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Research Article
Phylogenomic placement and revision of Iranattus Prószyński, 1992 jumping spiders (Salticidae, Plexippini, Plexippina)
expand article infoKiran Marathe‡§, Rishikesh Tripathi|, Ambalaparambil V. Sudhikumar|, Wayne P. Maddison‡
‡ University of British Columbia, Vancouver, Canada
§ National Centre for Biological Sciences, Tata Institute of Fundamental Research, Bengaluru, India
| Christ College, Irinjalakuda, India

Corresponding author: Kiran Marathe ( marathe12@gmail.com )

Academic editor: Danilo Harms
© 2024 Kiran Marathe, Rishikesh Tripathi, Ambalaparambil V. Sudhikumar, Wayne P. Maddison.
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.
Citation: Marathe K, Tripathi R, Sudhikumar AV, Maddison WP (2024) Phylogenomic placement and revision of Iranattus Prószyński, 1992 jumping spiders (Salticidae, Plexippini, Plexippina). Zoosystematics and Evolution 100(2): 531-542. https://doi.org/10.3897/zse.100.122034
ZooBank: urn:lsid:zoobank.org:pub:4488FFD3-5621-439E-9253-058E974EB0B3
Open Access

Abstract

The jumping spider genus Iranattus Prószyński, 1992, distributed from Africa to southwestern Asia, has been placed within the Harmochirina 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 I. principalis (Wesołowska, 2000) and I. rectangularis Prószyński, 1992. Additionally, the female of I. 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 Prószyński (1992) originally described the jumping spider genus Iranattus Prószyński, 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 extraordinarily 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 Kulczyński, 1895) have an apophysis on the male cymbium very similar to that of Iranattus. A relationship with Harmochirines was suggested by Wesołowska (2000), who, when describing Monomotapa Wesołowska, 2000 (later synonymized with Iranattus; Prószyński 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 Iranattus, currently composed of two species (World Spider Catalog 2024). Females were unknown until Wesołowska and Russell-Smith’s (2022) recent redescription of the African I. principalis (Wesołowska, 2000), known from Côte d’Ivoire, Nigeria, and Zimbabwe (Wesołowska 2000; Wesołowska and Russell-Smith 2011, 2022). We have recently collected I. rectangularis Prószyński, 1992, 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 phylogenomically using ultraconserved element (UCE) data and with information on female genitalic morphology. Additionally, we provide a comprehensive generic diagnosis and redescribe both species.

Materials and methods

Materials examined

The specimens of I. rectangularis were recently collected from the Desert National Park, Rajasthan, India. They are currently housed in the collection of the Centre for Animal Taxonomy and Ecology (CATE), Christ College, Kerala, with plans for eventual transfer to the Research 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. rectangularis used for taxonomic work, where #### represents a four-digit number.

The I. 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 ### is a two- or three-digit number. We interpret these to likely be Lamotte’s collection from Parc Nacional Banco (hence, “PNB”), Côte d’Ivoire, 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 ### is a three-digit number.

Morphology

We examined and photographed ethanol-preserved specimens using an Olympus OM-D E-M10 II camera mounted on an Olympus SZX12 or a Leica DMC4500 camera attached to a Leica M205 C stereoscope. We used a drawing tube attached to a Nikon ME600L compound microscope to prepare illustrations of I. 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 I. rectangularis specimens by digitally tracing the photographs.

Descriptions of color patterns are based on ethanol-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, tibia, metatarsus, and tarsus). Abbreviations used here are as follows: CO, copulatory opening; ECP, epigynal coupling pocket; PME, posterior median eye; PLE, posterior lateral eye; RTA, retrolateral tibial apophysis.

Taxon sampling for phylogenomic analysis

To test the phylogenetic placement of Iranattus, molecular data was gathered for I. rectangularis and added to Marathe et al.’s (2024) UCE phylogenomic dataset, which included 15 plexippines, two harmochirines, and one salticine. Because Iranattus’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 Iranattus 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.

Table 1.
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Specimens used in phylogenomic analysis.

Species Voucher Sex Locality Lat, long
Anarrhotus fossulatus Simon, 1902 AS19.1319 ♂ Singapore 1.379, 103.816
Artabrus erythrocephalus (C.L. Koch, 1846) AS19.2205 ♂ Singapore 1.355–7, 103.774–5
Baryphas ahenus Simon, 1902 d536 ♂ South Africa -25.95, 30.56
Bianor maculatus (Keyserling, 1883) NZ19.9864 ♂ New Zealand -42.1691, 172.8090
Carrhotus sp. AS19.4650 ♂ India 12.2145, 75.653–4
Chrysilla volupe (Karsch, 1879) AS19.6089 ♂ India 12.223, 76.627
Epeus sp. DDKM21.055 ♂ Singapore 1.355, 103.78
Evacin bulbosa (Żabka, 1985) AS19.2123 ♂ Singapore 1.406, 103.971
Evarcha falcata (Clerck, 1757) RU18-5264 ♂ Russia 53.721, 77.726
Ghatippus paschima Marathe & Maddison, 2024 IBC-BP833 ♂ India 12.220–1, 75.657–8
Habronattus hirsutus (Peckham & Peckham, 1888) IDWM.21018 ♂ Canada 48.827, -123.265
Hyllus keratodes (van Hasselt, 1882) DDKM21.028 ♂ Malaysia 3.325, 101.753
Hyllus semicupreus (Simon, 1885) AS19.4415 ♂ India 12.2156, 75.6606
Iranattus rectangularis Prószyński, 1992 DDKM21.091 juv. India 26.28, 70.40
Pancorius dentichelis (Simon, 1899) SWK12-0042 ♂ Malaysia 1.605–6, 110.185–7
Pancorius petoti Prószyński & Deeleman-Reinhold, 2013 SWK12-0195 ♂ Malaysia 1.603–4, 110.185
Pellenes limbatus Kulczyński, 1895 RU18-5679 ♂ Russia 50.0501, 89.3878
Plexippus paykulli (Audouin, 1826) AS19.7337 ♂ India 12.825–6, 78.252–3
Ptocasius weyersi Simon, 1885 DDKM21.069 ♂ Singapore 1.36, 103.78
Telamonia festiva Thorell, 1887 DDKM21.048 ♂ China 21.8105, 107.2925
Thyene imperialis (Rossi, 1846) AS19.6443 ♂ India 12.216, 76.625

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 PHYLUCE 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-i 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-i 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 analyses 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 inference, the option -m TEST (standard model selection followed by tree inference, edge-linked partition model, no partition-specific rates) was used with 10 search replicates. For the bootstrap analysis, a single IQ-TREE search was used for each of the 1000 search replicates.

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://doi.org/10.5061/dryad.ht76hdrpz).

Results

Phylogenetic results

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 fewer 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).

Table 2.
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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 Illumiprocessor; “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 QC Total UCE loci In at least 10 taxa After paralogy filter Filtered UCE sequence 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
Iranattus rectangularis DDKM21.091 SRR28802508 14825117 2926 2849 2818 2008593
Pancorius dentichelis SWK12-0042 SRR27728367 6025337 3092 3022 2988 2316987
Pancorius petoti SWK12-0195 SRR27728366 5116119 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

The phylogenetic results are shown in Fig. 1. The reciprocal monophyly of the subtribes Plexippina and Harmochirina 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 within Plexippina is largely consistent with Marathe et al. (2024) and has generally high bootstrap values.

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. Iranattus rectangularis is recovered distantly from the subtribe Harmochirina and placed as the sister lineage to Evarcha sensu lato within the subtribe Plexippina.

Iranattus 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, Pellenes limbatus, is placed as expected within the harmochirines. Thus, the similarities between Iranattus and harmochirines noted by Wesołowska (2000) and Maddison (2015) are convergences.

The placement of Iranattus in the Plexippina is also supported by the form of the epigyne. Wesołowska and Russell-Smith (2022) report a pair of coupling pockets in I. principalis, one on either side of a central atrium housing the copulatory openings, the same as we have found in I. 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 the Plexippina (e.g., Evarcha, Baryphas Simon, 1902; Pancorius Simon, 1902; Telamonia Thorell, 1887; Vicirionessa Wesołowska & Russell-Smith, 2022).

We therefore recognize Iranattus as a member of the subtribe Plexippina.

Taxonomic results

Family Salticidae Blackwall, 1841

Tribe Plexippini Simon, 1901

Subtribe Plexippina Simon, 1901

Iranattus Prószyński, 1992

Figs 2–7, 8–13, 14–19, 20–23, 24–29, 30–33, 34–41

Iranattus Prószyński, 1992: 97–98, f. 35–40.

Monomotapa Wesołowska, 2000: 159, f. 42–46 (synonymized by Prószyński, 2017: 36.).

Type species

Iranattus rectangularis Prószyński, 1992.

Species included

Iranattus principalis (Wesołowska, 2000); Iranattus rectangularis Prószyński, 1992.

Diagnosis

The remarkably long third legs of Iranattus (Figs 15, 18, 30, 32) and scoop-shaped cymbial apophysis (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 Iranattus, 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, Iranattus also has a shorter embolus lacking membrane (membrane-accompanied long embolus in Vailimia), a short RTA (long and curved in Vailimia), and two distinct deep conical ECPs (absent in Vailimia). 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 (Plexippoides Prószyński, 1984; Epeus Peckham & Peckham, 1886; and Erasinus Simon, 1899), but their apophyses are different in shape—in Iranattus, a long, broad blade with a rounded tip, concave in front so as to form a scoop; in Plexippoides, sharply pointed, for example.

Figures 2–7. 

Iranattus 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 (DDKM21.089); 6. Epigyne, ventral view (DDKM21.090); 7. Vulva, dorsal view (DDKM21.089). Scale bars: 0.2 mm.

Iranattus principalis (Wesołowska, 2000)

Figs 2–7, 8–13, 14–19

Monomotapa principalis Wesołowska, 2000: 160, 42–46.

Monomotapa principalis Wesołowska & Russell-Smith, 2011: 581, 96–98, 229–230.

Iranattus principalis Prószyński, 2017: 36, 14K, 17F (transferred from Monomotapa).

Iranattus principalis Wesołowska & Russell-Smith, 2022: 47, 29A–D, 30A–D.

Materials examined

In NHMUK, lacking complete labels. These are likely from Parc Nacional Banco, Côte d’Ivoire (see “Materials examined” for explanation). 2♀♀ (PNB21) • 1♂ 1♀ (PNB146) • 3♀♀ (PNB156) • 2♀♀ (PNB159) • 2♂♂ (PNB167) • 2♀♀ (PNB181) • 1♂ 1♀ (PNB192) • 2♂♂ 2♀♀ (PNB203).

Diagnosis

Larger than I. rectangularis, with an almost ovoid tegulum with a less prominent shoulder, RTA slightly bent near the tip (Figs 10, 11), and a multi-chambered spermatheca sandwiched between copulatory ducts dorsally and the epigynal plate ventrally.

Description

♂ (DDKM21.089). Measurements: Carapace 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); II–10 (2.8, 2, 2.4, 1.5, 1.2); III–16.6 (6.3, 2.9, 3.2, 2.5, 1.7); IV–10.1 (3.7, 1.6, 1.7, 1.8, 1.3). Leg formula III-IV-II-I. 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 anteriorly golden yellow, and remaining carapace dark brown. Lateral sides posteriorly and back sparsely covered with pale 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: III femur distinctly long. Femur golden yellow, distal segments yellowish-brown. Abdomen narrow, ovoid. Golden yellow with less prominent transverse pale bands. Spinnerets yellowish.

♀ (DDKM21.090). Measurements: Carapace 5.1 long, 5.1 wide. Abdomen length 6.4; width 4.8. Leg measurements: I–11.7 (3.8, 1.8, 2.7, 2.1, 1.3); II–11.6 (3.3, 2.8, 2.4, 1.8, 1.3); III–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 similar 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. Spinnerets yellowish. Epigyne (Figs 6, 7, 12, 13): Medially located copulatory opening flanked by conical-shaped ECP.

Figures 8–13. 

Iranattus 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.

Natural history

Wesołowska and Russell-Smith (2022) report Iranattus 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 (2♀ 8.40°N, 12.80°E) and Côte d’Ivoire (1♀ 8.40°N, 12.80°E; 2♂ 2♀ 8°44'N, 3°49'W) in the Musée royal de l’Afrique centrale, collected from the trees Cola laurifolia, Combretum fragrans, Anogeissus leiocarpus, and Crossopteryx febrifuga.

Distribution

Côte d’Ivoire, Nigeria, Zimbabwe, and Cameroon.

Iranattus rectangularis Prószyński, 1992

Figs 20–23, 24–29, 30–33, 34–41

Iranattus rectangularis Prószyński, 1992a: 97, f. 35–40.

Materials examined

1 ♂, 1♀, & 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 I. 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. 

Iranattus 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.

Figures 20–23. 

Iranattus 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. 

Iranattus rectangularis genitalia photographs. 24. Male left palp, ventral view (NRC-AA-7708); 25. Ditto, retrolateral 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.

Figures 30–33. 

Iranattus 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

♂ (NRC-AA-7708). Measurements: Carapace 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 III 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: III–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, posterior with pale hairs. Pale erect hairs on ocular area. Pale hair patch beneath PMEs. Black hair band starts anteriorly, 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 vertical, 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 covered with black hair. Abdomen narrow, ovoid. Brown with gray hair overlay. Spinnerets brown.

Figures 34–41. 

Iranattus rectangularis habitus. 34–38. Male; 39–41. Female.

♀ (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 II 2.41 [0.85, 0.47, 0.50, 0.32, 0.27], leg III 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 III–I–IV–II. Carapace shape similar to male. Ocular area orange anteriorly, white hairs sparsely 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. 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

Iranattus rectangularis was collected from the branches of non-native Vachellia tortilis alongside 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).

Figures 42, 43. 

Iranattus rectangularis habitat. 42. Vachellia tortilis woodland; 43. Aerial views of the landscape of the Desert National Park, Rajasthan, India.

Discussion

Iranattus 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: Aelurillina), 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 transfer of Iranattus 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 I. 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 Authority, Government of India, and supplemented by an additional 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. Beccaloni (NHMUK) for the loan of I. principalis specimens. We thank Galina Azarkina for providing information on additional material of I. principalis she examined. We thank Dmitry Logunov, Galina Azarkina, and Tamás Szűts for their time reviewing the manuscript and providing valuable comments. Funding to WPM was provided by an NSERC Canada Discovery Grant.

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