Indopadilla Caleb & Sankaran, 2019

Maddison, Wayne P., Beattie, Imara, Marathe, Kiran, Ng, Paul Y. C., Kanesharatnam, Nilani, Benjamin, Suresh P. & Kunte, Krushnamegh, 2020, A phylogenetic and taxonomic review of baviine jumping spiders (Araneae, Salticidae, Baviini), ZooKeys 1004, pp. 27-97 : 27

Indopadilla Caleb & Sankaran, 2019

Indopadilla Caleb & Sankaran, 2019. Type species Indopadilla darjeeling Caleb & Sankaran, 2019.

Species included.

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.

Diagnosis.

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).

Figures

Figures 4 - 35. Variation in traits among baviines 4, 5 oblique views of female prosoma 4 Bavia nessagyna (specimen IDWM. 20004; note convex front face of chelicerae, and typical thorax) 5 Indopadilla redunca (holotype IDWM. 20011; note concave front face of chelicerae, and thoracic bulges) 6 - 12 left male endite (9 also with chelicera) 6 Bavia nessagyna (holotype IDWM. 20005) 7 Indopadilla redunca (specimen SWK 12 - M 0009) 8 Stagetillus opaciceps (specimen JK. 08.08.19.0001) 9 Padillothorax semiostrinus (specimen JK. 20.06.20.001) 10 Padillothorax badut (specimen SWK 12 - 4688) 11 Maripanthus jubatus (specimen AS 19.4373) 12 Piranthus planolancis (specimen AS 19.5970) 13 - 20 carapace (all females except 15) 13 Bavia nessagyna (specimen IDWM. 20004) 14 Indopadilla redunca (holotype IDWM. 20011) 15 Stagetillus opaciceps male (specimen JK. 13.02.26.0017) 16 Stagetillus cf. opaciceps female (specimen MRB 079). 17 Padillothorax semiostrinus (specimen JK. 20.06.20.001) 18 Padillothorax mulu (from Mulu Nat. Pk.) 19 Maripanthus reinholdae (specimen SWK 12 - 1934) 20 Piranthus planolancis (specimen AS 19.5940) 21 - 28 Metatarsus of first leg, retrolateral view. All female except 24 male. All are of left legs except 23, 25, 28 which are of right leg, digitally flipped 21 Bavia nessagyna (specimen IDWM. 20004) 22 Indopadilla kahariana (specimen SWK 12 - 1876) 23 Stagetillus cf. opaciceps (specimen MRB 079) 24 Padillothorax semiostrinus (specimen SWK 12 - EP 0105) 25 Padillothorax mulu (specimen SWK 12 - EP 0105) 26 Maripanthus reinholdae (specimen SWK 12 - 1934) 27 Maripanthus draconis (from Gunung Belemut, Johor) 28 Piranthus planolancis (specimen AS 19.5940) 29 - 35 prolateral surface of male first leg femur. 29 Bavia cf. capistrata (specimen from Singapore) 30 Indopadilla kahariana (specimen from Lambir Hills Nat. Pk.) 31 Stagetillus opaciceps (specimen JK. 08.08.19.0001) 32 Padillothorax semiostrinus (specimen JK. 20.06.20.001) 33 Padillothorax badut (specimen SWK 12 - 4688) 34 Maripanthus draconis (specimen from Johor, Gunung Lambak) 35 Piranthus bakau (holotype). Scale bars: 0.1 mm (6 - 12, 21 - 35), 1.0 mm (4, 5, 13 - 20).

Figures 93 - 99. Indopadilla kodagura sp. nov., holotype. 93 male left palp, ventral view 94 same, retrolateral view of tibia 95 - 99 male. Scale bars: on palp 0.1 mm; on bodies 1.0 mm.

Figures 113 - 116. Indopadilla redynis sp. nov. 113 epigyne, ventral (holotype IDWM. 20012) 114 vulva, dorsal 115, 116 Female (SWK 12 - 0080). Scale bars on epigyne 0.1 mm; on bodies 1.0 mm.

Figures 121 - 128. Indopadilla vimedaba sp. nov. 121 male left palp, ventral view (holotype JK. 13.09.03.0011) 122 same, retrolateral view of tibia 123 epigyne, ventral (specimen SWK 12 - 3620) 124 vulva, dorsal 125 holotype male 126 - 128 female SWK 12 - 3620. Scale bars: on genitalia 0.1 mm; on bodies 1.0 mm.

Figures 76 - 79. Indopadilla bamilin sp. nov., male holotype SWK 12 - 1618 76 left palp, ventral view 77 same, retrolateral view of tibia 78, 79 holotype. Scale bars: on palp 0.1 mm; on bodies 1.0 mm.

Figures 100 - 103. Indopadilla nesinor sp. nov., female holotype MRB 076 100 epigyne, ventral 101 vulva, dorsal 102, 103 Body. Scale bars: on epigyne 0.1 mm.

Figures 129 - 142. Unidentified or undescribed Indopadilla, all in LKCNHM except 134 - 136, in UBCZ. 129 - 131 male " Yellow Long " (specimen JK 08.04.29.0029, Brunei, 4.6044 ° N, 114.6450 ° E), body, ventral palp, retrolateral palp tibia 132, 133 female " Yellow Long " (specimen JK. 12.03.14.0031, Brunei, 4.7036 ° N, 114.6264 ° E), epigyne, body 134 - 136 male " BVBTN " (specimen AS 19.2286, Singapore, 1.3562 ° N, 103.7748 ° E to 1.3572 ° N, 103.7734 ° E), body, ventral palp, retrolateral palp tibia 137, 138 female " Iridescent Ocular " (specimen JK. 13.02.16.3005, Brunei 4.5764 ° N, 115.0731 ° E), epigyne, body 139, 140 female " Orange Head " (specimen JK. 12.03.21.0011, Brunei, 4.5833 ° N, 114.5047 ° E), body, epigyne (shown alive as fig. B on p. 203 of Koh and Bay 2019) 141, 142 female " Gombak Reticulate " (specimen JK. 98.03.05.0001, Singapore, 1.3619 ° N, 103.7592 ° E), epigyne, body. Scale bars: on genitalia 0.1 mm; on bodies 1.0 mm.

Figures 1 - 3. Phylogeny. 1 Maximum Likelihood tree (best of 20 replicates) from combined data set of 1313 UCE loci, plus mitochondrial 16 SND 1 and COI regions, plus 28 S (dataset 4 in Methods). Baviines whose names are in bold, and all outgroups, have UCE data. Numbers are percentage of 500 bootstrap replicates showing the clade 2 Maximum Likelihood tree (best of 50 replicates) from concatenated data from 1313 UCE loci (dataset 1 in Methods). Numbers are percentage of 1000 bootstrap replicates showing the clade 3 Maximum Likelihood tree (best of 50 replicates) from concatenated data from mitochondrial data and 28 S (dataset 2 in Methods). Taxa in bold have data from the entire mitochondrial genome, or nearly so. Numbers are percentage of 1000 bootstrap replicates showing the clade for the full dataset ( 2), followed by the bootstrap percentage for the restricted mtDNA + 28 S dataset ( 3 in Methods). Spots at nodes show those clades that also appear in the ML tree in the restricted dataset. The branches to B. nessagyna and I. redynis are long, compacted visually by cutting and sliding part of the length over itself; the actual length therefore should be seen as longer by the length of the overlap.