Phylogenetic placement of Carrhotus Thorell, 1891 with three new species from Sri Lanka (Araneae: Salticidae)

. The Sri Lankan species of the genus Carrhotus Thorell,1891 have remained taxonomically unrevised. The present study reviews all species of the genus in the island. Here we describe and diagnose three new species: Carrhotus albosetosus sp. nov. ( ♀ ), C. atratus sp. nov. ( ♂♀ ) and C. lobatus sp. nov. ( ♂ ). Further, C. silanthi Caleb, 2020 is reported in Sri Lanka for the ﬁ rst time. Carrhotus taprobanicus Simon, 1902 and C. viduus (C.L. Koch, 1846) are redescribed based on material from Sri Lanka. A key to the Carrhotus species in Sri Lanka is given. of Carrhotus with a lighter pattern. Chelicerae unident with 2 promarginal teeth and 1 retromarginal tooth. RTA not prominent; cymbium in some species with apical process; tegulum oval, expands beyond apical border of tibia (PLP); embolus stout, set laterally on top of tegulum, wider base. Female epigyne moderately sclerotized, strongly haired. CD well sclerotized, straight, curved or coiled. Spermatheca one-chambered, oval to kidney-shaped . FD is well sclerotized. Accessory glands on external spermathecal walls. Legs I and II robust with scopulae, III and IV more delicate. Hairs and bristles blackish. Leg formula: 1-2-4-3


Introduction
Jumping spiders (Salticidae Blackwall, 1841) is the most diverse spider family and includes species commonly encountered in many habitats (World Spider Catalog 2022). This is also the case in Sri Lanka, where they are remarkably diverse. Currently, 86 species placed in 50 genera, with a large endemic component, are known from the island (Benjamin 2004(Benjamin , 2010(Benjamin , 2015Benjamin & Bambaradeniya 2006;Edwards & Benjamin 2009;. However, recent fi eldwork conducted around the country suggests that this might only be a small fraction of the island's jumping spider diversity.
The genus Carrhotus was erected by Thorell in 1891 with Plexippus viduus C.L. Koch, 1846 as its type species. It currently contains 33 species, mostly from the Oriental region (World Spider Catalog 2022). The fi rst species of the genus from Sri Lanka, the endemic Carrhotus taprobanicus Simon, 1902, was nominally described, sans illustrations a few years thereafter (Simon 1902). Recently, Carrhotus viduus (C.L. Koch, 1846) was recorded from the island by Prószyński (2009), with descriptions based on a pair of specimens collected by Fruhstorfer in 1889. Further, a very recent revision of the Indian Carrhotus provides descriptions of three new species and updates the taxonomy of the genus. One species newly described therein, C. silanthi Caleb, 2020, is recorded from Sri Lanka for the fi rst time (Caleb et al. 2020). Maddison (2015) based on an explicit phylogenetic framework placed Carrhotus in the tribe Salticini Blackwall, 1841 of the subfamily Salticinae Blackwall, 1841. Previously, Carrhotus was classifi ed in the Hylleae by Simon (1903) and later considered a Chrysillinae based on the conformation of its copulatory organs (Prószyński 1976(Prószyński , 2017. However, these placements were not tested within a phylogenetic framework, until recently. In this paper we review the taxonomy of Carrhotus spp. of Sri Lanka and describe three new species. Further, we infer the phylogenetic placement of the Carrhotus spp. of Sri Lanka using two target genes.

Morphology
Methodology and taxonomic descriptions are based on the format of , and Benjamin (2004Benjamin ( , 2010. Specimens for the morphological study were preserved in 70% ethanol and identifi ed using an Olympus SZX7 stereo microscope. Female genitalia were excised using insect pins and digested with the Sigma Pancreatin lp 1750 enzyme complex in a solution of sodium borate (Dingerkus & Uhler 1977). Male palps and epigynes were cleared with methyl salicylate and mounted for study. Drawings of male palps, epigynes and vulvae were made with the aid of a drawing tube attached to a Leica DM3000 LED compound microscope. Either a Nikon D80 or a D7000 camera with a macro lens was used to take photographs of live spiders. Photographs of palps, epigynes and intact spiders were taken with a Leica MC170 HD camera mounted on a Leica M205C stereo microscope using Leica Application Suite software (Leica Microsystems Limited, Germany). Images were merged with Helicon Focus image stacking software (ver. 6, Helicon soft Ltd). Images were then edited with Adobe Photoshop CC and assembled using Adobe Illustrator CS6. All measurements are in millimeters. Body length was measured as carapace length plus abdomen length (excluding spinnerets). Types and other specimens of the new species described herein are currently deposited in the National Institute of Fundamental Studies, Kandy, Sri Lanka (NIFS). For a complete list of taxonomic references for the previously described species see World Spider Catalog (2022).

Gene targets and primers
A multilocus molecular approach was used for this study and the target loci were selected based on prior molecular phylogenetic studies of salticids (e.g., Hedin & Maddison 2001;Bodner & Maddison 2012). A partial fragment of the mitochondrial protein-encoding gene Cytochrome c oxidase subunit I (CO1 -534 bp) and partial fragments of the nuclear gene, 28S ribosomal RNA (28S -796 bp) were amplifi ed. Details of each primer pair used, primer sequences, annealing temperatures and related references are given in Table 2.

DNA extraction, PCR and sequencing
Genomic DNA was extracted using the Qiagen DNeasy Tissue kit (Qiagen Inc., Valencia, CA, USA) following the manufacturer's protocols. Depending on size, either the fi rst leg only, or the fi rst and second legs from each specimen was removed. The remainder of the vouchered specimen is stored in 70% ethanol and deposited in the NIFS arachnid collection. Polymerase Chain Reaction (PCR) was carried out using either the Multiplex PCR kit (Qiagen) or the PuReTaq Ready-To-Go TM PCR beads (GE Healthcare, UK). For the former, the total reaction mixture consisted of 20 μL, including 2.5 μL undiluted DNA template, 1.6 μL forward and reverse primers, 10 pmμL -1 of Q-Solution, 10 μL 'Multiplex PCR Master Mix' containing hot start Taq DNA Polymerase, buffers and 2.3 μL of water (all the components come with the Multiplex kit). In the case of PCR beads, the total reaction mix was 25 μL and included 13 μL of ultrapure RO (Reverse Osmosis) water, 1 μL of each of the forward and reverse primers (10 pmμL -1 ) and 10 μL undiluted DNA template. A negative control (minus the template) was included to test for contamination during PCR runs. The primers used for amplifi cation and their sources are given in Table 2. PCR products were verifi ed on a 1% agarose gel and purifi ed using the Gene Clean TM Turbo kit (MP Biomedicals, LLC, USA). All purifi ed PCR products were Sanger sequenced (Sanger et al. 1977) in both directions by Macrogen (Seoul, South Korea).

Phylogenetic analysis
Sequences were assembled and edited using the Geneious ver. 11.0 software package. Sequence alignment was also done with the same software using default parameters and then edited manually using Mesquite ver. 3.51 (Maddison & Maddison 2018). The protein-coding COI was easily aligned. The alignment of the non-coding 28S was further refi ned manually in Mesquite. The fi nal alignments of the two gene fragments were then concatenated using the same program. Details of the fragments are provided in Table 2.
A model-based (maximum likelihood, ML) approach was used to infer the phylogenetic relationships of the targeted taxa. The best-fi t model for likelihood analysis was searched by running the 'fi nd best DNA/   Boc et al. 2012), were performed for maximum likelihood analysis using the non-parametric bootstrap program. The parameters used were as follows: substitution model (GTRGAMMAI), algorithm executed (Hill climbingdefault), and the bootstrap was estimated by 1000 alternative runs on distinct starting trees.

Phylogenetic results
The assembled matrix of the concatenated fragments included 38 taxa (20 ingroups, 18 outgroups) and the total length was 1330 bp. The length of the two individual sequenced fragments, excluding the primers, was as follows: 28S, 796 bp and COI, 534 bp. The best fi t model for the combined data matrix generated using MEGA was GTR+G+I and, accordingly, GTRGAMMAI was selected as the compatible model in RAxML.
The phylogenetic tree resulting from ML analysis of the combined data matrix is presented in Fig. 1

Diagnosis
Carrhotus can be readily distinguished from other salticid genera by the combination of the following characteristics: RTA stout, tegulum with at least a retrolateral and/or proximal lobe, embolus broadbased, tip mostly stout (

Description
Medium sized spiders. Cephalothorax longer than wide, eye fi eld shorter than 50% of total cephalothorax length. PME mid-way between anterior and posterior laterals. Abdomen egg-shaped, beige to brown with a lighter pattern. Chelicerae unident with 2 promarginal teeth and 1 retromarginal tooth. RTA not prominent; cymbium in some species with apical process; tegulum oval, expands beyond apical border of tibia (PLP); embolus stout, set laterally on top of tegulum, wider base. Female epigyne moderately sclerotized, strongly haired. CD well sclerotized, straight, curved or coiled. Spermatheca one-chambered, oval to kidney-shaped . FD is well sclerotized. Accessory glands on external spermathecal walls. Legs I and II robust with scopulae, III and IV more delicate. Hairs and bristles blackish. Leg formula: 1-2-4-3 (J astrzebski 1999).

Composition in Sri Lanka
Currently, the genus consists of 33 species worldwide [for a full list see World Spider Catalog (2022)].
Carrhotus albosetosus sp. nov. urn:lsid:zoobank.org:act: Diagnosis Females of C. albosetosus sp. nov. can be separated from other congeners by the white hairs on the legs ( Fig. 2A-B). The epigyne of this species is close to that of C. silanthi; however, it differs by the absence of prominent CD and FD as well as in the position of the opening of the accessory glands, which open at the 2'o clock position at the anterior region of the lateral walls of the spermathecae (Fig. 4A-B).

Etymology
The specifi c epithet is derived from a combination of Latin words; 'albi' meaning 'white' and 'comus' meaning 'legs', loosely referring to the white legs/white leg setae. COLOR AND BODY. Live spider with clypeal region brown; covered with iridescent hairs providing a metallic sheen to green colour; lateral margins of carapace lined by with thin line of white hairs; posterior region black ( Fig. 2A-B). ALEs and PLEs surrounded by blackish orbital setae. Chelicerae dark brown with curved outer margins and excavated inner margins: two promarginal and one retromarginal teeth. Sternum oval, brownish; labium and maxillae reddish-brown, with paler outer margins. Abdomen ovoid, densely covered with rusty brown hairs; anterior margin covered with a thin line of white hairs; pattern present on black background with pair of two white spots anteriorly and pairs of transverse stripes and two pairs of white spots following posteriorly. Mid-dorsum covered with scales of metallic sheen; venter yellowish brown, with a thin dark brown scattered pattern on median region. Spinnerets blackish, covered with a patch of white hairs dorsally (Fig. 3A-B).
EPIGYNUM. Epigyne with a pair of simple copulatory openings and short copulatory ducts diverge to join semioval-shaped spermathecae. Accessory glands open at 2'o clock position on anterior region of lateral walls of spermathecae (Figs 3C, 4A-B).

Distribution and habitat
Known only from the two localities mentioned above. The spiders were collected by beating foliage up to a height of 2 m. This species occurs in the lowland secondary rainforest of the dry zone and submontane forest of the central highlands of Sri Lanka (Fig. 17).   Diagnosis Males of C. atratus sp. nov. can be separated from those of other congeners by their triangular-shaped PLP and cone-shaped anticlockwise directed embolus tip (Figs 6E-F, 7A-B). The palp of C. atratus closely resembles that of C. erus Jastrzebski 1999; however, its PLP is relatively short, rounded and RTA is not pointed. RTA is smaller and tapered (Fig. 6F) than in other Carrhotus species from Sri Lanka. Females are distinguished by the prominent hood at the CO and hook-shaped CD (Fig. 7C-D).

Etymology
The specifi c epithet is derived from the Latin word 'atratus' meaning 'darkened', loosely referring to its blackish body colour.

Material examined
Holotyp e SRI LANKA • ♂; Northern Province, Jaffna District, Mandaitivu; 09°61′67″ N, 79°99′22″ E; 12 m a.s.l.; 12 Jan. 2020; S.P. Benjamin et al. leg  COLOR AND BODY (live spiders). Carapace black, sparsely covered with black hairs; lateral margins sparsely covered with white hairs; posterior region black ( Fig. 5A-B, E-F, I). Clypeal region blackish; eyes surrounded by blackish orbital setae. Chelicerae black, curved outer margins, excavated inner margins; two promarginal and one retromarginal teeth. Sternum oval, blackish; labium and maxillae blackish brown, paler outer margins. Abdomen ovoid, sparsely covered with black hairs; anterior margin sparsely covered by white hairs; no prominent pattern present on posterior region, beige pairs of transverse lines covered with white hairs on mid dorsum. Venter blackish with a very broad dark black median region covered with two thin greyish longitudinal lines. Spinnerets blackish, covered with a patch of black hairs dorsally (Fig. 6A-B). Leg I robust; femora I-IV black dorsally; patellae and tibiae black; tarsi and metatarsi of all legs blackish.
PALP. Dark black colour and embolus long, bow-shaped and tip directed anticlockwise, its base visibly separated from tegulum (Figs 6E-F, 7A-B). COLOR AND BODY (live spiders). Same as in male except for the following. Clypeal region covered with iridescent hairs providing a metallic sheen; lateral margins of carapace lined by a thin line of white hairs; posterior region black ( Fig. 5C-D, G-H). ALEs and PLEs surrounded by blackish orbital setae. Sternum oval, brownish; labium and maxillae reddish-brown, with paler outer margins. Abdomen ovoid, densely covered with rusty brown hairs; anterior margin covered with a thin line of white hairs; pattern present  on black background with pair of two white spots anteriorly and pairs of transverse stripes following posteriorly. Mid-dorsum covered with scales of metallic sheen; venter yellowish, with a broad dark brown median region (Fig. 6C-D).
EPIGYNUM. Epigyne with a pair of copulatory ducts with a loop or fl ap-like structure arising from elongated pear-shaped spermathecae, whose latter part is narrowing. Accessory glands open at 3 o'clock position at narrowing anterior region of lateral walls of spermathecae (Figs 6G, 7C-D).

Distribution and habitat
This species occurs in the mangrove forest, in the arid zone and lowland secondary rainforest of Sri Lanka (Fig. 17). Specimens were collected by beating foliage up to a height of 2 m.

Etymology
The specifi c epithet is derived from the Latin word 'lobatus' meaning 'lobed', loosely referring to its distinctive proximal lobe of the tegulum. robust; femora I-IV dark brown dorsally; patellae and tibiae dark brown; tarsi and metatarsi of all legs dark brown. Abdomen oblong, sparsely covered with black hairs; anterior margin covered with long black scales with sparse white hairs making an arc; no prominent pattern present on posterior region, beige pairs of transverse lines and pair of beige spots covered with white hairs on mid dorsum; venter blackish brown with a blackish brown median region. Spinnerets blackish brown, covered with a patch of white hairs dorsally (Fig. 8A-B).

Holotype
PALP. Dark brown; embolus short and hook-shaped and embolus tip directed in the clockwise path; its base separated from the tegulum. The double-lobed PLP is very distinctive compared to that of the other species considered here. RTA long and angle between the RTA and tibia is about 45º (Fig. 9A-B).

Distribution and habitat
This species occurs in the montane and submontane forests of the central highlands of Sri Lanka with limited distribution (Fig. 17). Specimens were collected by beating vegetation up to a height of 1-2 m.  Caleb, 2020 Figs 10A-F, 11A-H, 12A-D, 17 Carrhotus silanthi Caleb in Caleb et al., 2020: 58, Figs 33-56.

Diagnosis
Males of C. silanthi can be separated from other congeners by having the RTA markedly hook-shaped, slanted ventrally, and the distally pointed acuminate embolus (Fig. 12A-B). Females can be readily separated from those of C. albosetosus sp. nov. by the prominent ML and the position of the opening of the accessory glands on the internal spermathecal walls, close to the base of the copulatory ducts (Figs 11G, 12C-D).

Type material
Carrhotus silanthi Caleb in Caleb et al., 2020: 58, Figs 33-56 ( male   COLOR AND BODY. In live spiders, both sexes look similar in general body colour pattern and are clothed with iridescent hairs. Colour varies from metallic sheen to bronze-green as shown in Fig. 10A-F. Carapace: prosoma reddish-brown covered with iridescent hairs providing a metallic sheen; lateral margins of carapace lined by broad patches of white hairs; posterior region black. Hairs sparse. Sternum oval, brownish; labium and maxillae dark brown, with paler outer margins. Clypeal region brownish; eyes surrounded by yellowish orbital setae. Chelicerae dark brown with curved outer margins and excavated inner margins: two promarginal and one retromarginal teeth. Abdomen ovoid, densely covered with rusty brown hairs; anterior margin covered by white scales; pattern present on black background with pair of white spots anteriorly and pairs of transverse stripes following posteriorly (Fig. 11A-B). Middorsum covered with scales of metallic sheen; venter yellowish, with a broad dark brown median region. Spinnerets brownish, covered with a patch of white hairs dorsally. Leg I robust; femora I-IV dark brown dorsally; patellae and tibiae dark brown; tarsi and metatarsi of all legs dark brown. PALP. Dark brown; embolus short and thick with a blunt tip and embolus tip directed in clockwise path, its base separated from tegulum; bulbus with PLP; RTA markedly hook-shaped bent ventrally and angle between RTA and tibia about 45º (Figs 11E-F, 12A-B). COLOR AND BODY. Colouration pattern as in male, but differs as follows: AMEs surrounded by yellow orbital setae; clypeus covered with white hairs; carapace with white hairs making an arc behind PLEs. Abdomen ovoid, densely covered with rusty brown hairs; anterior margin covered by white scales; pattern present on black background with pair of white spots anteriorly and pairs of transverse stripes following posteriorly (Fig. 11C-D). Mid-dorsum covered with scales of metallic sheen; venter yellowish, with a broad dark brown median region. Spinnerets brownish, covered with a patch of white hairs dorsally. Legs with white border on distal margin of femur I-IV; other segments covered with sparse white hairs and black annulations alternatively.
EPIGYNUM. Epigyne with a pair of simple copulatory openings placed in small oval, yellowish depressions; copulatory ducts short, diverge to join stomach-shaped spermathecae. Accessory glands open on internal spermathecal walls close to base of copulatory ducts (Fig. 12C-D).

Intraspecifi c variation
The palpal conformation of NIFS_SAL_1116 closely resembles that of C. silanthi. However, it differs by having the PLP shorter and less rounded (Fig. 11F) vs oval-shaped PLP in C. silanthi (Fig. 11E), and a short, cone-shaped embolus (Fig. 11F) in NIFS_SAL_1116 vs a thinner embolus in C. silanthi (Fig. 11E).

Distribution and habitat
This species occurs in the lowland secondary rainforests of the dry and wet zones of Sri Lanka including the Ussangoda dry zone National Park in Hambantota District in the Southern Province (Fig. 17). Specimens were collected by beating vegetation up to a height of 1-2 m.  Simon, 1902 Figs 2E-F, 13A-G, 14A-D, 17

Diagnosis
Males of C. taprobanicus can be separated from those of C. viduus by their body colour pattern with no longitudinal stripes on the carapace and abdomen (present in C. viduus) (Fig. 15A, C). Further, the cheliceral front part is more wrinkled in C. viduus (Fig. 13B). The palpal conformation of C. taprobanicus closely resembles that of C. viduus; however, it can be distinguished by the shorter embolus and its rounded base (Fig. 13E). Further, the RTA is more slanted ventrally in C. taprobanicus (Fig. 14A-B). Females can be separated from those of C. viduus by the narrower ML and stouter CD and FD (Fig. 14C-D).

Type material
Carrhotus taprobanicus Simon, 1902 COLOR AND BODY. Live spider with carapace blackish-brown, covered with iridescent hairs providing a metallic sheen; lateral margins of carapace lined by broad patches of white hairs; posterior region black ( Fig. 2E-F). Clypeal region blackish; eyes surrounded by yellowish orbital setae. Chelicerae brown with curved outer margins and excavated inner margins: two promarginal and one retromarginal teeth. Sternum pentagonal, blackish brown; labium and maxillae yellowish brown, with paler outer margins. Abdomen ovoid, densely covered with black hairs; lateral sides of anterior margin with two white longitudinal lines; pattern present on black background with three pairs of white transverse stripes lateral margins of mid-dorsum. Mid-dorsum covered with scales of metallic sheen; venter yellowish, with a broad black median region covered with two broad whitish longitudinal bands. Spinnerets brownish, covered with a patch of white hairs dorsally (Fig. 13A-B). Leg I robust; femora I-IV dark brown dorsally; other segments covered with sparse black hairs and black annulations. PALP. Dark brown; embolus thinner and cone-shaped, its base separated from tegulum. PLP shorter and similar to that of C. viduus, angle between RTA and tibia about 45º vs about early 20º in C. viduus (Figs 13E-F, 14A-B).  COLOR AND BODY. Abdomen ovoid, sparsely covered with black hairs; lateral sides of anterior margin with four pairs of white herringbone-shaped pattern present on brown background with two pairs of white spots in mid-dorsum. ALEs surrounded by blackish orbital setae; clypeus covered with white hairs (Fig. 13C-D).
EPIGYNUM. Epigyne with a pair of copulatory openings, long copulatory ducts move anteriorly and almost sub-parallel to elongated, pear-shaped spermathecae. Accessory glands open at 2 o'clock position of mid-region of lateral walls of spermathecae (Figs 13G, 14C-D). Resembles the epigynes of C. viduus but differs in relative proportions of spermathecae, rather kidney-shaped in C. taprobanicus ( Fig. 14C-D), PEB less curved inwards and margins of CO less sclerotized than in C. viduus (Fig. 13G).

Remarks
Our identifi cation is based on Simon's (1902) description of the male and female syntypes. Though we were unable to examine the types of C. taprobanicus, the description by Simon (1902) was clear enough for an unambiguous identifi cation of the species, which we translate here to English: "♂. Length 6.5 mm. Cephalothorax darkened upper part, bronze-colour-haired, white hair belts at the thoracic margins, short clypeus and yellowish-red hairs around the eyes. Chelicerae long, planar and hairless with bronze and leathery transverse folds; curved outer margins and excavated inner margins: promarginal and one retromarginal teeth. Sternum; black, white-haired. Abdomen shorter and ovalshaped. Venter greyish and covered on both sides with white margins. Legs I, II darkened, tarsi lightened, metatarsi, lower tibia very slightly black-haired, patella inwardly white-haired. Legs: coxa, trochanter, metatarsi darker, metatarsi dark annulated.
♀. Length 7.8 mm. Cephalothorax black, bronze-colour-haired, reddish-white hair belts at the thoracic margins and around the eyes. The males can be distinguished from those of C. viduus (C.L. Koch, 1846) by their body colour pattern with no longitudinal stripes on the carapace and abdomen and the cheliceral front part is more wrinkled in C. viduus".

Distribution and habitat
This species occurs in the montane and submontane forests of the central highlands of Sri Lanka (Fig. 17). Specimens were collected by beating vegetation up to a height of 1-2 m.

Diagnosis
Males of this species are distinguishable by the presence of white longitudinal stripes on the carapace and abdomen, as well as the relatively larger retromarginal cheliceral tooth. The palpal conformation of C. viduus closely resembles that of C. taprobanicus; however, it can be distinguished by the longer, ventrally projecting embolus and its oval base (Fig. 16A). Further, the RTA is less slanted ventrally in C. viduus (Fig. 16A-B). Females can be separated from those of C. taprobanicus by the broader ML and longer CD and FD (Fig. 16C-D). Females can be readily distinguished from those of C. silanthi by the kidney-shaped spermathecae (Figs 12C-D, 16C-D).

Type material
Plexippus viduus C.L. Koch, 1846: 2 ♂♂; depository unknown. Our identifi cation is based solely on the description and illustrations of Prószyński (2009). He mentioned two possible specimens currently deposited in ZMB. However, their geographic origin is ambiguous (see Prószyński 2009).

Remarks
According to Sudhin et al. (2021), C. tholpettyensis Sudhin, Nafi n, Caleb & Sudhikumar, 2021 differs by the absence of a pair of white longitudinal stripes found on the abdomen of male C. viduus. The mentioned minor gentilic differences could easily be attributed to intraspecifi c variation. Thus, we consider C. tholpettyensis as a possible junior synonymy of C. viduus.

Material examined
SRI LANKA -

Description
Male MEASUREMENTS. TL 6.85, PL 2.58, PW at PLEs 2.31, AL 1.35, AW 0.87. Eye fi eld: diameter of AME 0.28; ALE 0.18; PME 0.1; PLE 0.14; PME-PME 0.82; PLE-PLE 0.84; ALE-PME 0.32; ALE-PLE 0.42. COLOR AND BODY. Live spider with carapace robust and relatively broad, blackish-brown covered with long black hairs sparsely and two longitudinal stripes of white hairs (Fig. 2C-D). Sternum oval, brown; labium and maxillae yellowish brown, with paler outer margins. Clypeal region brownish; eyes surrounded by blackish orbital setae. Chelicerae brown with curved outer margins and excavated inner margins: two promarginal and one retromarginal teeth (Fig. 15B). Abdomen blackish brown, ovoid with two longitudinal belts of white hairs and entire surface covered with a few whitish hairs. Middorsum with four median beige spots in different shapes, covered with black scales; venter greyish with a blackish-brown median region covered with two broad whitish longitudinal bands. Spinnerets brown, covered with a patch of black hairs dorsally. Leg I robust; femora I-IV dark brown dorsally; patellae and tibiae dark brown; tarsi and metatarsi of all legs dark brown. PALP. Dark brown; embolus short and thick with blunt tip; bulbus with PLP; RTA only slightly bent ventroapically, curved and claw-like (Figs 15E-F, 16A

Distribution and habitat
This species occurs in the mangrove forests in the arid zone and lowland secondary rainforests of the dry and wet zones of Sri Lanka (Fig. 17). Specimens were collected by beating vegetation up to a height of 1-2 m.
Key to the species of Carrhotus in Sri Lanka this study. We are indebted to N. Kanesharatnam and Dilini Bopearachchi for assistance with molecular laboratory work. Thanks to the Department of Wildlife and Forest Department of Sri Lanka for granting permits for fi eldwork. We also thank two anonymous reviewers for their suggestions and comments that improved the manuscript.