The Indian specimens examined in this study are deposited in two repositories. (i) The male holotype (I/SP-48) and a female paratype (I/SP-49) in the Southern Regional Centre, Zoological Survey of India (ZSIC), Chennai, Tamil Nadu, India and (ii) three paratypes (1 ♂ and 2 ♀♀) in the Biodiversity Lab Research Collections of the National Centre for Biological Sciences (NCBS), Bengaluru, India (http://biodiversitycollections.in/). Individual specimens deposited in NCBS are identified by three-digit voucher codes prefixed with “IBC-BX”. Non-Indian specimens are deposited in the University of British Columbia Spencer Entomological Collection.

We examined and photographed 70% ethanol-preserved specimens using a Leica MC190 HD camera attached to a Leica SAPO stereomicroscope automatically using the Leica Application Suite (LAS) v. 4.13. We examined and photographed the genitalia using a Leica MC190 HD camera attached to a Leica DM3000 LED compound microscope and prepared the drawings by digitally tracing the photographs. We photographed the living spiders with a Tamron 90 mm macro lens attached to a Nikon Z6 camera and a Godox flash covered with a Radiant Diffuser.

Descriptions of colour patterns are based on ethanol-preserved specimens. The male genitalic description is based on the left palp. 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 millimetres. Leg measurements are represented as follows: total length (femur, patella, tibia, metatarsus, and tarsus). Abbreviations used here are as follows: ALE, anterior lateral eye; ECP, epigynal coupling pocket; PME, posterior median eye; RTA, retrolateral tibial apophysis.

The morphology (conspicuous tegular lobe and two ECPs) of the species of Tenkana is consistent with their current placement among the plexippines. Therefore, we gathered molecular data for T. arkavathi, T. jayamangali, and T. manu and appended it to (Marathe et al. 2024b) plexippine-biased UCE phylogenomic dataset, which included 16 plexippines, five outgroups (three harmochirines, one salticine, and one chrysilline). We also included cf. Colopsus sp. from Lin et al. (2024) to allow cf. Colopsus, along with Pancorius from the Marathe et al. (2024b) dataset, a chance to capture Tenkana in the phylogenomic analysis. The total set of 25 species comprising 20 plexippines and five outgroups used in the phylogenomic analysis, with their taxonomic authorities indicated, is listed in Table 1.

To test Tenkana’s placement relative to the type species of Colopsus and others from Sri Lanka, we constructed matrices of mitochondrial cytochrome oxidase I (COI) and nuclear 28S, 18S and Histone 3 (H3) from publicly available data for Colopsus from Kanesharatnam and Benjamin’s (2021) study. We appended bycatch data for the same four gene regions present among the sequence capture genomic assemblies from the UCE dataset to the four gene matrices of C. cancellatus, C. ferruginus, and C. magnus. We followed a bycatch protocol similar to that described by Maddison et al. (2020), constructing local BLAST databases from SPAdes (Nurk et al. 2013) assemblies of each taxon in the UCE dataset. These assemblies were queried with publicly available COI, 28S, 18S, and H3 sequences from seven different salticid species (Aelurillus cf. ater (Kroneberg, 1875), Bianor maculatus (Keyserling, 1883), Colopsus cancellatus, Colopsus ferruginus, Hyllus treleaveni Peckham & Peckham, 1902, Pancorius athukoralai Kanesharatnam & Benjamin, 2021, and Salticius scenicus (Clerck, 1757)). Thus, the total set of 31 taxa (25 UCE bycatch and six Sri Lankan Colopsus of three species) are used in the four-gene phylogenetic analysis. The accession numbers are listed in Table 3.

Molecular data was gathered for UCE loci using target enrichment sequencing methods (Faircloth 2017), using the RTA_v3 probeset (Zhang et al. 2023) and following the protocols of Marathe et al. (2024a).

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_v3 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 setting within Mesquite v. 3.81 (Maddison and Maddison 2023b). As in the analysis of Maddison et al. (2020), loci suspected to include paralogies were deleted based on branch lengths in RAxML (Stamatakis 2014) inferred gene trees. Loci represented in fewer than 10 taxa total were deleted.

Maximum-likelihood phylogenetic and bootstrap analyses were performed with IQ-TREE v. 2.3.4 (Minh et al. 2020) 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 25 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.

The loci were aligned using MAFFT with the L-INS-i option, partitioned by locus, assigned codon positions to minimize stop codons for H3 and COI, and then concatenated in Mesquite v. 3.81. Maximum-likelihood phylogenetic and standard bootstrap analyses were performed with IQ-TREE v. 2.3.4 on the concatenated dataset using the Zephyr v. 3.31 package in Mesquite v. 3.81. The option -m MFP+MERGE (find the best partition scheme including FreeRate heterogeneity followed by tree inference, edge-linked partition model (-spp), with partition-specific rates) was used with 10 search replicates. For the bootstrap analysis, a single IQ-TREE search was used for each of 1000 search replicates. For both, four partitions were provided. IQ-TREE found the best partition scheme (Chernomor et al. 2016) and best fitting models (Kalyaanamoorthy et al. 2017) for phylogenetic tree inference: GTR+F+I+R2 model for 28S+H3 merged partition, GTR+F+I+G4 for 18S and COI separate partitions. Two trees were obtained: (1) unconstrained tree for 34 taxa, followed by standard bootstrap analysis (Fig. 3). (2) A constrained tree for the same dataset, followed by standard bootstrap analysis with an additional option of -g to specify (Tenkana, (Hyllus, Telamonia)) as the topological constraint (Fig. 2).

The raw sequence reads obtained from UCE capture are stored within the Sequence Read Archive (BioProject: PRJNA1145028, https://www.ncbi.nlm.nih.gov/bioproject/PRJNA1145028), and their accession numbers are listed in Table 2. The sequences obtained through UCE bycatch are available from the nucleotide database of the National Center for Biotechnology Information (NCBI, https://www.ncbi.nlm.nih.gov/), and their accession numbers are listed in Table 3. Concatenated UCE and legacy Sanger four-gene matrices used for phylogenetic and bootstrap analysis, along with trees, are available on the Dryad data repository (https://doi.org/10.5061/dryad.b8gtht7np).

Table 2 lists the sequence data recovered from the 25 taxa. 3404 UCE loci were initially recovered. Of these, 3302 remained after removing those suspected to include paralogies on branch lengths, and 3043 remained after removing those represented in fewer than 10 taxa. These were concatenated into the final matrix whose aligned length is 2557548 base pairs, in which each taxon had on average ~2 million base pairs of sequence data (min. 940794, max. 2281787). Table 3 lists sequence data for four genes for 34 taxa, including bycatch sequence data for 25 UCE taxa and Colopsus spp. gathered from NCBI.

The phylogenetic results are shown in Figs 1–3. In the UCE phylogeny (Fig. 1), the broader phylogenetic structure and the structure within Plexippina are consistent with (Marathe et al. 2024a, 2024b) and show generally high bootstrap values at the nodes as expected for high volume data. The constrained four-gene phylogeny (Fig. 2) respects the constraint used for tree search. Thus, the relationship of Tenkana is recovered similarly to the UCE phylogeny as (Tenkana, (Hyllus, Telamonia)), showing high bootstrap values. However, the general relationships among plexippines in unconstrained and constrained four-gene phylogeny are less reliable, as reflected in the low bootstrap values.

Figure 1. 

Maximum-likelihood tree from IQ-TREE analysis (best of 10 replicates) of a concatenated dataset of 3043 UCE loci. Numbers at the nodes are the percentage recovery of the clade based on 1000 bootstrap replicates. Tenkana is recovered as a sister genus to Hyllus and Telamonia in clade 3 and distantly from cf. Colopsus and Pancorius of clade 4.

Figures 2, 3. 

2 Maximum-likelihood tree from IQ-TREE analysis (best of 10 replicates) constrained for Tenkana clade (using clade 3 of the UCE tree, see Fig. 1) 3 Maximum-likelihood tree from IQ-TREE analysis (best of 10 replicates) without the constraint. The trees are inferred from a gene partitioned, concatenated nuclear 28S, 18S, H3 and mitochondrial COI genes. Numbers at the nodes are the percentage recovery of the clade based on 1000 bootstrap replicates and nodes without numbers suggests that those clades were not recovered in bootstrap analysis. Tenkana is recovered distantly (see Tenkana clade) from the type species of Colopsus, Colopsus cancellatus and other Sri Lankan Colopsus (see Colopsus clade) in both constrained and unconstrained trees.

Tenkana is nestled well within Plexippina as expected and recovered as the sister group to Hyllus C.L. Koch, 1846, and Telamonia Thorell, 1887 in clade 3 of the UCE phylogeny (see Fig. 1): (Tenkana, (Hyllus, Telamonia)). The type species of Colopsus and other members of the cancellatus group were found to be closely related to Pancorius Simon, 1902 (similarly to Kanesharatnam and Benjamin 2021). The manu species group is not closely related, however, to Pancorius and Colopsus. In the UCE phylogeny, it is recovered distantly from Pancorius and cf. Colopsus of clade 4 (see Fig. 1), while in unconstrained and constrained four-gene phylogenies, it is distant from Pancorius, cf. Colopsus, and the true Colopsus from Sri Lanka (Figs 2, 3). Their phylogenetic distance helps to explain the morphological differences between the manu group (non-glossy, non-elongate-bodied, tegulum with tegular lobe, medially located two ECPs) and the cancellatus group (glossy, elongate-bodied, without tegular lobe, laterally placed two ECPs).

Therefore, we propose Tenkana as a new genus to contain two species of the manu group currently placed within Colopsus and the new species described below.

Family Salticidae Blackwall, 1841

Subfamily Salticinae Blackwall, 1841

Tribe Plexippini Simon, 1901

Subtribe Plexippina Simon, 1901

 Tenkana jayamangali Caleb & Marathe, sp. nov.

https://zoobank.org/C4C0B4D7-20BF-4566-959B-C015940AF4E6

Figs 4–7, 8–15, 16–24

Kannada: ತೆಂಕಣ ಜಯಮಂಗಲಿ;

Devanagari: तेंकण जयमंगलि

Materials examined

India • Karnataka, Tumakuru; 13.3843°N, 77.2069°E; 987 m a.s.l.; 23 April 2023; coll. Y.T. Lohit & B.G. Nisha. Holotype: • ♂ (I/SP-48) in ZSIC, Paratypes: • 1 ♀ (I/SP-49) in ZSIC; • 1 ♂ (IBC-BX512); • 1 ♀ (IBC-BX513) in NCBS. Paratype: • 1 ♀ (IBC-BX511) in NCBS, 16 December 2023; coll. B.G. Nisha.

Etymology

The specific epithet ‘jayamangali’, a noun in apposition, is the name of a river originating in Devarayanadurga, Tumakuru, where this species was observed for the first time.

Diagnosis

The phylogenies recover Tenkana jayamangali as a sister species to T. arkavathi and T. manu. In the males of T. jayamangali, pale hairs occupy most of carapace surface area leaving small bald patch posteriorly, while in T. arkavathi and T. manu, pale hairs are gentler on carapace forming narrower bands on carapace laterally, tapering posteriorly. Ocular area of T. jayamangali covered with white hairs uniformly, while T. arkavathi has distinctive V-shaped bands and T. manu has bald ocular area. From ventrally, RTA can be seen extending much more laterally with slight bend sub-apically in T. jayamangali, while T. arkavathi with relatively short with prominent bend and T. manu with longer with no bend. Short sperm duct loop arising at 11 o’clock in T. jayamangali, while much longer sperm duct arising at 10 o’clock in T. arkavathi and T. manu). ECPs laterally placed T. jayamangali, while ECPs medially in T. arkavathi and T. manu).

Description

♂ (I/SP-48 in ZSIC). Total length 5.18; carapace 2.66 long, 2.09 wide; abdomen 2.52 long, 1.77 wide. Carapace, brown, white hairs laterally traversing posteriorly. In life, white hairs cover most of its surface area leaving small patch of bald black integument on thoracic slope. Tufts of thick bunch of hairs (‘eye lashes’) behind ALEs and below PMEs. In life, reddish-orange small hairs along the circumference of anterior eyes. Clypeus brown, length 0.19. Chelicerae brown. Legs brown, robust. Leg I and II ventrally fringed thickly with black hairs, but less densely on leg II. White leaf-like hairs interspersed prolaterally on patella and tibia of leg I; similar white hairs retrolaterally on femora of leg I and II and distal end and proximal and distal ends of leg III and IV femora. Leg measurements: I 5.66 (1.68, 1.16, 1.38, 0.88, 0.56); II 4.86 (1.62, 0.89, 1.05, 0.77, 0.53); III 5.54 (1.94, 0.94, 1.06, 0.95, 0.65); IV 5.56 (1.77, 0.76, 1.13, 1.19, 0.71). Leg formula 1432. Palp (Figs 4, 5, 8, 9) medium long membrane-accompanied embolus arising at 6 o’clock. Ovoid tegulum with prominent tegular lobe. RTA short, simple (Figs 5, 9). Abdomen brown and black mottlings on yellow integument, which is pronounced medially with band like appearance. Chevronous markings posteriorly. In life, bright orange coloured. Spinnerets brown, somewhat long.

Figures 4–7. 

Genitalia of Tenkana jayamangali sp. nov. 4 male left palp, ventral view (holotype I/SP-48) 5 ditto, retrolateral view (holotype I/SP-48) 6 epigyne, ventral view (paratype IBC-BX511) 7 vulva, dorsal view (paratype IBC-BX511). Scale bars: 0.2 mm. ECP, epigynal coupling pocket.

Figures 8–15. 

Photographs of genitalia and preserved bodies of Tenkana jayamangali sp. nov. Genitalia 8 male left palp, ventral view (holotype I/SP-48) 9 ditto, retrolateral view (holotype I/SP-48) 10 epigyne, ventral view (paratype IBC-BX511) 11 vulva, dorsal view (paratype IBC-BX511). Bodies: 12 male (holotype I/SP-48), dorsal view 13 ditto, ventral view 14 female (paratype IBC-BX511), dorsal view; 15, ditto, ventral view. Scale bars: 0.2 mm for genitalia; 1.0 mm for bodies. CO, copulatory opening; ECP, epigynal coupling pocket; MBE, membrane-bearing embolus.

♀ (IBC-BX511 in NCBS). Total length 5.24; carapace 2.53 long, 2.19 wide; abdomen 2.71 long, 1.83 wide. Carapace brown, more or less bald. White pale stripe posteriorly on thoracic slope. Tufts of thick bunch of hairs (‘eye lashes’) behind ALEs and below PMEs. In life, reddish-orange small hairs along the circumference of anterior eyes. Clypeus brown, length 0.16. Chelicerae brown. Legs brown, robust, largely bald. Leg measurements: I 5.36 (1.66, 1.05, 1.23, 0.85, 0.57); II 4.85 (1.60, 0.94, 0.97, 0.80, 0.54); III 5.68 (2.00, 1.02, 1.06, 0.98, 0.62); IV 5.57 (1.78, 0.82, 1.09, 1.21, 0.67). Leg formula 3412. Abdomen comparable as in male. In life, medial yellowish band surrounded by yellow and black mottlings. Epigyne (Figs 6, 7, 10, 11) medially located shallow two ECPs flanked by crescent shaped copulatory openings. Lamellar copulatory ducts join simple spermatheca ventrally.

Distribution

In addition to the type locality, iNaturalist observations (e.g. Mohan 2024; Raj 2024) appearing to represent this species are recorded around Bengaluru, Karnataka.

Natural history

Tenkana jayamangali was observed commonly in May; however, given the collecting of a female in December and iNaturalist observations, they may be adult year-round. Tenkana jayamangali were collected among dry leaf litter on the ground. A subadult was observed feeding on a bug nymph (Figs 23, 24).

Figures 16–24. 

Habitus of Tenkana jayamangali sp. nov. 16–18 male 19–21 female 22 male 23–24 subadult male feeding on a bug nymph. Photo credit: Nisha B.G. (16–21) and Lohit Y.T. (22–24).

Discussion

With Tenkana, the subtribe Plexippina now contains 36 genera; for India, the number of plexippines is 48 species in 19 genera (Maddison 2015; Marathe et al. 2024b; World Spider Catalog 2024). We continue to include Colopsus in the list of Indian plexippine genera, represented by Colopsus peppara Sudhin, Sen & Caleb, 2023. Colopsus peppara resembles Tenkana and Pancorius in body form; however, the features of the male palp could be attributed to Colopsus and Pancorius (simple round tegulum lacking tegular lobe and short embolus). What makes it puzzling is its epigyne, which has a medially located single ECP and does not match any of the three genera. The puzzling morphological features of C. peppara, along with a lack of clear synapomorphies for Colopsus, Tenkana, and Pancorius, compound the challenge of placing C. peppara definitively within a plexippine genus. Therefore, we propose maintaining the status quo until we can determine its placement using molecular data.

The authors have declared that no competing interests exist.

No ethical statement was reported.

Funding to WPM was provided by an NSERC Canada Discovery Grant. Museum and lab work in NCBS was supported by an NCBS research grant to KK.

KM, BGN, CCM, and YTL participated in the fieldwork. KM, JTDC and KK managed specimens. KM did the molecular work. KK provided space and resources for molecular work. KM and WPM planned the molecular phylogenetic study, processed the data, and performed the phylogenetic analyses. KM and JTDC studied the morphology of T. jayamangali. JTDC made decisions about new species. KM did the drawings of T. jayamangali genitalia. KM, WPM, and JTDC made decisions about the new genus. KM studied the other Tenkana spp. morphologically for generic diagnosis and description. KK helped with microscopy. KM wrote the first draft of the manuscript, excluding the species description. JTDC wrote the first draft of the species description. KM and WPM finalized the first draft. All other authors assisted with additions and corrections to the manuscript.

Kiran Marathe https://orcid.org/0000-0002-7364-3475

John T. D. Caleb https://orcid.org/0000-0002-9471-9467

Wayne P. Maddison https://orcid.org/0000-0003-4953-4575

B.G. Nisha https://orcid.org/0009-0006-7603-0785

Chinmay C. Maliye https://orcid.org/0009-0002-4050-9197

Y.T. Lohit https://orcid.org/0009-0007-8866-2226

Krushnamegh Kunte https://orcid.org/0000-0002-3860-6118

All of the data that support the findings of this study are available in the main text.