Zoosyst. Evol. 100 (2) 2024, 531-542 | DOI 10.3897/zse.100.122034 > PENSUFT. Gee Ee BERLIN Phylogenomic placement and revision of /ranattus Proszynski, 1992 jumping spiders (Salticidae, Plexippini, Plexippina) Kiran Marathe! 2", Rishikesh Tripathi**, Ambalaparambil V. Sudhikumar?, Wayne P. Maddison™ FP WNMY British Columbia, V6T 1Z4, Canada https://zoobank. org/4488 FF D3-562 1-43 9E-9253-058E974E BOB3 Corresponding author: Kiran Marathe (marathe1 2@gmail.com) Department of Zoology, University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia, V6T 1Z4, Canada National Centre for Biological Sciences, Tata Institute of Fundamental Research, GKVK Campus, Bengaluru, 560065, India Centre for Animal Taxonomy and Ecology, Department of Zoology, Christ College, Irinjalakuda, Kerala, 680 125, India Departments of Zoology and Botany and Beaty Biodiversity Museum, University of British Columbia, 6270 University Boulevard, Vancouver, Academic editor: Danilo Harms # Received 1 March 2024 # Accepted 18 April 2024 Published 14 May 2024 Abstract The jumping spider genus /ranattus Proszynski, 1992, distributed from Africa to southwestern Asia, has been placed within the Har- mochirina because of their male palp structures and elongated third legs. Here, we present phylogenomic evidence that it belongs instead to the subtribe Plexippina, further supported by the presence of two coupling pockets in the female epigyne. In this study, we redescribe /. principalis (Wesolowska, 2000) and /. rectangularis Proszyhski, 1992. Additionally, the female of 1. rectangularis, the type species of the genus, is described for the first time, and we report its range extension east to India. Key Words Afrotropics, Araneae, biodiversity research, classification, deserts, Harmochirina, Indomalaya, phylogenomics, systematics, taxonomy, xeric scrublands Introduction When Proszynski (1992) originally described the jump- ing spider genus /ranattus Proszynski, 1992, based on a single male specimen from Iran, he characterized it by features such as a simple tegulum (bulbus) and embolus, unusual cymbial apophysis, and an extraor- dinarily long pair of legs (which his text erroneously states are the fourth pair, but which in fact are the third, as in his figures 35-36). These traits led Maddison (2015) to place it within the Harmochirina, some of which have very long third legs (e.g., Neaetha Simon, 1885), and some of which (e.g., Pellenes limbatus Kul- czynski, 1895) have an apophysis on the male cymbi- um very similar to that of /ranattus. A relationship with * These authors are with equal authorship. Harmochirines was suggested by Wesotowska (2000), who, when describing Monomotapa Wesotowska, 2000 (later synonymized with /ranattus, Proszynski 2017), commented on its similarity in body and leg lengths with the harmochirines Neaetha Simon, 1885, and Pellolessertia Strand, 1929. Subsequent studies and new material now give the opportunity to reconsider the phylogenetic placement of /ranattus, currently composed of two species (World Spider Catalog 2024). Females were unknown until We- sotowska and Russell-Smith’s (2022) recent redescription of the African 1. principalis (Wesolowska, 2000), known from Cote dIvoire, Nigeria, and Zimbabwe (Wesolows- ka 2000; Wesotowska and Russell-Smith 2011, 2022). We have recently collected /. rectangularis Proszynski, 1992, Copyright Marathe, K. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 532 in India, allowing us to not only characterize it through living photographs and natural history information but also to describe its female for the first time and to gather genetic data. We set out to clarify its placement phyloge- nomically using ultraconserved element (UCE) data and with information on female genitalic morphology. Addi- tionally, we provide a comprehensive generic diagnosis and redescribe both species. Materials and methods Materials examined The specimens of /. rectangularis were recently collect- ed from the Desert National Park, Rajasthan, India. They are currently housed in the collection of the Centre for Animal Taxonomy and Ecology (CATE), Christ Col- lege, Kerala, with plans for eventual transfer to the Re- search Collections at the National Centre for Biological Sciences (NCBS), Bengaluru, Karnataka, India (http:// collections.ncbs.res.in), for permanent deposition. NRC- AA-#### represent NCBS voucher codes of I. rectangu- laris used for taxonomic work, where #### represents a four-digit number. The . principalis specimens used in this study were in vials in a large jar of poorly labeled salticid specimens in the Natural History Museum, London (NHMUK). All the vials in the jar contained, typically, African salticids. Their labels bore only codes of the form “PNB ###”, where ### 1S a two- or three-digit number. We interpret these to likely be Lamotte’s collection from Parc Nacional Ban- co (hence, “PNB”), Céte dIvoire, from which Wanless (1985) cites similar code labels under Sonoita lightfooti Peckham & Peckham, 1903, e.g., PNB 179, PNB 146. Some specimens are identified by voucher codes of the form DDKM21 .###, where ### 1s a three-digit number. Morphology We examined and photographed ethanol-preserved speci- mens using an Olympus OM-D E-M10 II camera mount- ed on an Olympus SZX12 or a Leica DMC4500 camera attached to a Leica M205 C stereoscope. We used a draw- ing tube attached to a Nikon ME600L compound micro- scope to prepare illustrations of /. principalis. We used clove oil for clear viewing of epigyne after digesting the internal epigynal soft tissues with pancreatin. We stacked photographs using Helicon Focus 7.6.6 Pro. We prepared the drawings of J. rectangularis specimens by digitally tracing the photographs. Descriptions of color patterns are based on etha- nol-preserved specimens. Carapace length is measured from the base of the anterior median eyes to the posterior margin of the carapace medially, while abdomen length is measured from the anterior to the end of the anal tubercle. All measurements are in millimeters. Leg measurements are represented as follows: total length (femur, patella, zse.pensoft.net Marathe, K. et al.: /ranattus jumping spiders tibia, metatarsus, and tarsus). Abbreviations used here are as follows: CO, copulatory opening; ECP, epigynal coupling pocket; PME, posterior median eye; PLE, pos- terior lateral eye; RTA, retrolateral tibial apophysis. Taxon sampling for phylogenomic analysis To test the phylogenetic placement of /ranattus, molec- ular data was gathered for 1. rectangularis and added to Marathe et al.’s (2024) UCE phylogenomic dataset, which included 15 plexippines, two harmochirines, and one salticine. Because /ranattus’s former placement in the Harmochirina was based in part on some Pellenes having a similar cymbial apophysis, one such Pellenes (Pellenes limbatus) was added to the dataset to give the harmochirines the best chance to capture /ranattus in the phylogenetic analysis. An extra outgroup taxon, Chrysilla volupe (Karsch, 1879), was also added. The total set of 21 species used in the phylogenomic analysis, with their taxonomic authority indicated, is listed in Table 1. Ultraconserved element (UCE) data Molecular data was gathered for UCE loci using target enrichment sequencing methods (Faircloth 2017), using the RTA_v2 probeset (Zhang et al. 2023), and following the protocols of Marathe et al. (2024). Raw demultiplexed reads were processed with PHY- LUCE v. 1.6 (Faircloth 2016), quality control and adapter removal were performed with Illumiprocessor wrapper (Faircloth 2013), and assemblies were created with SPAdes v. 3.14.1 (Nurk et al. 2013) using options at default settings. The UCE loci were recovered using RTA_v2 probeset (Zhang et al. 2023). The recovered loci were aligned with MAFFT using L-INS-1 option (Katoh and Standley 2013). The aligned UCE loci were then trimmed with Gblocks (Castresana 2000; Talavera and Castresana 2007) using —b1 0.5, —b2 0.7, —b3 8, —b4 8, —b5 0.4 settings and re-aligned with MAFFT using L-INS-1 option within Mesquite v. 3.81 (Maddison and Maddison 2023b). As in the analysis of Maddison et al. (2020), suspected paralogous loci were deleted based on branch lengths in RAxML (Stamatakis 2014) inferred gene trees. Loci represented in fewer than 10 taxa total were deleted. Phylogenetic analysis Maximum-likelihood phylogenetic and bootstrap analy- ses were performed with IQ-TREE v. 2.2.0 (Nguyen et al. 2015) using the Zephyr v. 3.31 package (Maddison and Maddison 2023a) in Mesquite v. 3.81 (Maddison and Maddison 2023b) on the concatenated, unpartitioned UCE dataset with 20 taxa. For the phylogenetic tree in- ference, the option -m TEST (standard model selection followed by tree inference, edge-linked partition model, Zoosyst. Evol. 100 (2) 2024, 531-542 Table 1. Specimens used in phylogenomic analysis. 533 Species Voucher Sex Locality Lat, long Anarrhotus fossulatus Simon, 1902 AS19.1319 3 Singapore 1.379, 103.816 Artabrus erythrocephalus (C.L. Koch, 1846) AS19.2205 3 Singapore 1.355-7, 103.774-5 Baryphas ahenus Simon, 1902 d536 3 South Africa -25.95, 30.56 Bianor maculatus (Keyserling, 1883) NZ19.9864 3 New Zealand -42.1691, 172.8090 Carrhotus sp. AS19.4650 es India 12.2145, 75.653-4 Chrysilla volupe (Karsch, 1879) AS19.6089 3 India 12223,-76;627 Epeus sp. DDKM21.055 3 Singapore 13355. 103.78 Evacin bulbosa (Zabka, 1985) AS19.2123 3 Singapore 1.406, 103.971 Evarcha falcata (Clerck, 1757) RU18-5264 3 Russia 53. 721,797.726 Ghatippus paschima Marathe & Maddison, 2024 IBC-BP833 re India 12.220-1, 75.657-8 Habronattus hirsutus (Peckham & Peckham, 1888) IDWM.21018 3 Canada 48.827, -123.265 Hyllus keratodes (van Hasselt, 1882) DDKM21.028 3 Malaysia 3,325; 101,753 Hyllus semicupreus (Simon, 1885) AS19.4415 3 India 12.2156, 75.6606 lranattus rectangularis Prészynski, 1992 DDKM21.091 Juv. India 26.28, 70.40 Pancorius dentichelis (Simon, 1899) SWK12-0042 3 Malaysia 1.605-6, 110.185-7 Pancorius petoti Proszynski & Deeleman-Reinhold, 2013 SWK12-0195 3 Malaysia 1.603-4, 110.185 Pellenes limbatus Kulczynski, 1895 RU18-5679 3 Russia 50.0501, 89.3878 Plexippus paykulli (Audouin, 1826) AS19.7337 3 India 12.825-6, 78.252-3 Ptocasius weyersi Simon, 1885 DDKM21.069 3 Singapore 1.36, 103.78 Telamonia festiva Thorell, 1887 DDKM21.048 3 China 21.8105, 107.2925 Thyene imperialis (Rossi, 1846) AS19.6443 3 India 12.216, 76.625 no partition-specific rates) was used with 10 search rep- Results licates. For the bootstrap analysis, a single IQ-TREE search was used for each of the 1000 search replicates. Phylogenetic results Data availability The raw sequence reads obtained from UCE capture are stored within the Sequence Read Archive (BioProject: https://www.ncbi.nlm.nih.gov/bioproject/1101580), and their accession numbers are listed in Table 1. The UCE loci matrices from SPAdes assemblies, pre-Gblocks, and the concatenated matrices used for phylogenetic and bootstrap analysis, along with trees, are available on the Dryad data repository (https://do1.org/10.5061/dryad.ht76hdrpz). Harmochirina _ ico 100 Table 2 lists the sequence data recovered from the 21 taxa. 3398 UCE loci were initially recovered. Of these, 3140 remained after removing those represented in few- er than 10 taxa, and 3104 remained after removing those suspected to include paralogies on branch lengths. These were concatenated into the final matrix, whose aligned length is 2779616 base pairs, in which each taxon had on average ~2.2 million base pairs of sequence data (min. 985191, max. 2462121). The phylogenetic results are shown in Fig. 1. The recipro- cal monophyly of the subtribes Plexippina and Harmochirina Chrysilla volupe [AS19.6089] Carrhotus sp. [AS19.4650] Bianor maculatus [NZ19.9864] Habronattus hirsutus [IDWM.21018] Pellenes limbatus [RU18-5679] Ghatippus paschima [AS19.3805] 100 Plexippus paykulli [AS19.7337] Ptocasius weyersi [DDKM21.069] 100 Artabrus erythrocephalus [AS19.2205] Plexippini 100 00 100 Plexippina 100 96 4100 Anarrhotus fossulatus [AS19.1319] Epeus sp. [DDKM21.055] Thyene imperialis [AS19.6443] Telamonia festiva [DDKM21.048] Hyllus keratodes [DDKM21.028] Hyllus semicupreus [AS19.4415] 100 100 Pancorius dentichelis [Swk12-0042] : Pancorius petoti [SWK12-0195] 0.1 100 100 Baryphas ahenus [4536] lranattus rectangularis [DDKM21.091] Evacin bulbosa [AS19.2123] Evarcha falcata [RU18-5264] Figure 1. The IQ- TREE-based maximum-likelihood tree, represented here, is the best of 10 replicates, inferred from a concatenated dataset of 3104 UCE loci. The numbers at the nodes are the percentage recovery of the clade based on 500 bootstrap replicates. Tranattus rectangularis 1s recovered distantly from the subtribe Harmochirina and placed as the sister lineage to Evarcha sensu lato within the subtribe Plexippina. zse.pensoft.net 534 Marathe, K. et al.: /ranattus jumping spiders Table 2. Specifics of molecular data used for this phylogenomic analysis. Molecular data was generated based on the RTA_v2 probeset. “SRA” is the Sequence Read Archive accession number available through NCBI; “Reads pass QC” is the number of reads after the removal of adapter contamination and low-quality bases using IlIlumiprocessor; “Total UCE loci” is the total number of UCE loci recovered with RTA_v2 probeset; “After paralogy filter” is the number of UCE loci after deletion of suspected paralogous loci based on branch length ratios; “In at least 10 taxa” is the number of UCE loci in at least 10 or more taxa after branch length criteria; “Filtered UCE sequence length” is the concatenated sequence length of filtered UCE loci; “Total loci” is the number of UCE loci represented among all taxa. Species Voucher SRA Reads pass Total UCE In at least After Filtered UCE Qc loci 10 taxa paralogy sequence filter length Anarrhotus fossulatus AS19.1319 SRR27728361 15542927 2525 2444 2414 2100562 Artabrus erythrocephalus AS19.2205 SRR27728359 14903498 2837 2792 2759 2333639 Baryphas ahenus d536 SRR27728358 2653688 2256 2243 2217 985191 Bianor maculatus NZ19.9864 SRR27728369 7914005 2962 2853 2820 2422490 Carrhotus sp. AS19.4650 SRR27728370 5272657 2920 2838 2806 2324883 Chrysilla volupe AS19.6089 SRR28802507 4968344 2877 2782 2752 2313910 Epeus sp. DDKM21.055 SRR27728357 13896435 2897 2834 2802 2452270 Evacin bulbosa AS19.2123 SRR27728356 10851810 2766 2684 2653 2157554 Evarcha falcata RU18-5264 SRR27728355 11538276 2762 2714 2683 2215341 Ghatippus paschima IBC-BP833 SRR27728354 7881860 2893 2836 2804 2430054 Habronattus hirsutus IDWM.21018 SRR27728360 6581974 2821 2732 2702 2218729 Hyllus keratodes DDKM21.028 SRR27728353 11349372 2926 2843 2811 2415960 Hyllus semicupreus AS19.4415 SRR27728368 9874003 2942 2874 2839 2422661 lranattus rectangularis DDKM21.091 SRR28802508 14825117 2926 2849 2818 2008593 Pancorius dentichelis = SWK12-0042 = SRR27728367 6025337 3092 3022 2988 2316987 Pancorius petoti SWK12-0195 =SRR27728366 85116119 2980 2908 2875 2304191 Pellenes limbatus RU18-5679 SRR28802506 4288156 2661 2603 2576 1977916 Plexippus paykulli AS19.7337 SRR27728365 7445183 2931 2852 2817 2186676 Ptocasius weyersi DDKM21.069 SRR27728364 9926900 2880 2821 2790 2326688 Telamonia festiva DDKM21.048 SRR27728363 7908436 2950 2889 2855 2462121 Thyene imperialis AS19.6443 SRR27728362 7797854 2893 2818 2789 2421843 Average: 2843 2773 2741 2228488 Minimum: 2256 2243 2217 985191 Maximum: 3092 3022 2988 2462121 Total loci: 3398 3140 3104 2779616 is consistent with previous molecular phylogenetic studies with both Sanger sequencing and UCEs (Maddison and Hedin 2003; Maddison et al. 2008; Bodner and Maddison 2012; Marathe et al. 2024). The phylogenetic structure with- in Plexippina is largely consistent with Marathe et al. (2024) and has generally high bootstrap values. Tranattus is nestled well within Plexippina, placed as a sister lineage to Evarcha Simon, 1902 sensu lato (see Fig. 1). The harmochirine included in the analysis with a similar cymbial apophysis, Pe//enes limbatus, is placed as expected within the harmochirines. Thus, the similar- ities between /ranattus and harmochirines noted by We- sotowska (2000) and Maddison (2015) are convergences. The placement of /ranattus in the Plexippina is also supported by the form of the epigyne. Wesoltowska and Russell-Smith (2022) report a pair of coupling pockets in /. principalis, one on either side of a central atrium housing the copulatory openings, the same as we have found in J. rectangularis (Figs 22, 28). This arrangement is discordant with that of harmochirines, which have a single epigynal coupling pocket placed centrally, anterior to the margin, flanked by copulatory openings on either side. Two pockets are typical, however, for members of zse.pensoft.net the Plexippina (e.g., Evarcha, Baryphas Simon, 1902; Pancorius Simon, 1902; Telamonia Thorell, 1887; Vicirionessa Wesotowska & Russell-Smith, 2022). We therefore recognize /ranattus as a member of the subtribe Plexippina. Taxonomic results Family Salticidae Blackwall, 1841 Tribe Plexippini Simon, 1901 Subtribe Plexippina Simon, 1901 Tranattus Prészynski, 1992 Figs 2-41 Iranattus Proszyhski, 1992: 97-98, f. 35—40. Monomotapa Wesolowska, 2000: 159, f. 42-46 (synonymized by Proszynski, 2017: 36.). Type species. /ranattus rectangularis Proszynski, 1992. Species included. /ranattus principalis (Wesotowska, 2000); Iranattus rectangularis Proszynski, 1992. Zoosyst. Evol. 100 (2) 2024, 531-542 535 Figures 2-7. /ranattus principalis genitalia drawings. 2. Male left palp, ventral view (DDKM21.089); 3. Ditto, oblique view (DDKM21.089); 4. Ditto, oblique view, closeup of the cymbial apophysis (DDKM21.089); 5. Ditto, retrolateral view(DDKM721.089); 6. Epigyne, ventral view (DDKM21.090); 7. Vulva, dorsal view (DDKM21.089). Scale bars: 0.2 mm. Diagnosis. The remarkably long third legs of /ranattus (Figs 15, 18, 30, 32) and scoop-shaped cymbial apophy- sis (Fig. 4) differentiate it from all other plexippines. The very robust carapace, bulging outward at the PLE and bearing the PLEs on tubercles, is unusual but shared also with Afrobeata Caporiacco, 1941, and Vailimia Kammerer, 2006. Vailimia especially might be confused with /ranattus, as they share erect hairs on the carapace (see Figs 34, 38, 41) and a compact crouch stance, but, besides the cymbial apophysis and long third legs, /ranattus also has a shorter embolus lacking membrane (membrane-accompanied long embolus in Vailimia), a short RTA (long and curved in Vail- imia), and two distinct deep conical ECPs (absent in Vail- imia). From Afrobeata, Iranattus differs in having longer third legs, a cymbial apophysis (lacking in Afrobeata), a shorter embolus (longer in Afrobeata), a simple short RTA (bifurcated in Afrobeata), shorter copulatory ducts (long in Afrobeata), and deep conical ECPs (shallow in Afrobeata). Some other plexippines have cymbial apophyses (Plex- ippoides Proszynski, 1984; Epeus Peckham & Peckham, 1886; and Erasinus Simon, 1899), but their apophyses are different in shape—in /ranattus, a long, broad blade with a rounded tip, concave in front so as to form a scoop; in Plexippoides, sharply pointed, for example. Tranattus principalis (Wesolowska, 2000) Figs 2-19 Monomotapa principalis Wesotowska, 2000: 160, 42-46. Monomotapa principalis Wesotowska & Russell-Smith, 2011: 581, 96-98, 229-230. Tranattus principalis Proszynski, 2017: 36, 14K, 17F (transferred from Monomotapa). Iranattus principalis Wesotowska & Russell-Smith, 2022: 47, 29A—D, 30A-D. Materials examined. In NHMUK, lacking complete la- bels. These are likely from Parc Nacional Banco, Cote dIvoire (see “Materials examined” for explanation). zse.pensoft.net 536 299 (PNB21) * 14 19 (PNB146) * 399 (PNB156) ° 29° (PNBI159) * 2¢¢ (PNB167) * 299 (PNB181) ° 145 192 (PNB192) * 24'3' 2929 (PNB203). Diagnosis. Larger than /. rectangularis, with an al- most ovoid tegulum with a less prominent shoulder, RTA slightly bent near the tip (Figs 10, 11), and a multi-cham- bered spermatheca sandwiched between copulatory ducts dorsally and the epigynal plate ventrally. Description. 3 (DDKM21.089). Measurements: Cara- pace 2.2 long, 2.1 wide. Abdomen length 1.7; width 1.4. Leg measurements: I-11.2 (3.4, 2.2, 2.4, 1.8, 1.3); I-10 (28425 24. eS 1.2) MN 16:66:38 72192352, 2 Sd lV 10.1 (3.7, 1.6, 1.7, 1.8, 1.3). Leg formula III-IV-0-I. Cara- pace wider than abdomen. Ocular area shaped like an isos- celes trapezoid, narrow at the anterior eye row and wide at the PLEs. PLEs on tubercles. Thoracic area slopes acute- ly downward behind ocular area. Ocular area anteriorly golden yellow, and remaining carapace dark brown. Lat- eral sides posteriorly and back sparsely covered with pale Marathe, K. et al.: /ranattus jumping spiders hairs. Clypeus narrow, yellowish-brown sparsely covered with hairs. Chelicerae vertical, narrow, yellowish brown. Palp (Figs 2—5, 8-11): Embolus medium-long, starting at 7 o’clock. RTA stout, short with blunt tip. Cymbium extends retrolaterally to form scoop-shaped apophysis. Tegulum prolaterally rounder; retrolaterally slightly angular at distal and proximal edges. Legs: II] femur distinctly long. Femur golden yellow, distal segments yellowish-brown. Abdomen narrow, ovoid. Golden yellow with less prominent trans- verse pale bands. Spinnerets yellowish. 9 (DDKM21.090). Measurements: Carapace 5.1 long, 5.1 wide. Abdomen length 6.4; width 4.8. Leg measure- ments: I-11.7 (3.8, 1.8, 2.7, 2.1, 1.3); I-11.6 (3.3, 2.8, 2.4, 1.8, 1.3); II-19.7 (6.9, 3.3, 4.7, 3.1, 1.8); IV-11.4 (2.9, 2.1, 2.3, 2.6, 1.5). Leg formula III-I-II-IV. Carapace shape simi- lar to male, width about same as abdomen. Brown, sparsely covered with pale hairs. Clypeus similar to male. Chelicerae similar to male. Legs similar to male. Abdomen ovoid, bulky, yellowish, covered with brown hairs, and more posteriorly. Figures 8—13. /ranattus principalis genitalia photographs. 8. Male left palp, ventral view (DDKM21.089); 9 Ditto, retrolateral view (DDKM21.089); 10. Ditto, oblique view (DDKM21.089); 11. Ditto, retrolateral view (DDKM21.089); 12. Epigyne, ventral view (DDKM21.090); 13. Vulva, dorsal view (DDKM21.089). ECP, epigynal coupling pocket. CO, copulatory opening. Scale bars: 0.2 mm. Arrows in Figs 10 and 11 point to the scoop-shaped retrolateral cymbial apophysis. zse.pensoft.net Dah Zoosyst. Evol. 100 (2) 2024, 531-542 Spinnerets yellowish. Epigyne (Figs 6, 7, 12, 13): Medially located copulatory opening flanked by conical-shaped ECP. Natural history. Wesotowska and Russell-Smith (2022) report /ranattus principalis as collected from the branches of savannah shrubs. G. Azarkina (pers. comm.) has seen material of this species from canopy fogging in tropical savannas in Cameroon (29 8.40°N, 12.80°E) and Céte d’Ivoire (19 8.40°N, 12.80°E; 2¢ 29 8°44'N, 3°49'W) in the Musée royal de |’ Afrique centrale, collect- ed from the trees Cola laurifolia, Combretum fragrans, Anogeissus leiocarpus, and Crossopteryx febrifuga. Distribution. Céte d’Ivoire, Nigeria, Zimbabwe, and Cameroon. Tranattus rectangularis Prészynski, 1992 Figs 20-41 Tranattus rectangularis Proszynski, 1992a: 97, f. 35-40. Materials examined. 1 @, 12, & 4 juveniles. From INDIA: RAJASTHAN: Jaisalmer: Thar Desert: Desert National Park, Myajlar area, 26.28°N, 70.40°E, 275 m elev., 20 Aug 2022, leg. R. Tripathi. Diagnosis. Smaller than /. principalis, with a bright orange face and erect hairs on the carapace, an angular tegulum with a prominent shoulder, a simple RTA, and a simple spermatheca with copulatory ducts ventrally. Figures 14-19. /ranattus principalis habitus. 14. Male, dorsal view (DDKM21.089); 15. Ditto, lateral view (DDKM21.089); 16. Ditto, ventral view (DDKM21.089); 17. Female, dorsal view (DDKM21.090); 18. Ditto, lateral view (DDKM21.090); 19. Ditto, ventral view (DDKM21.090). Scale bars: 1 mm. Arrows in Figs 15 and 18 point to the long third legs. zse.pensoft.net 538 Marathe, K. et al.: /ranattus jumping spiders Figures 20-23. /ranattus rectangularis genitalia drawings. 20. Male left palp, oblique view (NRC-AA-7708); 21. Ditto, retrolateral view (NRC-AA-7708); 22. Epigyne, ventral view (NRC-AA-7709); 23. Vulva, dorsal view (NRC-AA-7709). Scale bars: 0.1 mm. Figures 24-29. /ranattus rectangularis genitalia photographs. 24. Male left palp, ventral view (NRC-AA-7708); 25. Ditto, retro- lateral view (NRC-AA-7708); 26. Ditto, oblique view (arrow points to the scoop-shaped retrolateral cymbial apophysis); 27. Ditto, dorsal view (NRC-AA-7708); 28. Epigyne, ventral view (NRC-AA-7709); 29. Vulva, dorsal view (NRC-AA-7709). ECP, epigynal coupling pocket. CO, copulatory opening. Scale bars: 0.1 mm. zse.pensoft.net Zoosyst. Evol. 100 (2) 2024, 531-542 ao] Figures 30—33. /ranattus rectangularis habitus. 30. Male, dorsal view (NRC-AA-7708); 31. Ditto, lateral view (NRC-AA-7708); 32. Female, dorsal view (NRC-AA-7709); 33. Ditto, lateral (NRC-AA-7709). Scale bars: 1 mm. Arrows in Figs 30 and 32 point to the elongated third legs. Description. 4 (NRC-AA-7708). Measurements: Car- apace 1.46 long, 1.26 wide. Abdomen length 1.32, width 0.86. Leg measurements: Leg I 2.04 [0.69, 0.33, 0.48, 0.32, 0.22], leg II 1.88 [0.67, 0.33, 0.44, 0.28, 0.16], leg Il 3.56 [1.52, 0.55, 0.72, 0.43, 0.34], leg IV 1.94 [0.70, 0.31, 0.34, 0.38, 0.24. Leg formula: II—I-IV-II. Carapace wider than abdomen. Ocular area shaped like an isosceles trapezoid, narrow at the anterior eye row and wide at the PLEs. PLEs on tubercles. Thoracic area slopes acutely downward behind ocular area. Ocular area from base of front eyes to PMEs orange, covered with black hairs, pos- terior with pale hairs. Pale erect hairs on ocular area. Pale hair patch beneath PMEs. Black hair band starts anteri- orly, encircles carapace at ocular area edge. White band along lateral edge, narrow front, broadens posteriorly and behind. Clypeus narrow. Orange, covered with pale hairs, more densely near integument edge. Chelicerae verti- cal, narrow, yellowish brown. Palp (Figs 20, 21, 24—27): Embolus medium-long, starting at 9 o’clock, somewhat thick. RTA stout, short with blunt tip. Cymbium extends retrolaterally to form scoop-shaped apophysis. Tegulum prolaterally rounder; retrolaterally angular at distal and proximal edges. Legs: III femur distinctly long relative to others. Femur yellowish, distal segments yellowish zse.pensoft.net 540 er, Ye z; ee lg tae Ci OnE a = Figures 34—41. /ranattus rectangularis habitus. 34-38. Male; 39-41. Female. covered with black hair. Abdomen narrow, ovoid. Brown with gray hair overlay. Spinnerets brown. 9 (NRC-AA-7709). Measurements: Carapace 1.91 long, 1.56 wide. Abdomen length 1.83, width 1.27. Leg measurements: Leg I 2.67 [0.95, 0.53, 0.56, 0.36, 0.27], leg IT 2.41 [0.85, 0.47, 0.50, 0.32, 0.27], leg HI 4.30 [1.78, 0.70, 0.92, 0.50, 0.40], leg IV 2.44 [0.87, 0.40, 0.42, 0.44, 0.31]. Leg formula IN—I-IV-II. Carapace shape similar to male. Ocular area orange anteriorly, white hairs sparse- ly posteriorly. Pale erect hairs on ocular area. Thoracic slope covered with black hairs. Lateral sides covered with pale hairs, almost merging behind. Clypeus similar as in male. Chelicerae similar to male. Legs similar to male. zse.pensoft.net Marathe, K. et al.: /ranattus jumping spiders Abdomen shape comparable to male, but with a ‘kite’- shaped black color pattern between posterior edge and median. Epigyne (Figs 22, 23, 28-29): Medially located copulatory opening flanked by conical-shaped ECP. Natural history. /ranattus rectangularis was collected from the branches of non-native Vachellia tortilis along- side artificial water canals in the Desert National Park, a xeric and desert ecosystem located in Rajasthan, India (Figs 42, 43). The mosaic of orange, black, and grey body coloration helps them blend in with the branches, making them inconspicuous, except that in the field, the orangish faces of males (Fig. 34) sometimes stood out. Distribution. Iran, India (Rajasthan). Zoosyst. Evol. 100 (2) 2024, 531-542 ~ 42 ~ ly ng Figures 42, 43. /ranattus rectangularis habitat. 42. Vachellia tortilis woodland; 43. Aerial views of the landscape of the Des- ert National Park, Rajasthan, India. Discussion. /ranattus rectangularis is reported for the first time east of Iran, in western India. This seemingly ‘disjunct’ distributional pattern is quite possibly due to a lack of collecting between the sites and mirrors that of Stenaelurillus marusiki Logunov, 2001 (Salticidae: Aelu- rillina), where the type locality of S. marusiki is Iran. However, it has been reported much farther southeast in Maharashtra, India (Marathe et al. 2022). With the trans- fer of Jranattus to Plexippina, the subtribe now contains 35 genera, and the number of plexippines in India stands at 47 species and 18 genera. Acknowledgements Collection of J. rectangularis was facilitated by the Bustard Recovery Programme of the Wildlife Institute of India (WII), funded by the National Compensatory Afforestation Fund Management and Planning Authori- ty, Government of India, and supplemented by an addi- tional grant from the Rajasthan State Pollution Control Board. RT and AVS thank Rev. Fr. Jolly Andrews, CMI of Christ College (Irinjalakuda), for facilities and NCBS research collections. RT acknowledges the support of Dr. Sutirtha Dutta, Dr. Manju Siliwal, and Mr. Ashish Kumar Jangid from WII, as well as the Rajasthan State Forest Department, for the collecting permit. Special thanks to Sohan Lal Genwa and Amrat Genwa for their assistance during field activities. RT thanks Anshuman Pati and Jason D. Gerard for habitat photographs. RT acknowledges CSIR-UGC for fellowship. KM thanks Dr. Krushnamegh Kunte, NCBS, for providing the lab space and supplies. KM and WPM thank Carol Ritland and Allyson Miscampbell of the Genetic Data Centre at the University of British Columbia for assistance with lab facilities. We thank J. Beccalonit (NHMUK) for the loan of /. principalis specimens. We thank Galina Azarkina for providing information on additional ma- 541 terial of J. principalis she examined. We thank Dmitry Logunov, Galina Azarkina, and Tamas Szits for their time reviewing the manuscript and providing valu- able comments. Funding to WPM was provided by an NSERC Canada Discovery Grant. References Bodner MR, Maddison WP (2012) The biogeography and age of sal- ticid spider radiations (Araneae: Salticidae). Molecular Phyloge- netics and Evolution 65(1): 213-240. https://doi.org/10.1016/). ympev.2012.06.005 Castresana J (2000) Selection of conserved blocks from multiple align- ments for their use in phylogenetic analysis. Molecular Biology and Evolution 17(4): 540-552. https://doi.org/10.1093/oxfordjournals. molbev.a026334 Faircloth BC (2013) illumiprocessor: a trimmomatic wrapper for paral- lel adapter and quality trimming. https://doi.org/10.6079/J9OILL Faircloth BC (2016) PHYLUCE is a software package for the analysis of conserved genomic loci. Bioinformatics (Oxford, England) 32(5): 786-788. https://do1.org/10.1093/bioinformatics/btv646 Faircloth BC (2017) Identifying conserved genomic elements and designing universal bait sets to enrich them. Methods in Ecology and Evolution 8(9): 1103-1112. https://doi.org/10.1111/2041- 210X.12754 Katoh K, Standley DM (2013) MAFFT Multiple Sequence Alignment Software Version 7: Improvements in Performance and Usabili- ty. Molecular Biology and Evolution 30(4): 772-780. https://doi. org/10.1093/molbev/mst010 Maddison WP (2015) A phylogenetic classification of jumping spi- ders (Araneae: Salticidae). The Journal of Arachnology 43(3): 231. https://doi.org/10.1636/arac-43-03-23 1-292 Maddison WP, Hedin MC (2003) Jumping spider phylogeny (Araneae : Salticidae). Invertebrate Systematics 17(4): 529. https://doi. org/10.1071/1S02044 Maddison DR, Maddison WP (2023a) Zephyr: a Mesquite package for interacting with external phylogeny inference programs. http:// zephyr.mesquiteproject.org Maddison WP, Maddison DR (2023b) Mesquite: a modular system for evo- lutionary analysis. http://www.mesquiteproject.org/ [August 18, 2023] Maddison WP, Bodner MR, Needham KM (2008) Salticid spider phylogeny revisited, with the discovery of a large Australasian clade (Araneae: Salticidae). Zootaxa 1893(1): 49. https://doi. org/10.11646/zootaxa. 1893.1.3 Maddison WP, Beattie I, Marathe K, Ng PYC, Kanesharatnam N, Ben- jamin SP, Kunte K (2020) A phylogenetic and taxonomic review of baviine jumping spiders (Araneae, Salticidae, Baviini). ZooKeys 1004: 27-97. https://do1.org/10.3897/zookeys. 1004.57526 Marathe K, Sanap R, Joglekar A, Caleb JTD, Maddison WP (2022) Three new and notes on two other jumping spider species of the genus Stenaelurillus Simon, 1886 (Salticidae: Aelurillina) from the Deccan Plateau, India. Zootaxa 5125(1): 1-19. https://doi. org/10.11646/zootaxa.5125.1.1 Marathe K, Maddison WP, Kunte K (2024) Ghatippus paschima, a new species and genus of plexippine jumping spider from the Western Ghats of India (Salticidae, Plexippini, Plexippina). ZooKeys 1191: 89-103. https://doi.org/10.3897/zookeys.1191.114117 zse.pensoft.net 542 Nguyen L-T, Schmidt HA, Von Haeseler A, Minh BQ (2015) IQ- TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolu- tion 32(1): 268-274. https://doi.org/10.1093/molbev/msu300 Nurk S, Bankevich A, Antipov D, Gurevich A, Korobeynikov A, Lapidus A, Prjibelsky A, Pyshkin A, Sirotkin A, Sirotkin Y, Stepanauskas R, McLean J, Lasken R, Clingenpeel SR, Woyke T, Tesler G, Alekseyev MA, Pevzner PA (2013) Assembling Genomes and Mini-metagenomes from Highly Chimeric Reads. In: Deng M, Jiang R, Sun F, Zhang X (Eds) Research in Computational Molecular Biology. Lecture Notes in Computer Science. Springer Berlin Heidelberg, Berlin, Heidelberg, 158-170. https://doi. org/10.1007/978-3-642-37195-0_ 13 Proszynski J (1992) Salticidae (Araneae) of the Old World and Pacific Islands in several US collections. Annales Zoologici, Warszawa 44: 87-163. Proszynski J (2017) Pragmatic classification of the world’s Saltici- dae (Araneae). Ecologica Montenegrina 12: 1-133. https://doi. org/10.37828/em.2017.12.1 Stamatakis A (2014) RAXML version 8: A tool for phylogenetic analy- sis and post-analysis of large phylogenies. Bioinformatics (Oxford, England) 30(9): 1312-1313. https://doi.org/10.1093/bioinformatics/ btu033 zse.pensoft.net Marathe, K. et al.: /ranattus jumping spiders Talavera G, Castresana J (2007) Improvement of Phylogenies after Re- moving Divergent and Ambiguously Aligned Blocks from Protein Sequence Alignments. Kjer K, Page R, Sullivan J (Eds) Systematic Biology 56: 564-577. https://do1.org/10.1080/10635 150701472164 Wanless FR (1985) A revision of the spider genera Holcolaetis and Sonoita (Araneae: Salticidae). Bulletin of the British Museum (Natural History). Bulletin of the British Museum, Natural History. Zoology 48: 249-278. https://doi.org/10.5962/bhl.part.23463 Wesolowska W (2000) New and little known species of jumping spi- ders from Zimbabwe (Araneae: Salticidae). Arnoldia Zimbabwe 10: 145-174. Wesolowska W, Russell-Smith A (2011) Jumping Spiders (Araneae: Salticidae) from Southern Nigeria. Annales Zoologici 61(3): 553- 619. https://doi.org/10.3161/000345411X603409 Wesolowska W, Russell-Smith A (2022) Jumping spiders from Ivory Coast collected by J.-C. Ledoux (Araneae, Salticidae). European Journal of Taxonomy 841(1): 1-143. https://doi.org/10.5852/ ejt.2022.841.1943 World Spider Catalog (2024) World Spider Catalog. https://wsc.nmbe. ch/ [August 18, 2023] Zhang J, Li Z, Lai J, Zhang Z, Zhang F (2023) A novel probe set for the phylogenomics and evolution of RTA spiders. Cladistics 39(2): 116-128. https://doi.org/10.1111/cla.12523