Origin and diversification of free-living stick spiders of Sri Lanka including the description of four new species of Rhomphaea L. Koch, 1872 and two new species of Neospintharus Exline, 1950

Sri Lanka is a biologically diverse South Asian island, and together with the Western Ghats (Southern India) is one of the 36 world’s most biologically diverse areas. Here, we investigated the origin and diversification of Rhomphaea and Neospintharus of Sri Lanka using sequences of three genes: mitochondrial cytochrome c oxidase I (COI) and 16S rRNA (16S); and nuclear 28S rRNA (28S). Our phylogeny included 32 taxa (30 ingroup and 2 outgroup). We used Bayesian Inference and maximum likelihood methods to reconstruct the placement of species, divergence time estimations and their foraging behavior with an emphasis on species from Sri Lanka. Our phylogenetic hypothesis support the monophyly of Argyrodinae as well as the monophyly of Rhomphaea, where Rhomphaea is a sister group of Neospintharus. Further, our analysis also suggests that Sri Lanka was colonized by Argyrodinae several times. Additionally, the following new species are described: Rhomphaea shanthi sp. nov., Rhomphaea jacko sp. nov., Rhomphaea martini sp. nov., Rhomphaea marani sp. nov., Neospintharus kandelensis sp. nov. and Neospintharus ohiyiaensis sp. nov.


Introduction
Sri Lanka is a biologically diverse South Asian island. Sri Lanka together with the Western Ghats (Southern India) is one of the 36 world's most biologically diverse hotspots [1,2]. Large continental islands like Sri Lanka with faunas derived from a mixer of recent colonization by mainland species and long term in situ diversification have not been well studied [3,4]. However, such studies might offer new perspectives on processes generating island biodiversity [3,4]. Further, short-range endemics with very restricted distributions, as often found in Sri Lanka, may prove to be important flagship taxa for monitoring the effects of climate change and other threats on forest habitats [5].
Theridiidae Sundevall, 1833 commonly known as comb-footed spiders, is one of the largest families within Araneae comprising 2536 species classified in 125 genera [6]. Theridiids are climatic regions of Sri Lanka. Specimens were collected by hand or by beating vegetation up to around two meters. The collected spiders were preserved in 70% or 100% ethanol for morphological or molecular downstream application. Taxon sampling consist of 32 terminal taxa comprising of 30 ingroup taxa (14 were newly sequenced for this study) of the subfamily Argyrodinae and two outgroup taxa (Table 1). We included 14 new terminals: 7 specimens of R. shanthi sp. nov., 2 specimens each of R. marani sp. nov. and N. kandelensis sp. nov., 1 specimen each of R. jacko sp. nov. and R. martini sp. nov. Additional sequences were obtained from [18,19] to represent a more geographically evenly distributed sample. Accession numbers for all sequences as well as locality information are given in Table 1.

Morphology
Specimen preserved in 70% alcohol were examined using a Leica S9E binocular stereomicroscope (Leica Microsystems Limited, Wetzlar, Germany). Male palps (left) were dissected and immersed in Kaiser's glycerol gelatin (Merck KGaA, Darmstadt, Germany), slide mounted, observed and illustrated with the aid of Leica DM3000 LED stereo microscope with an attached drawing tube. Highly sclerotized or darker areas of palps and epigynum were shaded with an HB pencil. The female epigastric region was dissected and digested in a pancreatin solution for about 3-7 days, slide-mounted and illustrated as described above. Digital images of the specimens were taken with a Leica MC170 HD camera mounted on a Leica M205C stereomicroscope using the software package Leica application suite, LAS version 4.6.2. Acquired image stacks of different depths (20-50 images per stack) were assembled using Helicon Focus (version 6, Helicon Soft Ltd) to create a single image with the entire specimen in focus. Final images were edited using Adobe Photoshop Version CS6. Body length was considered as total length of prosoma + total length of opisthosoma (excluding spinnerets). In case of Rhomphaea, as their opisthosoma is elevated and folded, opisthosoma length was measured in two parts, anterior opisthosoma length (range from the point of pedicel in opisthosoma-spinnerets) and posterior opisthosoma length (point of spinnerets-posterior opisthosoma tip). Leg measurements are given in the following order: total (femur, patella, tibia, metatarsus, tarsus). Descriptions of morphological terminology follows Agnarsson [9]. All measurements are in millimetres. Types and other specimens of the new species described herein are currently in the NIFS and will be deposited in the National Museum of Sri Lanka, Colombo.

Molecules
Partial fragments of the 28S ribosomal RNA (28S) and mitochondrial protein-coding gene cytochrome c oxidase subunit 1 (COI) and 16S ribosomal RNA (16S) were amplified. COI and 16S gene regions are more suitable to resolve more recent evolutionary events, whereas 28S is more effective in resolving deeper nodes in phylogenetic trees [20]. Details of each primer pair used, expected amplicon length (bp, number of base pairs) annealing temperature/time, primer sequences and related references are given in Table 2.
Genomic DNA was extracted from 100% ethanol-preserved leg tissue using DNeasy 1 Blood and Tissue Kit (Qiagen, Hilden, Germany). Polymerase chain reaction (PCR) was carried out using primers used in previous studies [20][21][22][23]. All sequences were edited and aligned using Genious 11.1.5 and Mesquite v 3.51 [24]. The protein coding CO1 sequences were aligned easily. 16S and 28S sequences were subsequently treated with Gblock 0.91b [25] to cull ambiguous positions. Gblock parameters were defined as follows: minimum number of sequences for a conserved position (50%), maximum number of contiguous non-conserved positions (10), minimum number of sequences for a flanking position (46), minimum length of a block (2), allowed gap positions (With Half) and similarity matrices were used. All sequences have been deposited in Gene Bank and their accession numbers are given in Table 1.

Phylogenetic analysis
Bayesian and Likelihood methods were used for single and concatenated gene matrices. Prior to likelihood and Bayesian analyses, Partition finder software v 2.1.1 [30] was run to find the best fit model for each partition using linked Bayesian Information Criterion (BIC) ( Table 3). We tested the codon position specific models for each gene at 1 st , 2 nd , 3 rd positions. The evolutionary models for each codon positions were then applied in the subsequent Bayesian and Maximum likelihood analysis. The results of model selection and priors used for partitioned are listed in the Table 3. Phylogenetic trees were inferred using Bayesian method in MrBayes v 3.2 [31] and Maximum likelihood method in IQtree v 1.6.12 [32]. We conducted these analyses for each gene separately and also for the concatenated data matrix. Each Bayesian analysis comprised of two independent Markov chain Monte Carlo (MCMC) chains and 1×10 6 generations per run. In all MrBayes analyses we discarded the first 25% of the sampled trees as burn in. We sampled a tree every 1×10 3 generations in each analysis, then visually inspected the likelihood scores and posterior probability scores of trees in Fig Tree v 1.4.2 [33]. Maximum likelihood analysis for each dataset were conducted in IQtree with 1000 bootstrap replications.

Divergent time estimation
Estimates of divergence times were computed on the concatenated data matrix using BEAST v. 1.8.2. Beauti v. 1.8.2 [34] was used to generate the XML file. The clock models and substitution rate were unlinked and substitution models for each gene were set up as in MrBayes. The fossil information from a previous study was used to calibrate the tree [10,18]. The oldest date at the node representing the lineage of members of genera Rhomphaea and Neospintharus was set at 12.5 Mya with normal distribution prior and an arbitrary standard deviation of 0.01. We performed two independent runs of the analysis and in each run the first 25% of the trees were removed as burn-in. The MCMC chain length of each run was 1×10 7 with the frequency of 1×10 3 . The generated tree files were then used to generate a maximum clade credibility tree using TreeAnnotator v. 1.8.2 [34]. The final time-scaled tree was visualized using Figtree.

Distribution of species
The occurrence of the sampled species in Sri Lanka was mapped in QGIS v 3.14.16 [35] using the coordinates of sampled localities obtained from field works. Localities with uncertain coordinates were resorted to approximation of the points using Google Earth [36]. The list of abbreviations used in the study is given in Table 4.

Nomenclatural acts
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix "http://zoobank.org/". The LSID for this publication is: urn:lsid:zoobank.org:pub: urn:lsid:zoobank.org:pub:4399F1E7-0B98-4A9A-B098-ECA7FA18E6BB. The electronic edition of this work was published in a journal with an ISSN, and has been archived and is available from the following digital repositories: PubMed Central, LOCKSS [author to insert any additional repositories].

Phylogenetic analysis
The assembled matrix of the concatenated mitochondrial and nuclear markers included 69 sequences of 22 taxa; 22 of these sequences were newly generated for this study. The lengths of targeted fragments after excluding primers and Gblock treatment were as follows: 16S~536bp, 28S~544bp and CO1~1067bp. The assembled matrix of mitochondrial and nuclear markers includes 32 taxa (30 ingroups and 2 outgroups). The total length of final matrix was 2147bp.
The best-fit model for all data matrixes are given in Table 3.
The phylogenetic tree resulting from the Bayesian analysis of the combined data matrix is presented in Fig 1. The phylogenetic tree resulting from the ML analysis is presented in S1 Fig. Topologies of all trees are summarized in Fig 2. The trees resulting from both analyses are generally concordant with each other as well as with previous analyses [18,19] and are well-supported, with respect to our taxa of interest. Thus, the topology of both the Bayesian and ML trees are discussed together; the differences are presented in the summary tree (Fig 2). Both trees resulting from the analysis of the concatenated data recover the monophyly of Rhomphaea as well as Ariamnes post inclusion of Sri Lankan taxa. (Bayesian posterior probability = 0.98 and ML bootstrap support = 98; Figs 1 and S1). Sri Lankan Rhomphaea included in this study do not form a single clade, instead popup at various points within the Rhomphaea clade.

Divergent time estimation
Results from our dating analysis are presented in Fig 3. The dated phylogeny suggests that origin of Argyrodinae occurred around 19.6 Mya (96% Highest Probability Density, HPD interval of 22.5-16.75 Mya) which was between the late Paleogene and early Neogene (Fig 3). The origin of kleptoparasitism was dated in between the early to late Neogene (mean = 14.09 Mya with 95% HPD interval of 15.75-12.75 Mya). Based on our dating analysis, araneophagy may have evolved at least 5 My earlier than kleptoparasitism. Further, the origin and diversification of the genus Rhomphaea in Sri Lanka was in the early Neogene at around 10-5 million years ago.

Phylogenetic analysis
Phylogenetic relationships inferred in this study largely confirm previous findings of Su & Smith [18]. However, the phylogenetic placement of Rhomphaea and Neospintharus yielded mixed results. After the addition of 12 new terminals, comprising of sequences for five species to the matrix of Su & Smith [18], the resulting trees for both analyses recover a fairly well supported Rhomphaea clade. All putative Sri Lankan species cluster within this clade (Fig 1).
Our results provides further evidence in the classification of argyrodines by recognizing four distinctive generic clades of Argyrodes, Ariamnes, Rhomphaea and Neospintharus. However, our results indicate that circumscription of genera within Argyrodinae needs further attention. This is also suggested by the polytomies present in our preferred phylogenetic hypothesis. We found that, topology and clade composition of our phylogeny differs somewhat from previous studies [9,18]. However, we have here included only a limited sample of the diversity of Argyrodinae, focusing on taxa closely related to the stick spiders of Sri Lanka.
Members of genera Ariamnes and Rhomphaea are solitary, free living and specialized in 'silk throwing' [37,38] behavior associated with araneophagy and kleptoparasitic Argyrodes species [18,37,39]. Neospintharus shows both traits; directly feeding on prey and occasionally 'silk throwing' [18,37]. Unfortunately, behavioral data for Sri Lanka species included here are still not available. This study as well as unpublished data on the kleptoparatic Argyrodes, represents the most comprehensive molecular survey conducted for the subfamily Argyrodinae of Sri Lanka. Our molecular genetic data will enable further evolutionary studies on the diversification of the subfamily. The discovered molecular and morphological diversity, new species and their restricted range distributions with species inhabiting low land and highland cloud forest call for a larger assessment of their evolutionary history and conservation, based on broader sampling regimes and genomic analyses. Further, future studies will also have to address the systematics of the Argyrodinae as no clear evidence supporting the monophyly of generic groups or species groups were found.

Divergent time estimation
Based on our calibration (Fig 3) Sri Lankan Rhomphaea and Neospintharus species appear to have diverged very recently (less than 2Mya). Further, although our results are preliminary, it The tree recovered from the ML analysis of the concatenated molecular data is used here as it is better resolved. Key: "Navajo rugs" indicate posterior probability support of a node is larger than 0.7 in Bayesian analysis and larger than 70 in ML analysis of the individual gene or in concatenated data (black), or nodal supports values between 0.6-0.7 in Bayesian and 60-70 in ML analysis (red), absence of a given node in the tree specified in the legend (white). Diagnosis. Rhomphaea species differ from other Argyrodinae by the following characters: prosoma of males with elevated or projected eye region or sometimes with a lobe-like structure; clypeus slanted anteriorly; opisthosoma boomerang shaped [8], elongated, triangular or cylindrical, posterior half of opisthosoma (extending beyond the spinnerets) at least three times longer than the anterior half, sometimes tapered to form a single tip with a seta, in most of the species orientation of anterior and posterior opisthosoma halves form angle at spinnerets; elongated palpal tibia, embolus spiral directed clockwise in left palp, membranous conductor, in some species apex of TTA enlarged; epigynum with a socket-like ventral membranous structure (scape); rhomboid shaped egg sac [8,9].  [20]. Placement of oldest date marked as purple asterisk on the node. The yellow bar on the node shows the 95% highest probability density interval of the estimated time. https://doi.org/10.1371/journal.pone.0273105.g003

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Free-living stick spiders of Sri Lanka Description. Medium to large spiders. Prosoma pale yellow, with lateral black bands arising from the back of eye band. Eye band projected anteriorly, fused laterals. Clypeus slanted, projected anteriorly, distinct bi-lateral blackish-brown bands. Sternum slightly convex, blackish brown dusty flecks. Labium fused with sternum. Opisthosoma similar to prosoma in coloration with silver and brown-black flecks, triangular or cylindrical shaped, anterior and posterior opisthosoma divided by spinnerets, posterior half 3-5 times longer than anterior. Legs paleyellow, reddish brown bands/patches, leg formula I, IV, II and III. Pedipalp and legs similar in color and banding pattern. Palpal tibia elongated, narrow, tapered towards patella. Tibial rim scoop-shaped or flat with 3-5 retro lateral trichobothria. Cymbium entire with a characteristic cymbial hook or hood. Paracymbium absent. Membranous conductor closely associated with embolus tip. Embolusbifurcated in some species (embolus apophysis [9]), base lobbed, embolus spiral tip elongated and needle-like. TTA membranous or slightly sclerotized. Female epigynum not well sclerotized, ventral plate with a socket-like membranous hood (scape) [40], inconspicuous copulatory openings inside it. A pair of spermathecae oval shaped and larger. Copulatory duct long and encircled or short and straight. Fertilization duct shorter and twisted or straight.
Composition. 33 species; see World Spider Catalog [11] for a listing of all species. Distribution. Worldwide; See World Spider Catalog [11]. Etymology. This species is named after the first author's mother, Shanthinimala Tharmarajan. Used as a noun in apposition.

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Free-living stick spiders of Sri Lanka heavily ridged, not well sclerotized when compare to E and MA, located between E and MA, originates from tegular margin. MA unbranched, flexibly attached sclerite, mesial originated on T, outgrowth of T, small loop of SD can be seen, accommodates Chk corresponding pit. TTA membranous, vertical tube-like, unbranched, apex with 3-5 finger-like projections, resides back of E, attached to T, loosely associated. E rests in front of bulb, originates retroventrally, well sclerotized, longer than half of cymbium length, "inverted question mark" shaped, lobed base, spiral narrow cylindrical, tip hook-like. Palp measurements: length of Ti 0.5, length of Cy 0.3.
Diagnosis. Males of R. jacko sp. nov. closely resembles R. sagana but differ from other congeners by the length (longer) and shape of opisthosomsa (sickle-shaped, narrow, tapered and posterior part approximately about 8-10 times longer than anterior). The palpal sclerites (shape and location of embolus, conductor). Conductor (clearly visible, distal portion with hook-like apophysis (sub conductor). In R. sagana, conductor is not clearly visible [40]. Further, the lobed embolus base, embolus spiral narrow and grooved, the needle-like embolus tip is diagnostic.
Description. Male (holotype): based on alcohol preserved specimens. Prosoma yellow, flat but slightly convex in the middle with projected eye region, midline of dorsal prosoma and sternum with bilateral narrow faint-black lines, prosoma width equal to opisthosoma width, prosoma length 1.35, prosoma width 0.9. Clypeus slanted, length 0.26. Sternum narrow, 3 times longer than wide, widest at the 2 nd coxae and pointed between 4 th coxae. Labium semicircular and re-bordered. Eyes positioned on upper projection of prosoma, wider eye band, larger medians, measurements: AME 0.1, ALE 0.08, PME 0.08, PLE 0.08, PME-PME 0.1, PLE-PLE 0.3, ALE-PME 0.08, ALE-PLE 0. Opisthosoma sickle shaped, triangular, very long, posterior opisthosoma almost 10 times longer than anterior part, posterior opisthosoma covered with shiny larger silver spots, cylindrical, elongated upwardly and tapered towards the posterior tip with a characteristic long strong seta, length; pedicel-spinnerets 0. Tibia scoop shaped, distal end slightly wider than base, rim strongly asymmetric, protruding, strongly exaggerated on one side, characteristic row of long strong setae, 4-5 retro lateral trichobothria, two prolateral trichobothria. Cymbium entire, retromargin with a small distal cymbial hood. Chk faces downward inside Cy, distal portion blunt. T less than half of cymbial cavity. C located between E and MA, slightly sclerotized, associated C and MA, distal portion branched with hook-like apophysis (sub conductor), folded to hold embolus tip. MA distal pit accommodates BC lock mechanism, margin well sclerotized, unbranched, placed on tegulum, loop of sperm duct not clearly visible. TTA membranous, not sclerotized, tubelike elongated, retro laterally originated, very closely associated with C and E, apex with a knob in which embolus tip rests. E retro ventrally originated on tegulum, not hidden by cymbium, slightly curved, spiral tip entire not forked and needle-like with a median groove.
Female: unknown. Distribution. See Fig 11. Rhomphaea martini sp. nov. Etymology. The species is named after Sri Lankan novelists Lama Hewage Don Martin Wickramasinghe. He is known for his trilogy Gamperaliya, Yuganthaya, and Kaliyugaya. He also lived on the same street in Colombo as the second author.
Diagnosis. Males of this species differ from others congeners by the ocular area of male (projected anteriorly with a characteristic seta). Median eyes distantly separated from each other. Prosoma bordered laterally with two black dusty stripes continued from clypeus, stripes much darker and wider than those of R. jacko sp. nov. Opisthosoma of males shorter, triangular, rounded tip. Tibia of male palp much shorter than others. Embolus bifurcated with an apophysis. Females differ from others congeners by the conical, long, triangular opisthosoma with silvery patches and pointed tip, when compared to R. shanthi sp. nov. ventral epigynum socket (scape) smaller. CD with two loops, enters S anterioly (Fig 7C).
Distribution. See Fig 11. Rhomphaea marani sp. nov. Etymology. This species is named after the author's brother Seermaran for assistance and support during this study.
Diagnosis. This species differs from others congeners by the following characters: prosoma shaded with brown with a median yellow stripe. Thoracic region with deep incision. Eye region not projected as R. martini sp. nov. furrowed between median eyes. Narrow cylindrical opisthosoma with dark reddish-brown patches all over the body, which gives them a unique twig-like appearance. Male palp with elongated tibia and bifid embolus. Epigynum with large S, short CD. CD longer in both R. shanthi sp. nov and R. martini sp. nov.
Description. Male (holotype): based on alcohol preserved specimens. Prosoma pale yellow with two broad black bands on both sides extended up to clypeus, deep incision in the middle, two humps, prosoma length 1.05, prosoma width 0.75. Clypeus length 0.27. Sternum pale yellow with similar pattern of patches, wider than long, widest at 1 st coxae and pointed at 4 th coxae. Eye region projected anteriorly with usual eye band arrangement, larger AMEs, medians separated from each other and close to laterals, measurements: AME 0.1, ALE 0.1, PME 0.1, PLE 0.1, PME-PME 0.1, PLE-PLE 0.3, ALE-PME 0.1, ALE-PLE 0. Opisthosoma narrow cylindrical, pale yellow with prominent silver and brownish black patches, similar pattern of patches on ventral side, same width throughout the whole length, posterior region almost two times longer than anterior, posterior end shaded with dark brown and with 3-4 characteristic small humps, cross section of opisthosoma end triangular, length Pedicel-spinnerets 0.8, spinnerets-tip of opisthosoma 1.9, opisthosoma width 0.66. Spinnerets located in 1/3 rd length of opisthosoma anteriorly and positioned angularly. Legs; pale yellow, with black bands. Palp similar to legs in coloration and pattern of bands, measurements: length of palp tibia 0.7, length of cymbium 0.4. Tibia distal end slightly wider than base and tapered, base narrower than patella at its articulation, tibia rim exaggerated in the middle and facing bulb, scoop shaped, rim with regularly arranged row of uniform strong long setae, 4-5 retro lateral trichobothria, 1-2 pro lateral trichobothria. Cymbiumentire without cymbial hood. Chk distally placed inside of the cymbium, facing downward, distal portion tapered to a sharp tip. MA with distal narrow pit where Chk locks. ST large. T not larger than half of the cymbial cavity, with tegular pit at its ectal margin connected to embolus base. Conductor extension of tegulum, located between embolus and MA, membranous, larger and broader, distal portion bent but not folded. MA mesial originated, outgrowth of tegulum, unbranched, loop of sperm ducts not clearly visible, narrow pit-like hood on the distal MA. TTA membranous, long tube-like, apical knob of TTA overlaid on embolus spiral tip, closely associated with C and E, attached to T via extra tegular apophysis (ETA). ETAlocated in the ectal margin of tegulum and closely related to TTA. Bifid embolus with embolus apophysis, embolus base lobed and placed retro ventrally. Embolus spiral narrow, cylindrical and tapered distally to form a needle-like sharp tip.
Distribution. See Fig 11. Genus Neospintharus Exline, 1950. Neospintharus Exline, 1950. Type species: Argyrodes parvus Exline & Levi, 1962. Diagnosis: Prosoma of male with anteriorly projected caput process and clypeus separated by a groove, two horn-like projections finished with long setae. Anterior median eyes in males placed on both sides of groove between eye projection and clypeal processes. Labium fused with sternum and not re-bordered. Opisthosoma triangular, posterior opisthosoma tip bifurcated, accommodates humps or forked (fish tail-like) process. Conductor distal tip is entire and lobed. TTA with enlarged and ridged distal tip. Embolus short or medium size.
Description. Male (holotype): based on alcohol preserved specimens. Prosoma yellowbrown with some grey hue, wider, prosoma length 1.06, prosoma width 0.72. Caput process projected anteriorly to form a small high lobe with blunt tip of the process bearing 3-4 long and strong setae. Clypeal process slightly longer than head process, narrow, cylindrical, tapered towards tip bearing 3-4 long and strong setae as in caput process, projected parallelly to caput process, not equal length of caput process. Sternum, coxa pale yellow with some grey hue except coxa legs yellow-brown in colour. Sternum shield-like. Labium fused with sternum and not re-bordered. Eyes located between caput process and clypeal process, AMEs larger

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Free-living stick spiders of Sri Lanka than PMEs, square orientation of AMEs and PMEs. Separation of posterior medians slightly more than the separation of anterior medians. Eye measurements: AME 0.1, ALE 0.06, PME 0.1, PLE 0.06, PME-PME 0.1, PLE-PLE 0.36, ALE-PME 0.06, and ALE-PLE 0 (fused laterals). Opisthosoma nearly triangular, light grey with silvery hue or spots, dorsal opisthosoma more concentrated with a band of silver hue, posterior opisthosoma blunt darker grey hue and characteristically both lateral sides with a larger circular marking filled with darkened grey flecks. Spinnerets; light grey, placed in the middle and lower edge of opisthosoma, total length 1.98, opisthosoma length 1.08 and opisthosoma width 0.74. Leg measurements: leg I: 4.48 Palp distinctively broadened, tibial distal end asymmetrically slightly protruding, tibial rim facing bulb of the palp. Tibia with two retrolateral trichobothria and one prolateral trichobothria. Cymbium entire retro margin, distal pro margin, no cymbial hood. Chk blunt tip placed distally and facing downward. T size less than half of cymbial cavity and located on top of ST. C originated from T, entire not folded or grooved and distal tip of embolus and conductor associated with each other, C distal portion not pointed, subequal to base, sclerotized. MA mesially originated, easily distinguishable sclerite, placed on tegulum. TTA unbranched, located between MA and C, closely associated with E and C, distal tip enlarged and strongly sclerotized ridged surface. E originated retro ventrally on tegulum, embolus spiral short, embolus base slightly lobbed, measurements: palp tibia 0.2 and length of cymbium 0.32. Female (paratype). based on alcohol preserved specimens. All characters as in male, except the following: larger than males, total length 2.12, pattern similar to males, darker and browner than males with silvery spots. Prosoma; middle dark patches, posterior wider, length 1.04, width 0.74. Clypeus without modifications, high, rounded. Eye region; slightly elevated with a groove under AMEs, measurements: AME 0.1, ALE 0.1, PME 0.1, PLE 0.1, PME-PME 0.