM.20004 from Lepoh-Ridan Trail, 4.2022°N, 114.0279°E to 4.2019°N, 114.0278°E 170 m el. 2 April 2012 Maddison/Piascik WPM#12-124; female IDWM.20006 from Bukit Pantu Trail, 4.2028°N, 114.0305°E to 4.2032°N, 114.0305°E, 210 m el. 5 April 2012 Maddison/Piascik/Ang WPM#12-134; female IDWM.20003 from Pantu Trail, 4.2030°N, 114.0399°E to 4.2032°N, 114.0396°E, 150–160 m el. 6 April 2012 Piascik/Ang WPM#12-145.

From the Greek nessa, duck, and gyne, female, referring to the resemblance of the epigyne to a duck’s bill. Other names: In WPM’s lab notebooks the informal code for this species was “BVDUC-S”.

One of the more delicate Bavia, along with B. capistrata, having legs II–IV very much paler than I, and thus resembling Indopadilla. Its distinction from B. fedor is slight: the embolus of both appears as a curved and narrowing blade with a series of retrolateral teeth. The teeth are short, triangular, and closely spaced in B. nessagyna, but larger in B. fedor, appearing as broad pillars whose bases are well separate (photographs of holotype kindly supplied by J. Boone, Bishop Museum). In B. nessagyna the teeth are not on the embolus proper but on a TmA that parallels the embolus (Fig. 64). In B. fedor it is unclear whether the teeth are on a TmA or on the embolus itself; Prószyński’s illustration (Berry, Beatty and Prószyński 1997) shows no division into two processes, but it may be that they are closely adpressed. Even still, the overall shapes of the embolic division differ: broad at the base in B. nessagyna but abruptly narrowing; narrower at the base in B. fedor and narrowing more gradually. Epigyne with arcing ridges lateral to the openings in B. fedor; without such a ridge in B. nessagyna. B. nessagyna differs from B. cf. capistrata in the distinct embolus, less contrasting markings, and the abdominal markings more transverse than longitudinal.

Male (based on holotype, specimen IDWM.20005). Carapace length 3.6; abdomen length 4.8. Carapace (Fig. 71): Broad, seeming even broader because of small tuft of setae at widest point just posterior to PLEs. Integument of ocular area black, thorax dark brown, except in alcohol pale orange yellow area just behind the ocular area. Bands of cream coloured scales lie just posterior to and lateral to the ocular area; thorax with a few small spots of cream scales, the two posterior ones of which are prominent and well separated, as in B. capistrata. Clypeus dark and glabrous, but a disorderly fringe of long cream setae overhangs the chelicerae (Fig. 70). Chelicerae simple and mostly vertical, dark, with cream setae that, with those of the clypeus, give the appearance of an unkempt moustache. At least two teeth on retromargin (paratype with 6). Palp as in Fig. 64; embolus thin and accompanied by toothed TmA. Femur dark basally, but terminally is pale, as are all more distal segments. Endites with thumb-like lobe laterally (Fig. 6). Legs II–IV notably paler than legs I, which bear annulae of white scales on patella and distally on tibia. Metatarsus I dark; tarsus pale. Ventral fringe of black setae beneath tibia and metatarsus I. Abdomen with dorsal markings primarily transverse, with four transverse pale bands separated by three dark bands.

Female (based on paratype IDWM.20003). Carapace length 3.3; abdomen length 5.9. Carapace as in male, but lacking tuft at widest point. Clypeus and Chelicerae as in male. Five retromarginal teeth. Legs similar to those of male, but with first leg not quite so dark, and without the ventral fringe so distinctly developed. Abdomen with only a hint of transverse bands, instead dominated by longitudinal stripes: a narrow central pale band flanked by wide dark bands. Epigyne with ECP beneath a posteriorly-projecting mound (Figs 66–67).

All specimens from Lambir Hills were collected on big-leaved foliage (e.g., palms) except for IDWM.20004 whose collecting record says simply “foliage”.

One female from Malaysia: Sarawak: Lambir Hills Nat. Pk., Inoue Trail, 4.2002°N, 114.0346°E to 4.2004°N, 114.0342°E, 200 m el. 4 April 2012 Maddison/Piascik/Ang WPM#12-130. One male from Malaysia: Sarawak: Bako Nat. Pk., Mangroves, beach forest, 1.722°N, 110.446°E, 0 m el. 8 March 2012 Maddison/Piascik/Ang/Lee WPM#12-003.

Indopadilla annamita (Simon, 1903), comb. nov., transferred from Bavia

Indopadilla bamilin Maddison, sp. nov.

Indopadilla darjeeling Caleb & Sankaran, 2019

Indopadilla kahariana (Prószyński & Deeleman-Reinhold, 2013), comb. nov., transferred from Bavia

Indopadilla kodagura Maddison, sp. nov.

Indopadilla insularis (Malamel, Sankaran & Sebastian, 2015)

Indopadilla nesinor Maddison, sp. nov.

Indopadilla redunca Maddison, sp. nov.

Indopadilla redynis Maddison, sp. nov.

Indopadilla sabivia Maddison, sp. nov.

Indopadilla sonsorol (Berry, Beatty & Prószyński, 1997), comb. nov., transferred from Bavia

Indopadilla suhartoi (Prószyński & Deeleman-Reinhold, 2013), comb. nov., transferred from Bavia

Indopadilla thorelli (Simon, 1901)

Indopadilla vimedaba Maddison, sp. nov.

Front face of chelicerae concave or flat in both males and females, bordered laterally by a sharp ridge (Fig. 5). The ridge projects laterally in some males (e.g., Fig. 98, triangle, or see the male of I. vimedaba figured by Prószyński 1987, p. 105, as “Stagetilus semiostrinus”). (Males of Bavia may have chelicerae with a flat front face, and males of Padillothorax may have the front face concave medially, but none have such a sharp or extended lateral ridge.) Clypeus very narrow at centre, exposing a broad expanse of arthrodial membrane that in most species is white (Figs 96, 115, 128; Żabka 1988; Prószyński and Deeleman-Reinhold 2013 fig. 12; Malamel et al. 2015 fig. 9; Caleb et al. 2019 figs. 4, 21; dark in I. kahariana, I. darjeeling, I. bamilin). Spermatheca anterior to copulatory openings (posterior in other baviines). Retromarginal teeth of male not clustered onto a raised hump; rather, the teeth occur on a long sharp ridge that extends medially from base of fang (e.g., Prószyński 1987, p. 105). Male endite rounded, lacking lobe or sharp corner (Fig. 7). Thorax with a bulge just behind the PLE that gives the carapace a “muscular” appearance (Figs 5, 14; see triangle on Fig. 14). (This area contains the muscle attachments for the palps, first legs, and possibly the second legs.) Abdominal markings varied but include a pale longitudinal stripe along each side that is broken approx. half way toward the back, followed by a more posterior pale spot that extends slightly dorsally (Figs 79, 99, 103). When the embolus is long (e.g., Fig. 93), it arises simply as an extension of the tegulum, not being clearly divided from the tegulum as in Piranthus and Maripanthus.

This distinctive group may have many dozens or even hundreds of species, judging from the rate of discovery of new species among the few specimens being collected. We show in Figs 129–142 some of the apparently-new species that we are not naming. Matching males and females with such sparse collecting can be difficult. We have co-collected both males and females of I. kahariana, and they fall together on the molecular phylogeny (Fig. 3). Among the new species, only two are represented by both sexes, I. redunca and I. vimedaba. We interpret the male I. bamilin, I. kodagura, and I. sabivia as not being matches for the female I. nesinor because they are distinct on the molecular phylogeny or have mismatching genitalia (length of embolus vs. copulatory ducts; elaboration of RTA vs. ECP).

Embolus length varies through Indopadilla, short in many species, in other species (e.g., I. kodagura, I. suhartoi) as long as in some Piranthus and Marapathus. While we might be tempted to split the group into two – Indopadilla with a long embolus in South Asia and southeast Asia, and a new genus with a short embolus in southeast Asia – the most complete molecular data nests the long-embolus I. kodagura among short embolus species (Fig. 1). Also, even if the long and short embolus species formed mutually monophyletic groups, dividing them would break one very distinctive and easily recognized clade (cheliceral ridge, thoracic bulges, wide arthrodial membrane on face, etc.) into two much more subtle groups. We therefore maintain the whole group as a single distinctive genus.

The peculiar carapace bulge and exposed arthrodial membrane on the clypeus hint to the possibility that Indopadilla may use unusual biomechanics. The third leg claw tufts appear noticeably larger than the others. Indopadilla are excellent jumpers, difficult to collect even on a beating sheet, from which they can escape in a single decisive bound. They are usually collected from foliage. A video of living males of I. kodagura and the undescribed species “BVBTN” (Figs 134–136) is available in Maddison (2020).

Holotype male (specimen SWK12-1618, in UBCZ) from Malaysia: Sarawak: Mulu Nat. Pk., Clearwater Cave Trail, 4.0597°N, 114.8292°E to 4.0592°N, 114.8291°E, 60 m el. 14 March 2012 Maddison/Piascik/Ang WPM#12-027.

An arbitrary combination of letters, ungendered. Other names: In Maddison (2015b) and WPM’s lab notebooks the informal code for this species was “BVBML”.

Both embolus and RTA short and simple (Fig. 76). Other species with such a short embolus have a distinct TmA (I. annamita, I. kahariana, I. redunca, I. sabivia), which seems lacking in I. bamilin. First legs and body dark. None of the known females is an obvious match for this male in terms of markings or molecular phylogenetic placement, but also the unmatched females have longer copulatory ducts than would be expected for such a short embolus, or have a more dramatic ECP than expected for the simple RTA.

Male (based on holotype, specimen SWK12-1618). Carapace length 2.3; abdomen length 3.0. Carapace dark brown, slightly paler around fovea, with a few patches of pale scales on thorax (Figs 78, 79). Clypeus dark, narrow; status of arthrodial membrane unclear, as chelicerae are somewhat sunken into the prosoma. Chelicerae vertical, brown, concave in front and with lateral ridge. Retromarginal teeth at least three, on long ridge. Palp femur dark; more distal segments pale. Embolus short, undivided (i.e., no TmA; Fig. 76). Tibia with dorsal lobe distally (Fig. 77). Endite rounded laterally, without corner or lobe. Legs strongly contrasting between black leg I and pale legs II–IV. First patella entirely black. Abdomen dark red in life, with longitudinal band of pale scales on each side broken as typical for the genus.

We include illustrations of the described Indopadilla kahariana, to show living specimens and to show the genitalia in detail. Christa Deeleman-Reinhold kindly compared the type specimens to our illustrations and confirmed the match. A photograph of a living female is shown by Koh and Bay (2019, page 203, fig. A). In WPM’s field and lab notebooks the informal code for this species was “BVSMB”.

Holotype male (NCBS-BN351, also known as AS19.4314), in NCBS collection, from India: Karnataka: Kodagu: Yavakapadi, Honey Valley area, forest & edge, 12.215°N, 75.659°E to 12.216°N, 75.661°E, 1300 m elev. 25 June 2019 W. Maddison & K. Marathe WPM#19-077.

In the Kodava language, kodagura means from Kodagu. Other names: In WPM’s field or lab notebooks the informal code for this species was “BVHVW”.

Very similar to I. insularis, contrastingly marked in dark brown and yellow, with the face appearing white because the clypeus is withdrawn toward the eyes to expose a bright white arthrodial membrane. Like I. insularis, I. darjeeling, I. sonsorol, I. suhartoi, and I. thorelli in having a long thin embolus, but even longer than in those species, arising from the retrolateral basal corner of the bulb.

Male (based on holotype, specimen NCBS-BN351). Carapace length 2.6; abdomen length 4.0. Carapace integument dark brown to black except to either side of fovea and narrow medial stripe on thorax, and with small patches of yellow scales in pattern typical for Indopadilla. Clypeus dark, extremely narrow, exposing white arthrodial membrane. Chelicerae dark, with lateral ridge bearing a tooth (Fig. 98, triangle). At least five retromarginal teeth, on long ridge. Palp yellow except for black cymbium. Embolus simple, long (Fig. 93); RTA a flat blade. Legs strongly contrasting between dark brown to black first legs, and yellowish posterior legs. Darkness of first legs relieved by pale tarsus, and honey coloured path on patella. Abdomen black except for two prominent yellow-white patches along each side.

A video of the living holotype is available in Maddison (2020).

Holotype female (specimen MRB076), in LKCNHM, from Singapore: Nee Soon Swamp Forest, beating vegetation, 1.39°N, 103.81°E, 12 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-015. Paratype female (specimen JK.14.05.19.0015) from Singapore: Central Catchment Nature Reserve Upper Peirce Reservoir 1.3811°N, 103.8156°E, J. K. H. Koh 19 May 2014.

An arbitrary combination of letters, ungendered. Other names: In WPM’s lab notebooks the informal code for this species was “BVNES”.

As in I. vimedaba, the face appears white because of the exposed arthrodial membrane, whose boundary with the very narrow clypeus is indistinct. Differs from I. vimedaba in having chevroned abdominal markings visible in alcohol and epigynal openings small and copulatory ducts densely tangled and fused (Fig. 101).

Female (based on holotype, specimen MRB076). Carapace length 2.5; abdomen length 3.6. Carapace integument in alcohol dark brown, paler in ocular area and along midline of thorax, with patches of yellow scales (Fig. 102). Clypeus extremely narrow, but broad arthrodial membrane is exposed and bright white. Chelicerae brown, with concave front and lateral ridge. At least four retromarginal teeth. Legs pale except first, whose femur is brown darkening to black ventrally, and whose tibia and metatarsus are red-brown in the middle. Abdomen dark except for large patches of yellow scales along each side. Epigyne with shallow ECP and small openings (Fig. 100).

All from Malaysia: Sarawak: Mulu Nat. Pk. Holotype. Female (specimen IDWM.20011, in UBCZ), from Botanical Trail, 4.0380°N, 114.8137°E, 50 m el. 18 March 2012 W. Maddison WPM#12-059. Paratype female (specimen SWK12-1831, in UBCZ) from Botanical Trail, 4.0404°N, 114.8151°E to 4.0405°N, 114.8154°E, 50 m el. 16 March 2012 Maddison/Piascik/Ang WPM#12-039. Paratype. Male (specimen SWK12-M0009, in UBCZ) from Nightwalk Trail, 4.0446°N, 114.8154°E to 4.0450°N, 114.8156°E, 50 m el. 24 March 2012 Maddison/Piascik/Ang WPM#12-090. Paratype male (specimen JK.12.01.22.0024, in LKCNHM) from Sungai Paku Waterfall Trail, 04.0372°N, 114.8247°E, J. K. H. Koh 22 January 2012.

Latin, meaning bent backward, referring to both the RTA and the ridge in front of the epigynal openings. Other names: In Maddison (2015b) and WPM’s field or lab notebooks the informal code for this species was “BVMTT”. In J. Koh’s notebooks it was referred to as “Bent RTA”.

The living paratype male 12.01.22.0024 is shown as figure E on p. 203 of Koh and Bay 2019.

Male palp distinctive for bent RTA and thick curled embolus accompanied by TmA (Fig. 104). Epigyne similar to that of I. redynis, with cavernous ECP, but with each opening framed by an anterior curved ridge. Posterior third of abdomen dark except for prominent pale transverse band just in front of anal tubercle. First femur pale basally, lacking dark spot of I. redynis.

Male (based on paratype, specimen SWK12-M0009). Carapace length 2.9; abdomen length 3.6. Carapace in alcohol dark brown except medium brown near fovea, and a small medium brown stripe medially on thoracic slope. Clypeus dark, glabrous, very narrow in middle, exposing a white arthrodial membrane beneath. Chelicerae dark, concave and with lateral ridge. At least four retromarginal teeth, on long sharp ridge. Palp pale except dark gray retrolateral face of femur. Embolus thick and curved, accompanied by similarly curved TmA (Figs 104, 106). RTA projects out away from axis of palp, curved toward the proximal (Figs 105 106). Legs strongly contrasting between the first legs (dark, except for patella, tarsus, and much of femur). Abdomen as described under Male-female matching.

Female (based on holotype, specimen IDWM.20011). Carapace length 3.2; abdomen length 4.1. Carapace dark brown, paler around fovea and in a narrow medial band on thoracic slope. Clypeus narrow, dark, glabrous, exposing white arthrodial membrane. Chelicerae vertical, concave in front. Five retromarginal teeth. Legs pale except for first leg, whose tibia and metatarsus are dark brown in the middle, and the femur which grades to dark brown terminally. Abdomen marked as described under Male-female matching. Epigyne (Figs 107, 108) with large ECP medially. Openings behind arcing ridges.

The male and female, both collected at Mulu National Park, are matched primarily on the basis of markings and expected genitalic correlations. They share abdominal markings (Figs 110–112): just in front of the anal tubercle is a prominent pale transverse mark, more prominent than in other species, and in front of the pale mark is a dark more or less unmarked area that extends to cover the posterior third of the abdomen. Just in front of that, in the middle third, are two uneven narrow broken longitudinal bands, besides which are similar uneven pale bands on the side. While other Indopadilla species have uneven longitudinal bands, no others known have the bands so restricted by a dark posterior third, nor have the pale pre-anal band so prominent. In genitalia, the large ECP is expected to correspond to a dramatic RTA, which the matched male has. A female was chosen as holotype as it is in best condition, and the molecular data are from a matching female.

Holotype female (specimen IDWM.20012, in UBCZ) from Malaysia: Sarawak: Kubah Nat. Pk., roadside, 1.603–4°N, 110.185°E, 350 m el. 7 March 2012 Maddison/Piascik/Ang/Lee WPM#12-002. Paratype female (specimen SWK12-0080, in UBCZ) from Malaysia: Sarawak: Kubah Nat. Pk., Waterfall Trail, 1.605–6°N, 110.185–7°E, 300 m el. 7 March 2012 Maddison/Piascik/Ang/Lee WPM#12-001.

An arbitrary combination of letters, ungendered. Other names: In WPM’s lab notebooks the informal code for this species was “BVMT2”; it was also sometimes confused with I. redunca and labelled “BVMTT”.

Similar in colour and epigyne to I. redunca. The epigyne differs in having the anterior part of the cavernous ECP with a more sharply defined boundary, the openings not behind a curved ridge, and the edge of the opening clearly sinuate (Fig. 113). The abdominal dorsum has fine reticulate markings throughout, not showing the unmarked posterior third. First leg markings match those of I. redunca except that anterior base of first femur with dark patch. Pale area on thoracic slope much broader than in I. redunca.

Female (based on holotype, specimen IDWM.20012). Carapace length 3.6; abdomen length 5.7. Carapace integument medium to dark red-brown, except upper part of thorax between and beside eyes, continuing as a broad medial pale area to pedicel. Clypeus dark, extremely narrow, exposing broad white arthrodial membrane. Chelicerae light brown, concave in front. Five teeth on retromargin. Legs pale except dark brown on middle of first tibia and metatarsus, and darker patch at front base of first femur. Abdomen red-brown, with delicate reticulate pale markings. Epigyne (Fig. 113) with large cavernous ECP; openings sinuate, resembling those of dendryphantines.

Holotype male (specimen d107, in UBCZ) from Malaysia: Sabah: Village of Kiabau. Central Sabah. 5.8315°N, 117.2245°E, 23 November 2000 K. Ober #00.437.

An arbitrary combination of letters, ungendered. Other names: In WPM’s lab notebooks the informal code for this species was “BVSAB”.

Palp similar to I. kahariana with a short sharp embolus with a TmA behind it, but the TmA is broader and more retrolaterally placed than in I. kahariana. RTA with sharp point, unlike the broad flat RTA of I. kahariana. These two species are sister groups on the molecular phylogeny (Fig. 1).

Male (based on holotype, specimen d107). Carapace length 2.8; abdomen length 3.8. Overall appearance similar to that of I. kahariana, paler coloured than many male Indopadilla, honey to medium brown. Carapace medium brown to honey coloured, darker on sides, with ocular area and larger portion of dorsal thorax quite pale. Clypeus narrow, exposing white arthrodial membrane. Chelicerae brown, with lateral ridge. Six retromarginal teeth, on long ridge. Palp pale except basal part of femur. Embolus short and sharp, with broad TmA behind (Fig. 117). Legs mostly honey coloured, first somewhat darker and with ventral fringe on patella through metatarsus. Abdomen in alcohol similar to that of female I. kahariana, with series of darker paired spots dorsally and elongated pale spots along the side.

Holotype male (specimen JK 13.09.03.0011), in LKCNHM, from Singapore: Nee Soon Swamp, 2 September 2013. J. K. H. Koh.

An arbitrary combination of letters, ungendered. Other names: In Maddison (2015b) and WPM’s field or lab notebooks the informal code for this species was “BVMDB”.

As noted under Padillothorax, this species was mistaken for Padillothorax semiostrinus, and illustrated by Prószyński (1987) under that name. He illustrates a male and female in the same vial 15151 in the MNHN Paris from the “Malayana” peninsula.

Palp with embolus longer that half the length of the bulb, straight and tapering (Fig. 121). Other Indopadilla have the embolus either longer and curved, or shorter. Epigyne with small but distinct ECP on back margin medially; openings simple and posterior (Fig. 123). Body dark brown except for highly contrasting but narrow pale cream markings, reticulate on abdomen.

Male (based on holotype, specimen JK 13.09.03.0011). Carapace length 2.4; abdomen length 3.5. Carapace in alcohol dark brown to black, slightly paler around fovea and in narrow medial band along thoracic slope, which also has some white scales; other patches of white scales along borders of ocular area. Clypeus extremely narrow at centre, and beneath it is a broad expanse of white arthrodial membrane; precise boundary between the clypeus and arthrodial membrane indistinct. Chelicerae with strong lateral ridge that extends into a flange near the fang, as drawn by Prószyński (1987: 105). Three promarginal teeth and at least five retromarginal. Palp honey coloured except darker base of femur. Embolus a narrow long triangle (Fig. 121). Legs honey coloured except much larger first leg, which bears a ventral fringe. Abdomen dark above and below, with two pair of lateral pale spots, the anterior of which connects with a few thin pale lines.

Female (based on specimen SWK12-3620). Carapace length 3.3; abdomen length 5.4. Carapace integument in alcohol medium red-brown except for narrow medial pale band along thoracic slope. Clypeus extremely narrow, exposing broad white arthrodial membrane (Fig. 128). Chelicerae concave in front with lateral ridge. At least four retromarginal teeth. Legs pale except first leg, dark except tarsus. Abdomen brown with distinct reticulation of pale scales. Epigyne (Fig. 123) with simple openings, from which broad copulatory ducts proceed anteriorly before narrowing considerably and proceeding to the posterior, then back anteriorly to the spermathecae.

The male and female described above were not co-collected, but they match well the male and female (MNHN 15151, photographs examined) described by Prószyński, which were in the same vial, and thus likely co-collected. These males and females match in markings: the first leg is all dark except the tarsus; the abdominal dorsum is dark with some narrow pale lines, the venter dark; the pale medial band of the integument of the thoracic slope is narrow; there is a narrow band of pale scales on the midline low on the thoracic slope; the triangle of scales near the fovea is narrow. The male has fewer lines in its abdominal markings than the female, but those it has are precise matches to the female. Other Indopadilla differ; e.g., another candidate female, I. nesinor, has the first leg considerably more strongly banded. suggesting her male should have a first patella paler than seen in Fig. 125, more like that of I redunca (Koh and Bay 2019:. 203, fig. E), whose female’s leg is banded similarly to I. nesinor. Also, I. nesinor differs in having a pale underside of the abdomen, and different thoracic markings. The relatively long but wide embolus of the male is a good fit to the openings and first broad part of the duct of the matched female, being ~ 0.07 mm wide at its base and ~ 0.3 mm long. In contrast, the openings of I. nesinor are only ~ 0.02 mm wide, enough to accommodate only a small portion of the embolus.

One female (specimen SWK12-3620, in UBCZ) from Malaysia: Sarawak: Mulu Nat. Pk., headquarters area, on & in buildings, 4.042°N, 114.814°E, 50 m el. 26–27 March 2012 Maddison/Ang WPM#12-100.

Stagetillus banda (Strand, 1911), comb. nov.

Stagetillus opaciceps Simon, 1885

Stagetillus irri Maddison, sp. nov.

Hyctiota banda Strand, 1911 is based on a juvenile (SMF 2511 in SMF, examined) and not easily placed. Its being a baviine is suggested by its elongate body (most like that of Indopadilla or Stagetillus), plurident chelicerae, and Asian origin. Its macrosetae sockets resemble those of baviines (Figs 21–23). It lacks the thoracic bulge and ridged chelicerae of Indopadilla, and the flat carapace of Padillothorax. Like Stagetillus, the fovea is displaced slightly to the posterior. The two dark bands running through the PLE shown in Strand’s (1911) original figure (no longer visible in specimen) resemble those of male S. opaciceps. The carapace is widest posteriorly, though not so clearly as in Figs 15 and 16. Although its synonymy with Stagetillus is not certain, by placing it with Stagetillus, a salticid genus otherwise incertae sedis is provisionally settled.

Carapace distinctive in shape, widest point toward the posterior, approx. half way between the back eyes and the pedicel, and in colour, orange or yellow with the white digestive diverticulum showing beneath the transparent ocular quadrangle. Palp much like that of Bavia, with short blade-like embolus.

Prószyński’s (1987) illustrations of Simon’s type specimen characterize the male well. Two additional males illustrated here (Figs 143–147) are conspecific or at least very close (the male from Brunei, Figs 146, 147, has a slightly narrower embolus). Females of the species have not been reported, although we now tentatively identify, as the female of S. opaciceps, the specimen MRB079 from Ulu Gombak used by Maddison et al. (2014) for molecular phylogeny (Figs 148–150). That the female is a Stagetillus is suggested by its similarities to the male in size and body form, carapace shape, yellow coloration, and visibility of the ocular digestive diverticulum. The modest ECP and epigynal openings are what might be expected from the small simple RTA and embolus of the known male of S. opaciceps. Given that Ulu Gombak is only 70 km from a known male of S. opaciceps (Figs 143, 144), the match of the Ulu Gombak female to S. opaciceps is credible. We therefore label the female, used here for molecular data, as S. cf. opaciceps (Figs 148–150). In WPM’s lab notebooks the informal code for the female was “STULG”.

Male (specimen JK.13.02.26.0017, in LKCNHM) Malaysia: Negeri Sembilan, Hutan Lipur Ulu Bendul, 2.73°N, 102.0789°E, J. K. H. Koh 26 February 2013; Male (JK.08.08.19.0001, in LKCNHM) Brunei: Belait, Disturbed forest off Labi Road, 4.5858°N, 114.5067°E, J. K. H. Koh 19 August 2008. Female (specimen MRB079, in UBCZ) Malaysia: Selangor: Ulu Gombak Field Station. 3.325°N, 101.753°E, 250 m el. 16–19 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang, WPM#05-026.

Holotype male (specimen IDWM.20023, in FSCA) and paratype female from Philippines: Luzon: Laguna Province, Los Baños, International Rice Research Institute, February 1993. R.R. Jackson (batch RRJ Ph 246/93). Other paratypes from same locality are 1 male (specimen IDWM.20022), December 1996, R.R. Jackson (batch RRJ Ph 292/96, in UBCZ); 3 females (specimens IDWM.20014, S202, and S203, in UBCZ), 1–16 December 1993, R.R. Jackson & G.B. Edwards; and 2 males 3 females, March 1993, R.R. Jackson (batch RRJ Ph 323/93, in FSCA).

From the acronym for the type locality. Other names: This species was referred to by Maddison and Hedin 2003 as “unident. (Phil.)” In WPM’s lab notebooks the informal code for this species was “STPHL”.

The FSCA holds more material of this species, including from Mt. Makiling, near the type locality but at higher elevation (G. B. Edwards, pers. comm.).

Similar in overall appearance to S. opaciceps, from which it differs in palp (narrower and longer embolus) and epigyne (shorter openings and having a central mound, presumably bearing the ECP). Male lacking the dense fringe beneath the first leg (Figs 155 vs. 146), but with a spur on the paturon (Fig. 155, arrow) that is lacking in S. opaciceps (Fig. 146). Male endite lacks the lateral bulge seen in S. opaciceps.

Male (based on holotype, specimen IDWM.20023). Carapace length 3.3; abdomen length 4.6. Carapace yellow-orange with two darker stripes passing along PME, PLE, and to posterior margin, with transparent ocular areas showing bright white digestive diverticular beneath. Shape as in S. opaciceps (Fig. 150). Clypeus narrow, more or less glabrous. Chelicerae orange, projecting slightly, with a spur on each paturon just above the fang base (arrow, Fig. 155). Palp pale except darker cymbium. Embolus arising at ~ 9:00 on the bulb, narrowing to a fine tip, with transparent flange retrolaterally (Fig. 151). Long lobe of tegulum overhangs the tibia. RTA broad and obtuse. Legs pale yellow to dark orange, first pair darkest, with femora of first two pair darker below. First leg long, with only a weak fringe beneath. Abdomen with dark central band with three wider areas, flanked by bright white.

Female (based on paratype, specimen IDWM.20014). Carapace length 2.9; abdomen length 5.2. Entirely light in colour, from white to medium orange, except for the black of eyes, and small black patch at the front distal tip of the first leg femur. Carapace pale orange, with transparent ocular area showing bright white digestive diverticular beneath. Shape as in S. opaciceps (Fig. 157). Chelicerae orange, with five retromarginal teeth. Legs pale to dark honey-orange, except for black on prolateral tip of femur I. Abdominal integument transparent in alcohol, showing white digestive glands to either side of beige heart and mottled stercoral pocket. Epigyne (Fig. 153) similar to those of Bavia, with sclerotized openings on either side of a central mound.

Padillothorax badut Maddison, sp. nov.

Padillothorax casteti (Simon, 1900), comb. nov., transferred from Bavirecta

Padillothorax exilis (Cao & Li, 2016), comb. nov., transferred from Bavirecta

Padillothorax flavopunctus (Kanesharatnam & Benjamin, 2018), comb. nov., transferred from Bavirecta

Padillothorax mulu Maddison, sp. nov.

Padillothorax semiostrinus Simon, 1901

Padillothorax taprobanicus Simon, 1902

The synonymy of Bavirecta with Padillothorax can be established now that the identity of the type species of the latter has been clarified (see below). The synapomorphies uniting them include the position of macrosetae on the first femur, flattened carapace, placement of fovea, pale thoracic “window” (Kanesharatnam and Benjamin 2018), and palp form. The deep genetic divide seen in Fig. 3 between P. flavopunctus and the P. badut species group (P. badut and P. mulu) might permit us to divide this into more than one genus, but this is unnecessary and not yet justified. The known Padillothorax species together make a recognizable genus, holding together well through their synapomorphies. In addition, there is no known data guiding us as to how to divide them and maintain monophyly, as we see no strong evidence resolving the trichotomy (P. semiostrinus, P. flavopunctus, P. badut + P. mulu).

Distinctive for the macroseta(e) in the middle of the front surface of the first leg femur, the palp with narrow distally-pointing embolus, the pale trapezoidal “window” dorsally on the thorax (Kanesharatnam and Benjamin 2018), and the flat carapace with fovea well posterior to the eyes. The first femur’s prolateral macrosetae, either one or two, are present in both males and females (Figs 32, 33). While salticids usually have macrosetae dorsally or on the front surface distally (Figs 29–31, 34, 35), the more central location (more ventral and more basal, Figs 32, 33) of Padillothorax is unusual, known from only a few other genera (Epocilla, Padilla, some marpissines). The thoracic window is also different from the pale area on the top of the thorax seen in other baviines (e.g., Bavia nessagyna in alcohol), where it is neither as pale nor does it extend so far posteriorly as in Padillothorax. Stagetillus spp., and Maripanthus draconis have a pale thorax, but it is not framed by dark, as most of the thorax is pale. Palp is also distinctively simple: embolus narrow, pointing distally (i.e., appears erect in standard ventral view), fused to the tegulum, with no tegular fold covering its base as in Bavia and many Indopadilla. Epigynal openings anterior with simple copulatory ducts proceeding posteriorly. Abdomen with distinct transverse bands. First legs distinctly the longest. Male endite with lobe or sharp corner, unlike Indopadilla (whose endite is rounded, without lobes or corners).

There has been confusion about the identity of P. semiostrinus. Prószyński (1987 p. 105) illustrated as P. semiostrinus a male and female from Simon’s collection that clearly belong to the group here called Indopadilla, as Prószyński’s illustrations show the group’s typical diagnostic features. If his illustrations had shown P. semiostrinus, then Indopadilla would have been junior synonym of Padillothorax. However, Prószyński’s illustrations are misidentified, representing a species quite distinct from P. semiostrinus, and which we describe above as Indopadilla vimedaba. Simon’s (1901a, b) descriptions are sufficient to show the distinctions. He notes the thorax of P. semiostrinus almost twice as long as the ocular quadrangle (approx. equal in Prószyński’s drawings and in Indopadilla generally), the carapace flatter than Bavia (as high or higher in Indopadilla), the fovea well back of the posterior eyes (immediately behind in Indopadilla), the retromarginal cheliceral teeth on a conical elevation (Indopadilla without elevation), and the male endite with a corner (well rounded in male Indopadilla). The last two distinctions are clear also in Simon’s illustration (1901b, p. 461), which shows the teeth on a mound and the lateral margin of the chelicera simple, in contrast to Prószyński’s illustration which shows the teeth not on a mound but spread along a ridge, and the lateral margin of the chelicera with an extended ridge (as in Indopadilla). Even Simon’s name, Padillothorax, emphasizes the similarity to Padilla, whose distinctive carapace (flat, long, with short ocular quadrangle) is quite unlike the more standard carapace shown in Prószyński’s drawings and Indopadilla in general. Thus, Prószyński’s drawings are misidentified. He was likely misled by the vial's label, which appears much like those of Simon’s other types.

As to what is Padillothorax semiostrinus, we have not been able to examine the type specimens, as they have not yet been located in the MNHN Paris. However, specimens found recently in Singapore and Taiwan match well Simon’s (1901a) description, which we translate here to English (with the assistance of Charmaine Gorrie and Anna Bazzicalupo):

"♂. Length 7.5 mm. Cephalothorax low, long and oval, red-brown, darker towards the border; texture very wormy-coriaceous except for the middle of the thoracic part which is smoother. Cephalic area in front and at both sides, [and?] near the eyes, decorated with white-silver hairs. Two wide medial thoracic bands, nearly contiguous; a thin marginal line decorated with white-silver hairs. Few white hairs around the eyes. Clypeus very narrow, bald. Abdomen narrow and very long, decorated above with dark violet, a medial band that is wide, entire, and yellow-brick red, bordered in front with lines and behind with a series of spots covered with silver-white hairs, marked on each side with a straight line in front and two white oblique [or crosswise?] lines behind. Venter reddish-yellow. Spinnerets dark. Chelicerae shiny black, short and diverging, convex outside, inside somewhat ribbed, inferior margin having a sunken furrow, then very raised and armed with a series of contiguous teeth, the middle larger. Mouth area black. Laminae truncate at the tip, convex, but with a compressed corner that is slightly extended. Sternum yellow. First pair of legs much longer and thicker than the others, femur clavate, tibia long and ovate, dark brown, coxa and femur black, tarsus yellow, tibia and metatarsus with fringe of reasonably long but not very dense black hairs. Remaining legs pale yellow, armed by a few small spines, as in Bavia. Palps reasonably small, pale yellow, thick with white hairs; tibia and patella rather short, equipped at the outside with a [long?] apophysis with a straight front and a black and acute tip."

There are two apparent or possible conflicts between this description and the specimen seen in Figs 158–164. The most serious conflict is that the RTA in Fig. 158 is short, but Simon’s description suggests the RTA is long. However, we were unable to understand the apparent use of genitive “apophysi” in “tibia patella breviore, extus ad apicem apophysi sat longa, antice directa, apice nigra et acuta, instructa”. What exactly is long: the tip of the apophysis, the apophysis, or the outer edge of the tibia to the tip of the apophysis? Second, his description refers to marks at the back of the abdomen that are “obliquus”, which would differ from Figs 164, 166 if it were translated as “oblique”, but match if translated as “crosswise”. Otherwise Simon's description and his mouthparts illustration are an excellent match to the Singapore specimen, including the distinctive pair of nearly-touching wide thoracic bands, the pattern of the abdomen, the thorax rugosity except in the middle, the shape of the carapace and position of the fovea, the shininess and shape of the chelicerae, the mound bearing the retromarginal cheliceral teeth, the shape of the endite, the colours of the legs, and the yellow sternum.

We therefore provisionally identify the specimen of Figs 9, 17, 24, 32, 158–164 as P. semiostrinus. Although we might have added “cf.” to its label to indicate our uncertainty (“P. cf. semiostrinus”), we avoid this so as to propose a stable concept of the species that could endure if Simon’s types are never found.

A juvenile found in Singapore (Fig. 165) suggests the likely appearance of females. Males and females of this or a very similar species have been found in Taiwan, not yet examined, but with photographs posted in the website Facebook (Figs 166, 167).

Natural history. The male in Singapore was found in the open on a simpoh air leaf (Dillenia suffruticosa). A video of the living juvenile (specimen AS19.2448) is available in Maddison (2020).

Adult male (specimen JK.20.06.20.001), in LKCNHM, from Singapore: Mandai Road, 1.4106°N, 103.7631°E. Y. Ng 20 June 2020. Juvenile (specimen AS19.2448), in UBCZ, from Singapore: Palau Ubin, 1.406°N, 103.971°E, 11 June 2019 Maddison/Morehouse et al. WPM#19-048.

This species shares the diagnostic features of the genus, including macrosetae on the middle of the front face of the first femur. P. flavopunctus has two such macrosetae on each femur, as in P. badut and P. mulu, but larger and placed even more proximally. We have concerns that the paratype female (IFS_SAL_679) is not conspecific with the male holotype, but if not conspecific, it is a closely related species, both by its strong morphological similarity (body form, markings) to the male and by its molecular proximity to a juvenile (IFS_SAL_1017) collected alongside the male at the type locality.

All from Malaysia: Sarawak: Lambir Hills Nat. Pk., and in UBCZ. Holotype male (specimen IDWM.20007) from Bukit Pantu Trail, 4.2035°N, 114.0304°E to 4.2039°N, 114.0303°E, 210 m el. 5 April 2012 Maddison/Piascik/Ang WPM#12-136; paratype female (specimen SWK12-4350) from Lepoh-Ridan Trail, 4.2019°N, 114.0278°E to 4.2019°N, 114.0275°E, 190 m el. 2 April 2012 Maddison/Piascik WPM#12-125; paratype male (specimen SWK12-4688) from Inoue Trail, 4.2000°N, 114.0353°E to 4.2002°N, 114.0350°E, 190 m el. 4 April 2012 Maddison/Piascik/Ang WPM#12-128; paratype female (specimen IDWM.20008) from Bukit Pantu Trail, 4.2035°N, 114.0304°E to 4.2039°N, 114.0303°E, 210 m el. 5 April 2012 Maddison/Piascik/Ang WPM#12-136.

From the Malay word badut, meaning clown. In the field we called these (and P. mulu) the “banded clowns”. Other names: In Maddison (2015b) and WPM’s field or lab notebooks this species was grouped with P. mulu under the informal name “BVBND”, until it was distinguished from that species as “BVBND-L”.

Very similar to P. mulu, differing most notably in details of genitalia. The embolus of P. badut lacks the prolateral basal ridges and has a longer terminal part; the epigyne has the openings hidden beneath a fold.

This and the other new Malaysian species (P. mulu) are very similar in appearance, thin and banded; together we consider them as the P. badut species group. When the first legs are held forward in life (e.g., Fig. 174), the spider appears as a series of transverse white bands approximately evenly spaced from anterior to posterior: the first leg annuli, the two transverse bands on the carapace, and the four transverse bands on the abdomen. Their bodies are more delicate and parallel-sided than other Padillothorax, and they have more complex emboli.

Male (holotype, specimen IDWM.20007). Carapace length 2.3; abdomen length 3.1. Carapace (Fig. 172): Black to dark brown except for band just back of the anterior eye row, and the pale trapezoidal window on thorax. Unusually flat (carapace height 0.72), with ocular area and most of the thorax on a plane, declining suddenly only near back of thorax. Fovea well behind the PLEs (by more than their diameter). Clypeus very narrow, dark. Chelicerae vertical, glabrous, dark. Teeth not examined in holotype, but another male from Lambir Hills has five retromarginal teeth, together in a mound as in P. semiostrinus. Palp pale except for base of femur. Base of embolus with two peaked projections on the retrolateral side (Fig. 168). Endite with lateral lobe (Fig. 10). Legs: First legs very distinctly longest and darkest, red-brown to black except for strong annulae with white scales. Legs II–IV pale yellow with just a smudge of dark laterally on a few segments of the fourth leg. Abdomen: narrow, with four white transverse bands.

Female (paratype, specimen IDWM.20008). Carapace length 2.1; abdomen length 3.1. Colour and structure matches that of male in nearly all aspects, with the most distinct difference being the slightly shorter first legs. Cheliceral teeth not examined in this specimen, but another female from Lambir Hills has four retromarginal teeth, together in a mound. Epigyne (Fig. 170) with two anterior openings, hidden under folds, and simple copulatory ducts visible without clearing. The copulatory duct has a diverticulum for the accessory gland extending laterally just before entering the simple spermatheca.

On large-leaved understory plants such as palms. In life, they often hold the front legs out or to the front.

All from Malaysia: Sarawak: Lambir Hills Nat. Pk., collected 4–6 April 2012 by Maddison/Piascik/Ang, in UBCZ. One female (specimen d548) from Inoue Trail, 4.2000°N, 114.0353°E to 4.2002°N, 114.0350°E, 190 m el. WPM#12-128. One male two females from Bukit Pantu Trail, 4.2028°N, 114.0305°E to 4.2032°N, 114.0305°E, 210 m el. WPM#12-134. One female from Bukit Pantu Trail, 4.2043°N, 114.0302°E to 4.2047°N, 114.0303°E, 210 m el. WPM#12-138. One female two juveniles from Bukit Pantu Trail, 4.2047°N, 114.0303°E to 4.2052°N, 114.0303°E, 200 m el. WPM#12-139. One female from Pantu Trail,4.2027°N, 114.0401°E to 4.2030°N, 114.0399°E, 150 m el. WPM#12-144.

All from Malaysia: Sarawak: Mulu Nat. Pk., in UBCZ. Holotype male (specimen IDWM.20009) from Summit Trail near Camp 1, 4.0496°N, 114.8589°E to 4.0493°N, 114.8594°E, 220 m el. 21 March 2012 Maddison/Piascik/Ang WPM#12-067. Paratype female (specimen SWK12-EP0105) from Summit Trail near Camp 1, 4.0489°N, 114.8606°E to 4.0486°N, 114.8610°E, 280 m el. 21 March 2012 Maddison/Piascik/Ang WPM#12-071. Paratype male (specimen SWK12-2556) from Summit Trail near Camp 1, 4.0491°N, 114.8601°E to 4.0489°N, 114.8606°E, 270 m el. 21 March 2012 Maddison/Piascik/Ang WPM#12-070. Paratype female (specimen SWK12-EP0108) and male (specimen IDWM.20010) from Base Trail near Camp 1, 4.0543°N, 114.8534°E to 4.0544°N, 114.8531°E, 130 m el. 23 March 2012 Maddison/Piascik/Ang WPM#12-084.

From the name of the type locality. Other names: In Maddison (2015b) and WPM’s field or lab notebooks this species was grouped with P. badut under the informal name “BVBND”, until it was distinguished from that species as “BVBND-M”.

Very similar to P. badut, differing most notably in details of genitalia. The embolus of P. mulu is shorter, and has a toothed ridge on the prolateral base; the epigyne has the openings exposed on a more or less flat surface.

Male (holotype, specimen IDWM.20009). Carapace length 2.5; abdomen length 3.5. Colour and structure matches that of P. badut as described above. Cheliceral teeth not examined. Palp pale except for base of femur. Base of embolus with various fine teeth; embolus shorter than in P. badut (Fig. 176).

Female (paratype, specimen SWK12-EP0105). Carapace length 2.1; abdomen length 3.3. Colour and structure matches that of P. badut as described above. Cheliceral teeth not examined in this specimen, but another female from Mulu has five retromarginal teeth. Epigyne (Fig. 178) similar to that of P. badut, except for the openings being exposed, not under a fold.

As for P. badut, on large-leaved understory plants. They often hold the front legs out or to the front.

All from Malaysia: Sarawak: Mulu Nat. Pk., in UBCZ. One female (specimen SWK12-EP0107) from Base Trail near Camp 1, 4.0543°N, 114.8534°E to 4.0544°N, 114.8531°E, 130 m el. 23 March 2012 Maddison/Piascik/Ang WPM#12-084. One female (specimen SWK12-2574) from Summit Trail near Camp 1, 4.0491°N, 114.8601°E to 4.0489°N, 114.8606°E, 270 m el. 21 March 2012 Maddison/Piascik/Ang WPM#12-070.

Maripanthus draconis Maddison, sp. nov.

Maripanthus draconis Maddison, sp. nov.

Maripanthus jubatus Maddison, sp. nov.

Maripanthus menghaiensis (Cao & Li, 2016), comb. nov. (transferred from Nannenus)

Maripanthus reinholdae Maddison, sp. nov.

Maripanthus smedleyi (Reimoser, 1929), comb. nov., transferred from Bavia

An arbitrary combination of letters, reminiscent of Marpissa (as the females resemble) and Piranthus (to which it is closely related). To be treated grammatically as masculine.

Epigynal atria long and gaping, anteriorly placed. Embolus long and beginning on the basal side of the tegulum, apparently freely articulated from the tegulum (as in the related Piranthus). Retromarginal cheliceral teeth close together, forming a single short ridge. Male endite with sharp corner (Fig. 11). Abdomen with central longitudinal pale band flanked by dark and stuttered into a chevron. Body unremarkable for the Salticinae, in contrast to its sister genus Piranthus which are unusually flat and robust. The small M. menghaiensis and M. reinholdae in some respects resemble Indopadilla or small Bavia, but they lack the ridged chelicera l paturon of the former and the short embolus of the latter.

Holotype male (specimen AS19.2232), two paratype females (specimens AS19.2250 and d547), all in LKCNHM, from Singapore: Bukit Timah Nature Reserve, stream at Jungle Falls Path. 1.3562°N, 103.7748°E to 1.3572°N, 103.7734°E 110–150 m elev. 12 June 2019 Maddison, Morehouse, & Marathe WPM#19-051. Paratype male (specimen WSG018) from Singapore: Nee Soon Swamp Forest. Beating vegetation. 1.39°N, 103.81°E 12 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-015.

Greek, δράκων, referring to the fiery colours of the male. Other names: In Maddison (2015b) and WPM’s field or lab notebooks the informal code for this species was “CFMAR”. A specimen that is likely a closely related but distinct species is shown in Koh and Bay 2019 as “Bavia” sp. B Black-collared long-bellied jumping spider.

Most similar to M. smedleyi, of which only the female is known; M. draconis differs in having longer epigynal atria (greater than half of epigynal length) and less distinct atrial cliff (“ac”, Fig. 191). Similar to M. jubatus in being large bodied, but M. draconis differs in having a honey-coloured translucent thorax, which in the female is speckled with black, and which in the male contrasts strongly against the black ocular area. Male front is impressively red-orange, with red-orange first femora, palps, and rings around eyes, and a golden face. Embolus slightly shorter than the other three species for which males are known. Ventral bump on palp femur.

Male (based on holotype, specimen AS19.2232). Carapace length 4.0; abdomen length 5.0. Carapace (Figs 196, 197) black in ocular area and in a thin line along ventral and posterior margin; thorax strongly contrasting honey coloured, glabrous except for a few vertical black setae. Clypeus covered with yellow-cream setae. Chelicerae vertical, a soft orange-red, covered with yellow-cream setae. Three promarginal and five retromarginal teeth. Palp femur pale, darkening to red-brown distally. Bulb elongate (Figs 188, 189); embolus arising on the prolateral basal corner and proceeding distally. Femur with small but distinct distal ventral bump. Endite with small corner projection. Legs with pale femora and darker markings more distally. First femur striking orange-red, patella to tarsus dark brown to black. First patella through metatarsus with sparse ventral fringe of setae that are black at base, white at tip. Abdomen with brown longitudinal medial band dorsally, flanked by scattered red scales, with some scattered white scales basally.

Female (paratype, specimen AS19.2250). Carapace length 4.2; abdomen length 6.1. Carapace (Fig. 199) integument black in ocular area, except for a pale spot on each side just medial to the PME (in other females, these pale spots form a transverse band that intersects with a longitudinal medial pale band, forming a pale cross in the ocular area). Thorax honey-coloured with distinct black speckles. Clypeus with scattered white setae. Chelicerae brown with darker patches, with scattered white setae. Three promarginal and five retromarginal teeth. Legs pale honey-coloured with various darker patches. Abdomen with central longitudinal pale chevroned band flanked by red scales. Epigyne with large atria leading to broad ducts (Figs 191, 192). Atria shallow laterally, declining rapidly at an “atrial cliff” (“ac”, Fig. 191) medial to which the surface is distinctly deeper. The ECP is apparently medial and small.

In Singapore, beating vegetation in forest understory.

In UBCZ. Singapore: Bukit Timah Nature Reserve, stream at Jungle Falls Path. 1.3562°N, 103.7748°E to 1.3572°N, 103.7734°E, 110–150 m elev. 12 June 2019 Maddison, Morehouse, & Marathe WPM#19-051 (one female); Bukit Timah Nature Reserve. 1.355°N, 103.78°E, 29 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-041 (1 female 1 juvenile). Malaysia: Johor: Gunung Belumut Recreational Forest. 2.066°N, 103.527°E, 60–100 m el. 24 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-038 (1 male, 1 female, 6 juveniles); near Kluang, Gunung Lambak. 2.025°N, 103.344°E, 50–100 m el. 25 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-039 (1 male). Pahang: Tanah Rata. Jungle Trail 9 from Robinson Falls. 4.46°N, 101.40°E, 1200–1500 m el. 21–22 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-035 (1 female). Selangor: Ulu Gombak Field Station, 3.325°N, 101.753°E, 250 m el. 16–19 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-026 (1 female 1 juvenile); canyon near Ulu Gombak, 3.325°N, 101.765°E, 275 m el. 17 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-027 (1 female 1 juvenile). Sarawak: Fairy Caves, near Kuching, 1.381–2°N, 110.117–9°E, 20 m el. 10 March 2012 Maddison/Piascik/Ang/Lee WPM#12-011 (1 female). The female from Sarawak is listed with some hesitation. It may be conspecific with a male Maripanthus from Brunei, which appears to be a closely related but distinct species, with embolus initially directed distinctly more to the dorsal, and slight different carapace markings (specimen JK 08.08.23.0004, in LKCNHM, from Brunei: Ulu Temburong National Park, Ashton Trail 4.5428°N, 115.1528°E J K H Koh 23 August 2008).

Holotype male (specimen NCBS-BN352, also known as AS19.4373) and paratype female (specimen NCBS-BN353, also known as AS19.4996), in NCBS collection, from India: Karnataka: Kodagu: Yavakapadi, Honey Valley area, buildings and roadside, 12.22°N, 75.66°E, 1100 m elev. 23–28 June 2019 W. Maddison & K. Marathe WPM#19-069.

Latin, meaning maned or crested, referring to the field of short black setae on the male ocular area. Other names: In WPM’s field or lab notebooks the informal code for this species was “CFMA2”.

Similar size and body form to M. draconis, but differs most notably in solid dark integument of the carapace and face. Male palp with dramatically long embolus and RTA.

Male (based on holotype, NCBS-BN352). Carapace length 3.9; abdomen length 4.8. Carapace (Figs 207–211) black, with scattered white scales on the top of the thorax. Front half of ocular area covered with vertical black setae, giving the appearance of a mane from in front (Fig. 210). Clypeus black, with long black setae. Chelicerae vertical, black. At least five retromarginal teeth. Palp black, but cymbium with white setae. Embolus long and swooping (Figs 202–204), beginning with a broad base over the tibia, proceeding prolaterally, then dorsally, then distally. RTA long and more or less straight. Femur lacks the ventral bump seen in M. draconis. Endite with small corner projection. Legs similar to those of M. draconis, with some segments of first legs dark, but differs in having the first femur pale honey darkening terminally to brown, and tarsus pale. Abdomen light brown mid-dorsally, darker laterally and with scattered cream coloured scales.

Female (based on paratype, NCBS-BN353). Carapace length 4.2; abdomen length 5.1. Carapace black except medium brown areas (yellow in alcohol) around fovea and along margin, covered with yellow cream scales in band along ventral margin, and dorsally on ocular area and anterior part of thorax (Figs 212–214). Clypeus covered with long cream-coloured setae. Chelicerae dark, covered with long cream setae. Three promarginal and five retromarginal teeth. Legs pale honey-coloured with various darker patches. Abdomen like that of male, but paler (Fig. 213). Epigyne (Fig. 205) similar to that of M. draconis, but with ECP split into a pocket on each side.

Found in dry hanging banana leaves.

All in NCBS collection. One male (specimen NCBS-BN354, also known as AS19.4403) from India: Karnataka: Kodagu: Yavakapadi, on top of car, 12.2408°N, 75.6547°E, 23 June 2019 K. Marathe WPM#19-068. One male one female (specimens NCBS-BN355 and NCBS-BN355) from India: Karnataka: Kodagu: near Madikeri, Rainforest Retreat, banana plantation, 12.480°N, 75.709°E, 30 June 2019 K. Marathe WPM#19-103.

Nannenus menghaiensis is here transferred to Maripanthus (and thus to the Baviini) based on its many close similarities with M. reinholdae, which itself is placed in Maripanthus by both morphological and molecular data. M. menghaiensis has an elongate body and pattern of thoracic and abdominal markings very much like those of other baviines (and unlike Nannenus, which is a compact-bodied ground dweller). See Diagnosis of M. reinholdae for distinctions therefrom.

A male (specimen IDWM.20013) that is either M. menghaiensis or a very closely related species is shown in Figs 223 and 224 (in UBCZ, from Malaysia: Selangor: Ulu Gombak Field Station, 3.325°N, 101.753°E, 250 m el. 16–19 May 2005. W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-026). Its palp is very much like that figured by Cao, Li and Żabka (2016), but it differs slightly in the base of the embolus and a narrower embolus.

Holotype male (specimen JK 11.12.24.0006), in LKCNHM, from Brunei: Ulu Temburong National Park, Canopy Walk Trail, 4.5522°N, 115.1578°E, J. K. H. Koh 24 Dec. 2011. Paratype female (specimen SWK12-1934, in UBCZ), from Malaysia: Sarawak: Mulu Nat. Pk., Botanical Trail, 4.0406°N, 114.8170°E to 4.0404°N, 114.8176°E, 50 m el. 16 March 2012 Maddison/Piascik/Ang WPM#12-044. Paratype female (specimen SWK12-1991, in UBCZ), also from Mulu Nat. Pk., Kenyalang Trail, 4.0229°N, 114.8128°E to 4.0228°N, 114.8134°E, 55 m el. 17 March 2012 Maddison/Piascik/Ang WPM#12-047.

Named in honour of Christa Deeleman-Reinhold, whose extensive work on southeast Asian spiders has greatly increased our knowledge of the area’s fauna. She has discovered and described over 350 new species, including 54 new salticids. Other names: In Maddison (2015b) and WPM’s field or lab notebooks the informal code for this species was “BVBGB”. This species is shown in Koh and Bay 2019 as “Bavia” sp. C Two-lined long-bellied jumping spider; the upper photograph on p. 208 is of the holotype.

Very similar to M. menghaiensis, and like it smaller and more Indopadilla-like in body form than M. draconis and M. jubatus (shorter first legs, more elongate abdomen). Differs from M. menghaiensis in the longer epigynal openings, and in details of the palp’s bulb. In view from the retrolateral, the embolus is first directed to the distal then quickly turns dorsally (M. menghaiensis, embolus begins toward the dorsal). When the embolus comes out from behind the tegulum it is directed slightly proximally (slightly distally in M. menghaiensis). The embolus is thinner near the tip than in M. menghaiensis.

Male (based on holotype, specimen JK.11.12.24.0006; living holotype shown on p. 208 of Koh and Bay 2019). Carapace length 2.9; abdomen length 4.0. Carapace (Fig. 222) black except for brown around fovea. Cream scales form a band crossing the thorax behind the PLEs, a small streak on the midline of the thoracic slope, and a narrow band along the lateral margins. Clypeus dark but centrally with a cluster of erect white scales that overhang the chelicerae. Chelicerae simple, vertical, dark. Plurident. Palp pale except for dark femur. Embolus long, arising retrolaterally before curling under the bulb prolaterally then proceeding distally (Figs 215–217). A short spur diverges from the embolus before it turns distally (Figs 215, 216). RTA vertical except for a bend and curl distally. Endite with small sharp corner, similar but smaller to that in the larger species of Maripanthus. Legs pale honey coloured except for first, which is black in all segments except the tarsus. Third and fourth legs darker near more distal joints. Abdomen thin, dark, with a pale mid-dorsal band, just lateral to which are indistinct longitudinal streaks of white scales.

Female (based on paratype, specimen SWK12-1934). Carapace length 3.7; abdomen length 5.0. Carapace as in male, but slightly paler in integument. Chelicerae with three promarginal and five retromarginal teeth. Legs as in male but with first legs only slightly darker than the others. Abdomen brown with central pale chevroned band, and thin white streaks as in male. Epigyne (Fig. 218) similar in conformation to M. draconis and M. jubatus, but with atria smaller and copulatory ducts not so compacted.

One female (specimen JK.12.02.04.0010, in LKCNHM) from Brunei: Belait, Trail To Wasai Teraja Secondary Forest, 4.2911°N, 114.4231°E, J. K. H. Koh 4 February 2012.

This species is close to M. draconis, with similar body form and markings (Figs 225, 226), but differing in details of the epigyne (Figs 227, 228). The atria are shorter, ~ ½ length of the epigyne. Within the atria, the central depressed area is separated from the lateral raised areas by a more distinct and more heavily sclerotized “atrial cliff” (“ac”, Fig. 227). The spermathecae are smaller.

Piranthus api Maddison, sp. nov.

Piranthus bakau Maddison, sp. nov.

Piranthus decorus Thorell, 1895

Piranthus kohi Maddison, sp. nov.

Piranthus mandai Maddison, sp. nov.

Piranthus planolancis Malamel, Nafin, Sudhikumar & Sebastian, 2019

Carapace surface rugose, with a coarse reticulate sculpturing throughout. Carapace flat (height well less than half the length), with ocular area and front part of thorax on a plane, and fovea well back of PLE, 1.3–1.5 × further from front of carapace than is the back of the PLE. Legs robust, especially the first pair. Embolus begins at prolateral basal corner of bulb; epigyne with central septum. Tip of abdomen black.

Two of the species (P. bakau and P. kohi) are distinctive for their black-and-white banding, three others (P. decorus, P. mandai, and P. planolancis) are more simply marked with brown and black, while the last (P. api) is a red-orange-black ember.

The four new species described here extend the range of Piranthus eastward as far as Borneo. The two previously described species, P. decorus (Thorell 1895; Caleb and Sanap 2017) and P. planolancis (Malamel et al. 2019; Nafin et al. 2020), are from Myanmar and India. A video of a living female of P. planolancis (specimen AS19.5940) is available in Maddison (2020).

Holotype female (specimen AS19.3205), in LKCNHM, from Singapore: Sungei Buloh Wetland Reserve, near Visitor Centre, 1.440°N, 103.734°E, 19 June 2019 Maddison/Marathe/Morehouse/et al. WPM#19-064.

From the Malay word, “api”, meaning “fire”, referring to the colour. Other names: In WPM’s field or lab notebooks the informal code for this species was “PIORG”.

A distinctively narrow species with bright red-orange legs.

Female (holotype, specimen AS19.3205). Carapace length 3.0; abdomen length 3.5. Carapace narrow and low, orange-red-brown with central black area covering ocular area and medial part of thorax (Figs 229, 233). Clypeus black. Chelicerae short and vertical, dark. Legs red-orange except for black patella and tibia of first legs. Abdomen dark brown above, with median longitudinal paler band that in alcohol appears as to uneven parallel cream bands, extending backward until the black end of the abdomen. Epigyne (Fig. 235) with medial septum and a small medial ECP.

The two specimens were both found deep within large grass tussocks overhanging a moist ditch. A video of the living holotype is available in Maddison (2020).

A second female (specimen AS19.3217, in UBCZ), raised in captivity, same data as the holotype.

Holotype male (specimen SWK12-0561, also known as d424), in UBCZ, from Malaysia: Sarawak: Bako Nat. Pk., Mangroves, beach forest, 1.722°N, 110.446°E, 0 m el. 8 March 2012 Maddison/Piascik/Ang/Lee WPM#12-003.

Referring to the type locality and to the holotype’s habitat, mangroves (Malay, bakau = mangrove). Other names: In Maddison (2015b) and WPM’s lab notebooks the informal code for this species was “BKOMG”. This species is shown in Koh and Bay 2019 as the male of “Bavia” sp. D Strong-armed flat jumping spider.

This and the closely similar P. kohi differ from other known Piranthus in having white transverse banding on the body and legs, and the posterior legs striped with black and translucent white. P. bakau (Figs 237–254) differs from P. kohi (Figs 255–269) in having:

• Second white transverse band on the dorsum of the abdomen (i.e., the first behind the basal band) complete or broken by only a slight space;
• Sides of thorax lacking the three distinct narrow vertical lines seen in P. kohi (at most only a hint of two);
• Carapace lateral to the PLE with a bare patch, lacking golden scales, extending from PME back to behind PLE, and lateral to the bare patch is a stripe of denser golden scales (Figs 245, 249);
• First and second tibia bicoloured (black with white tip, arrow in Fig. 248);
• Second femur bicoloured (white basally, black terminally, Fig. 249);
• Black dorsal band on the fourth tibia incomplete, beginning mid-segment and reaching to the end (Figs 246, 247).
• Carapace slightly flatter than in P. kohi.
• Embolus (Fig. 237) notably longer than in P. kohi (Fig. 255), closely resembling that of P. planolancis (Nafin et al. 2020);
• Epigyne with cavernous atria framed by a curved medial ridge, and relatively long copulatory ducts leading to a posterior tangle of tubes and spermathecae.

Juveniles have markings consistent with those of adults, and thus can be distinguished by the non-genitalic features above.

Male (based on holotype, SWK12-0561). Carapace length 2.7; abdomen length 3.0. Carapace with rugose surface, black on ocular area, dark brown otherwise, covered thinly with narrow golden to white scales except bare patch lateral to PLE, and on posterior slope. Clypeus narrow, dark, with a few white setae. Chelicerae small and vertical, dark, with a few pale setae. Palp (Figs 237–239) black except band of white scales terminally on femur, and cymbium, which is dark basally but then fades to white distally. Embolus long, arising on prolateral basal corner of bulb, looping first dorsally, then proximally, then distally. RTA a long blade. Endite margin rounded laterally. Legs relatively larger (compared to the body) than in many baviines, with first pair especially robust. First and second legs mostly dark and with terminal white annuli on the segments. First patella and tibia with ventral fringe of black setae. Third and fourth leg segments from patella to tarsus translucent white with black bands and stripes. Fourth tibia and patella with a pattern of short black and white bands. On the tibia, a black dorsal band begins not basally but half way to the end, and extends distally to the tip; basally, it is interrupted by white (Fig. 246; compare with Fig. 266). Abdomen with three narrow transverse white bands, a basal one, a second one behind that, and a third one behind that. Posterior to the third band the abdomen is a shiny black, but anterior it is dusted with golden setae.

Female (based on specimen AS19.2895). Carapace length 3.0; abdomen length 4.1. The specimen was collected and photographed as a small juvenile (Figs 251–254); as an adult its basic appearance is similar (Figs 249, 250). Structure and markings of carapace and legs as in male. Abdomen with second transverse band oblique. Epigyne (Fig. 240) with medial septum dividing deep broad atria. What serves as the ECP is not obvious; there may be two, folds posterior to the atrium on each side, midway between the epigastric furrow and the posteriormost part of the medial septum. If so, it may be intermediate between P. decorus, with a medial ECP (Caleb and Sanap 2017), and P. planolancis (Malamel et al. 2019; Nafin et al. 2020), with a pair of distantly separated lateral ECPs (Nafin et al. 2020: fig. 17).

P. bakau and P. kohi are similar in general appearance, have overlapping geographical ranges, and have been collected to date with only adult males or adult females at a locality, not both. This leads to a question of which male matches which female. Unless there are additional closely similar species in the same areas, the inferred matching is well supported by the differences in markings, carapace shape, and lengths of embolus/copulatory ducts. The male of P. bakau and the female from Tengkorak inferred to match it share the diagnostic traits mentioned above. Doubt might arise because of one difference in their markings: the female has the second transverse abdominal band more oblique, with its two sides meeting at a central peak, while in the holotype the band is straight across. However, the second male, from Brunei, shows a peak (Fig. 242). A juvenile co-collected with the male holotype has the band peaked, and looks very much like the female when immature. The female is not designated as a paratype, however, because of the possibility it is a different but very closely related species, given the geographical distance between it and the holotype and their different habitats.

Holotype male collected from mangroves; female from Tengkorak collected by shaking vines and understory trees near waterfall.

Two juveniles with same data as holotype. Also, one male (specimen JK.11.04.17.0040), in LKCNHM, from Brunei: Tutong, Tasek Merimbun, Zone C2, Palau Luba, Sungai Melunchur, 4.5817°N, 114.6872°E, J. K. H. Koh 17 Apr. 2011. One female (specimen AS19.2895), in UBCZ, from Malaysia: Johor: Gunung Belemut Forest, Lata Tengkorak, 2.055°N, 103.543°E, 250 m elev. 16 June 2019 Maddison/Morehouse/et al. WPM#19-057.

Holotype male (specimen AS19.1813), in LKCNHM, from Singapore: Sungei Buloh Wetland Reserve, 1.440–1.447°N, 103.730–103.735°E, 10 June 2019 Maddison/Morehouse/et al. WPM#19-045. Paratype female (specimen JK 19.07.19.0001), in LKCNHM, from Singapore: Pulau Ubin, Balai Quarry Trail, 1.4178°N, 103.9850°E, leg. P. Ng 19 July 2019.

This elegant species is named in honour of Joseph Koh Kok Hong, arachnologist, conservationist, and diplomat. Koh has worked tirelessly to build peace with nature. Through his collecting and excellent books (Koh 1989; Koh and Leong 2013; Koh and Bay 2019), he has substantially promoted our knowledge of southeast Asian spiders and stimulated interest in their diversity and beauty. Koh collected the first known specimen of this species, in Brunei, and enabled our field work in Singapore during which the holotype and other specimens were collected. Other names: In WPM’s lab notebooks the informal code for this species was “SUBSG”. This species is shown in Koh and Bay 2019 as the female of “Bavia” sp. D Strong-armed flat jumping spider.

P. kohi (Figs 254–269) shares with P. bakau (Figs 237–254) the pattern of transverse white bands, but differs from it in having:

• Three distinct vertical lines on each side of the thorax;
• Carapace lateral to the PLE more or less uniformly covered in white to gold setae;
• Second transverse dorsal band of the abdomen well broken at the middle;
• First and second tibiae and femora solid dark, not bicoloured;
• Black dorsal stripe extends the full length of tibia 4 (Figs 266, 267);
• Shorter embolus and copulatory ducts.

Juveniles can be distinguished by the non-genitalic features above.

Male (based on holotype, AS19.1813). Carapace length 2.5; abdomen length 2.5. Carapace with rugose surface, black, dusted above with narrow golden scales. Sides and back of thorax bare except for three narrow and distinct vertical lines of pale setae. Clypeus black. Chelicera vertical and black. Palp black except for white cymbium. Embolus arising on prolateral basal corner, proceeding ventrally then curving distally (Figs 255, 256) . RTA a long blade. Legs robust, especially the first pair. First two pairs black except for tarsus and metatarsus, and white annulae terminally on femur. First patella and tibia with ventral fringe of black setae. Fourth tibia and patella with a pattern of long black and white bands. On the tibia, a black band reaches from the base (on the prolateral side) all the way to the dorsal tip, forming an oblique dorsal band (Fig. 266; compare with Fig. 246). Abdomen similar to that of P. bakau, with three transverse white bands, anterior to the third of which the abdomen is dusted with golden setae, posterior to it a shiny black. Second transverse white band with a broad gap in both male and female.

Female (based on specimen, 12.04.11.0032). Carapace length 3.4; abdomen length 4.0. Structure and markings as in male, but generally more reddish, especially first and second legs, which are red-orange-brown in the femur and patella (and tibia of the second pair). Epigyne (Fig. 258) with central septum, but lacking the large cavernous atria of other Piranthus species.

See comments under P. bakau. Male and female P. kohi share the diagnostic traits mentioned above. The matching is supported by both males and females occurring in Singapore and in similar mangrove habitats – eight specimens from Sungei Buloh including 4 males; 6 specimens from Palau Ubin including 2 females.

Specimens in Singapore were found beating trees and vines in a mangrove area. It appeared that our greatest success in finding them was when shaking woody vines. Their motion when alive has a different sense than other baviines; rather than the sharply-jumping Indopadilla, or the frequently waving Padillothorax badut group, or the more sedate Piranthus planolancis and P. api, P. kohi is constantly flicking up and down the first legs, palps, and abdomen, somewhat like ant mimicking salticids. A video of the living holotype is available in Maddison (2020).

Singapore: Sungei Buloh Wetland Reserve, 1.440–1.447°N, 103.730–103.735°E, 10 June 2019 Maddison/Morehouse/et al. WPM#19-045 (3 additional males raised in captivity, 2 juveniles, UBCZ); Sungei Buloh Wetland Reserve, Coastal Trail, 1.446°N, 103.730°E to 1.445°N, 103.735°E, 19 June 2019 Maddison, Marathe, Ng WPM#19-063 (2 juveniles, UBCZ); Pulau Ubin, Chek Jawa, 1.4122°N, 103.9908°E, 11 June 2019 Maddison, Sung, & Outomuro WPM#19-047 (1 female raised in captivity, 2 juveniles, UBCZ); Lim Chu Kang Mangroves, Tree trunks and limbs, 1.44°N, 103.70°E, 13 May 2005 W. Maddison, I. Agnarsson, J. X. Zhang. WPM#05-020 (2 juveniles, UBCZ). Malaysia: Selangor: Ulu Gombak Field Station, 3.325°N, 101.753°E, 250 m el., 16–19 May 2005 W. Maddison, D. Li, I. Agnarsson, J. X. Zhang. WPM#05-026 (1 juvenile, specimen MRB109, UBCZ). Brunei: Belait, Kuala Balai, Sungai Mendarum Damit Freshwater Swamp Forest, 4.4386°N, 114.3581°E, J. K. H. Koh 11 Apr. 2012 (1 female, specimen JK 12.04.11.0032, in LKCNHM). The specimen from Ulu Gombak, used in the molecular study, matches P. kohi well in markings, but it is a juvenile and thus labelled conservatively in the phylogenies as P. cf. kohi.

Holotype male (specimen JK.91.05.31.0001), in LKCNHM, from Singapore: Mandai Track 15 Trail, 1.4106°N, 103.7783°E, J K H Koh 31 May 1991.

Named for the type locality. Other names: In lab notebooks the informal code for this species was “SGOMG”.

In colouration similar to P. planolancis, browns and blacks, but with body more compact and robust, as in P. bakau and P. kohi. Embolus with only a small loop before proceeding distally, and thus P. mandai is second in sequence from least to most rotated embolic bases: P. kohi (Fig. 255), P. mandai (Fig. 270), P. planolancis (Nafin et al. 2020), P. bakau (Fig. 237).

Male (holotype, specimen JK.91.05.31.0001). Carapace length 2.7; abdomen length 2.5. Carapace with rugose surface, black and dark brown, with a sparse covering of narrow scales that is more or less uniform: there is no bare patch beside the PLE, and no distinct three vertical thoracic lines, but there is a slight condensation of scales into a single vertical thoracic line, similar to that in P. bakau. Clypeus narrow and dark. Chelicerae vertical, dark, with a few pale scales basally. Palp brown. Palp similar to that of P. planolancis, but RTA much shorter (Figs 270–272), the shortest known among Piranthus. Legs light to dark brown, with indistinct markings (Figs 273, 274) similar to those of P. planolancis; darkest in the femur, patella, and tibia of the first leg, and the femora of the other legs. Abdomen brown above, lacking the transverse bands of P. bakau and P. kohi, but with the dark end of the abdomen (Fig. 274).

At least 16 species of elongate dull brown salticids collected in field work in Asia and New Guinea were initially thought to be baviines (example photographs, Figs 275–282), but an analysis of two genes (Fig. 283) in nine of the species show that they form a clade with the similar Nungia epigynalis, nested within the Viciriini. Photographs of them were released by Maddison (2015b) under the name Nungia. Their distinction from baviines is also morphological. They have unident or fissident chelicerae and a carapace that is dark, matte (not shiny), distinctly flat-topped, and relatively square. The male first legs are fringed below; the embolus is usually short and fixed to the tegulum. The group is unreported but diverse in New Guinea (based partly on collecting by WPM, Maddison and Zhang 2011; Nungia there mistakenly reported under “Bavia and related genera”), as might be expected for a member of the Astioida. Capeyorkia falls within this group (Fig. 282, 283; NPNGE and NPNGF are listed as Capeyorkia because of similarity of palps), and likely Pungalina as well. Both Pungalina and Capeyorkia may merit synonymy with Nungia. At least some species of Muziris Simon, 1901 belong here as well. Also within the group are two species misplaced in Bavia, one in Diplocanthopoda, and one in Cosmophasis. While we might have delayed their consideration until a future revision of Nungia, we address them here as part of organizing the baviines. We therefore make the following new combinations:

Nungia hatamensis (Thorell, 1881), comb. nov., transferred from Diplocanthopoda Abraham, 1925. See Fig. 280.

Nungia modesta (Keyserling, 1883), comb. nov., transferred from Bavia. Based on Keyserling’s (1883) figures, body form and colouration typical of the New Guinea Nungia.

Nungia papakula (Strand, 1911), comb. nov., transferred from Bavia. Male and female syntypes in SMF examined; appears closely related to N. hatamensis and a likely senior synonym of Muziris wiehlei Berland, 1938. See Fig. 281.

Nungia xiaolonghaensis (Cao & Li, 2016), comb. nov., transferred from Cosmophasis Simon, 1901. See Fig. 279.

Some of these new combinations, and others made above in baviines, correct mistaken placements that may have resulted from convergence in the general form of the male palp. The palps of Nungia hatamensis, Nungia xiaolonghaensis, Maripanthus menghaiensis, Bavia capistrata, and Bavia maurerae do in fact resemble those of the genera in which they had been placed, respectively, Diplocanthopoda (a hasariine), Cosmophasis (a chrysilline), Nannenus (a nannenine), Evarcha (a plexippine), and Epidelaxia (a nannenine). Those genera are all in the Saltafresia, a group phylogenetically distinct from the astioids and baviines. The resemblance is largely restricted to the palps, as the remainder of the body is quite distinct in each case. Convergent evolution of the basic form of the palp is widespread in salticids, insofar as the palps are simple and variation in a single dimension (embolus length) can generate palps that look superficially quite similar. The general shape of the palp should be used with caution in determining relationships.