Phylogenetic placement of Sri Lankan goblin spiders (Araneae, Oonopidae): integrating new taxa and taxonomic implications

Phylogenetic placement of Sri Lankan goblin spiders (Araneae, Oonopidae): integrating new taxa and taxonomic implications U. G. S. L. Ranasinghe1, Suresh. P. Benjamin1 1 National Institute of Fundamental Studies, Hantana Road, Kandy, Sri Lanka https://zoobank.org/EC62195B-BD4F-43CF-B80F-681934F28260 Corresponding author: Suresh. P. Benjamin ([email protected]) Academic editor: Stephanie F. Loria ♦ Received 12 August 2025 ♦ Accepted 3 November 2025 ♦ Published 14 November 2025 Abstract Sri Lanka harbours an exceptionally high diversity of goblin spiders (family Oonopidae), with 45 currently recognised species across 13 genera, of which 38 are endemic. In this study, we present a molecular phylogenetic analysis based on two nuclear ribosomal markers (18S and 28S rRNA), incorporating 28 species from eight genera: Aprusia, Brignolia, Ischnothyreus, Opopaea, Orchestina, Pelicinus, Silhouettella, and Xestaspis. These taxa were analysed in the context of the existing global phylogeny, enabling both regional and broader evolutionary comparisons. Our results reveal that several genera, particularly Brignolia, Opopaea, and Ischnothyreus, are paraphyletic, highlighting the need for taxonomic revision. Endemic clades such as Aprusia, Brignolia, and Xestaspis show strong evidence of recent, within-island speciation and microendemism. The genus Orchestina is recovered as monophyletic, with Sri Lankan species forming a distinct clade. The findings support the hypothesis that reduced body sclerotisation is a plesiomorphic trait in oonopids. Overall, the study underscores the exceptional microendemism and evolutionary distinctiveness of Sri Lanka’s oonopid fauna, likely driven by both ecological isolation and in situ speciation. However, given the incomplete representation of Indian and Southeast Asian taxa, and limited molecular sampling, further studies incorporating broader taxon and gene sampling are needed to fully resolve the origins and diversification of this ecologically important spider family. Key Words Araneae, Biogeography, Ceylon, Phylogenetics, short-range endemics Introduction Short-range endemic species are important for understanding both local biodiversity and broader biogeographic patterns (Harvey et al. 2011). Compared to more widespread and less habitat-specific invertebrates, they offer greater potential to reveal phylogeographic patterns and the influence of historical and present-day landscape processes (Harvey et al. 2011). They might be more susceptible to extinction (Mammola et al. 2018) due to climate change (Harvey and Dong 2023), agroforestry, urbanisation and pollution (Branco and Cardoso 2020). Generally, goblin spiders (Araneae: Oonopidae), which are considered shortrange endemics with very restricted distributions, and may prove to be important indicator taxon for monitoring the effects of climate change and other threats on forest habitats (Baehr 2011). Goblin spiders, currently include over 1978 described species in 115 genera (World Spider Catalog 2025). Members of this family are small (1–4mm), ecribellate, haplogyne (Jocqué and Dippenaar-Schoeman 2006), free hunting spiders that live in leaf litter, canopy, under stones, rocks, tree barks or even within buildings (Fannes et al. 2008; Grismado et al. 2011; Thoma et al. 2014). Goblin spiders are usually yellowish, orange or reddish-brown (Fig. 1), although some species can be whitish or pink (Saaristo 2001; Eichenberger et al. 2012). Although small in size, they exhibit a remarkable diversity of morphological characters (Platnick and Brescovit 1995; Fannes and Jocqué 2008; Platnick and Dupérré 2009a, b; Baehr and Ubick 2010; Baehr et al. 2010; Platnick and DupérEvolutionary Systematics. 9 2025, 259–270 | DOI 10.3897/evolsyst.9.168533 Copyright U. G. S. L. Ranasinghe & Suresh. P. Benjamin. 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. evolsyst.pensoft.net U. G. S. L. Ranasinghe & Suresh. P. Benjamin: Sri Lankan goblin spiders 260 ré 2010a, b; Ubick and Griswold 2011; Brescovit et al. 2012a, b; Platnick and Berniker 2014; Platnick and Berniker 2015; Tong and Li 2015). To date, thirteen genera of goblin spiders have been reported in Sri Lanka: Aprusia Simon, 1893 (Grismado et al. 2011; Ranasinghe and Benjamin 2018a), Brignolia Dumitrescu & Georgescu, 1983 (Platnick et al. 2011; Ranasinghe and Benjamin 2016a), Camptoscaphiella Caporiacco, 1934 (Baehr and Ubick 2010), Cavisternum Baehr, Harvey & Smith, 2010 (Ranasinghe and Benjamin 2018b), Gamasomorpha Karsch, 1881 (Eichenberger et al. 2012), Grymeus Harvey, 1987 (Ranasinghe and Benjamin 2018b), Ischnothyreus Simon, 1893, Opopaea Simon, 1892 (Platnick and Dupérré 2009), Orchestina Simon, 1882 (Ranasinghe and Benjamin 2025), Pelicinus Simon, 1892 (Ranasinghe and Benjamin 2018b), Silhouettella Benoit, 1979 (Ranasinghe and Benjamin 2018b), Xestaspis Simon, 1885 (Eichenberger et al. 2012; Ranasinghe and Benjamin 2016b) and Xyphinus Simon, 1893 (Ranasinghe and Benjamin 2016c), which includes 45 Oonopidae species, highlighting the diversity of the family on the island. Large continental islands such as Sri Lanka, where faunal assemblages are shaped by both recent colonisation from the mainland and longterm in situ diversification (Mayr 1947) remain relatively understudied in terms of their evolutionary dynamics (Beenaerts et al. 2010; Benjamin 2010; Pyron et al. 2013; Karunarathna et al. 2019; Meegaskumbura et al. 2019; Sudasinghe et al. 2020; Perera and Suranjanfernando 2024). Within this context, the family Oonopidae (goblin spiders) presents a compelling study group. The first molecular phylogeny of Oonopidae by Busschere et al. (2014), based on ribosomal RNA sequences from 37 genera, laid the groundwork for understanding the global diversity of this family. Field surveys across more than 100 localities have revealed that many of these species exhibit highly restricted distributions, some occurring at only a few sites, while others are known from a single forest patch and are absent even in the immediate surrounding forests (Ranasinghe and Benjamin 2016a, b, c, 2018a, b). These patterns highlight the significance of exploring the phylogenetic relationships of Sri Lanka’s short-range endemic oonopids. Incorporating these taxa into a broader molecular framework will not only enhance the global phylogeny of the group but also provide critical insights into the evolutionary mechanisms driving microendemism on continental islands. In this study, we present the first phylogeny of Sri Lankan goblin spiders and investigate the relationships of these short-range endemic species in the island to better understand how they contribute to the global diversity of the group (Busschere et al. 2014). Figure 1. A–D. Live goblin spiders, genus: Xestaspis sp. photo credit: SPB. Evolutionary Systematics 9 2025, 259–270 evolsyst.pensoft.net 261 Methodology Taxon Sampling Specimens were collected from the natural forest of major climatic-physiographic zones of Sri Lanka (Pethiyagoda and Manamendra-Arachchi 1998), i.e. Central Mountain Range, Knuckles Range, Low Country wet zone and the dry zone, with the aim of maximizing the number of species collected (Fig. 2). Locations are listed alphabetically by provinces and districts in Appendix 1 (extended version of Batuwita and Benjamin 2014). Specimens were collected by sifting litter and leaving the residue overnight in a Winkler extractor or by hand sorting the residue. The collected specimens were preserved in 100% ethanol. The preserved specimens were examined using an Olympus SZX 7 stereomicroscope and identified up to generic level using recently published international revisionary works of Oonopidae (Platnick and Dupérré 2009; Baehr and Ubick 2010; Harvey and Smith 2010; Grismado et al. 2011; Platnick et al. 2011; Eichenberger et al. 2012; Henrard and Jocqué 2012; Platnick et al. 2012a, b). DNA Extraction, amplification, purification and sequencing Genomic DNA was extracted from 100% ethanol-preserved leg tissue using the DNeasy Blood and Tissue Kit (Qiagen, Hilden, Germany) following the manufacture’s protocol. Initially, Polymerase chain reactions (PCR) were carried out using 10 primer pairs (Suppl. material 1: table S1) for 43 DNA samples (Suppl. material 1: table S2). However, the amplification of 16S, CO1 and H3 fragments was only partially successful. Thus, four promising primer pairs (18Sa2.0/9R (650 bp), 18S3F/18Sbi (850 bp) 18S1F/18S5R (820 bp) and 28SZX/28SC (1200 bp) for three overlapping regions of 18S and partial fragment of 28S were selected for further PCRs. PCR was performed with an initial denaturation at 95 °C for 2'30", then 35 cycles started with denaturation at 95 °C (30"), annealing step of 30" 18Sa2.0/9R: 48 °C, 18S3F/18Sbi and 18S1F/18S5R: 56 °C, 28SZX/28SC: 48.5 °C and extension at 72 °C for 1'30" for 35 cycles. The PCR reaction mixture (20 µl) included Gotaq DNA Polymerase, 5× colorless Gotaq Flexi Buffer, Magnesium Chloride 25 mM, dNTP Mix (Promega Corporation, Madison, USA) and undiluted DNA template. PCR for older specimens was carried out using the Qiagen Multiplex PCR’ kit (Qiagen, Hilden, Germany) in total reaction mixes of 20 µL, including 2 µL of undiluted DNA template, 1.6 µL of each primer (10 pM /mL), 2 µL of ‘Q Solution’ and 10 µL of the ‘Multiplex PCR Master Mix’, containing hot start TaqDNA polymerase and buffers. However, contrary to the manufacture’s protocol the PCR reactions were not multiplexed, but run individually (Dimitrov et al. 2012; Huber et al. 2014). Thermal cycling was performed on a MyCycler (Biorad, California, USA). PCR products were visualized by 1% agarose gel in 1× TBE buffer. Bands were visually characterized as either weak or strong based solely on brightness. Strong bands were then purified using the QIAquick PCR Purification Kit (Qiagen). The weak bands were purified using the Illustra ExoProStar (GE Healthcare, UK) PCR clean-up system, according to manufacturer’s protocol. All purified PCR products were Sanger sequenced in both direction by MACROGEN (Seoul, South Korea). Figure 2. Sampling localities around Sri Lanka; L44: Knuckles forest reserve, L10: Ethagala forest, L20: Hiyare; L29: Meemure. evolsyst.pensoft.net U. G. S. L. Ranasinghe & Suresh. P. Benjamin: Sri Lankan goblin spiders 262 Sequence editing, alignment and phylogenetic tree construction Sequences were edited and assembled using Geneious 6.1.5 software package (Kearse et al. 2012). Sequence alignment was done by Geneious 6.1.5 software using default parameters and then further refined and edited manually if needed using Mesquite (version 3.04; Maddison and Maddison 2011). Three data matrixes were assembled for analysis, 18S (1796 bp), 28S (1123 bp) and a concatenated 18S+28S alignment. The concatenated 18S + 28S matrix was 2782 bp in length. Maximum likelihood (ML; Felsenstein 1973) searches were performed in IQ-TREE version 1.6.12 (Nguyen et al. 2015) under the (GTR+F+I+G4) model of nucleotide substitution that was inferred as the best-fit model by ModelFinder (Kalyaanamoorthy et al. 2017). A total of 1000 ultrafast bootstrap (Hoang et al. 2018) replicates were done to assess branch supports. Additional sequences of the Oonopidae 18S region (n=111) and 28S region (n=57) were obtained from GenBank (ncbi.nlm.nih.gov), and the whole dataset was assembled and aligned using the Mesquite software package (version 2.72; Maddison and Maddison 2011). Final analyses included 154 taxa representing 40 genera with a worldwide distribution (142 ingroup and 12 outgroup). The ingroup consisted of Oonopidae taxa and the outgroups were representatives of Dysderidae, Orsolobidae and Segestriidae. Representative Filistatidae and Liphistiidae taxa were used to root the tree (Busschere et al. 2014). Results We present a phylogenetic analysis based on two nuclear ribosomal loci (18S and 28S rRNA), resolving the relationships among 43 goblin spider taxa from Sri Lanka. The dataset includes representatives from the genera Aprusia, Brignolia, Camptoscaphiella, Ischnothyreus, Opopaea, Orchestina, Pelicinus, Silhouettella, and Xestaspis. These taxa were integrated into a global phylogenetic framework (now consisting of 154 taxa representing 40 genera) with a worldwide distribution (142 ingroup and 12 outgroup taxa). This includes representatives of four families in the superfamily Dysderoidea: Dysderidae, Oonopidae, Orsolobidae, and Segestriidae, as well as the three subfamilies of Oonopidae: Oonopinae, Orchestininae, and Sulsulinae (as in Busschere et al. 2014). Results from molecular analyses based on the 18S, 28S, and combined 18S+28S datasets suggest that the family Oonopidae is monophyletic, supporting the findings of previous studies (Busschere et al. 2014). The genus level relationships within Oonopidae are well resolved in our analysis and largely align with morphologically defined groups, with a few exceptions. In contrast to previous works (Busschere et al. 2014), the genera Aprusia, Camptoscaphiella and Xestaspis are included here for the first time, while multiple species now represent Brignolia and Pelicinus. Moreover, monophyletic clades were recovered for the genera Xestaspis, Escaphiella, Orchestina, Pelicinus, and Neotrops. In contrast, Brignolia, Opopaea, Aschnaoonops, Paradysderina, Unicorn, and Silhouettella were not resolved as monophyletic in any of the analyses. In the updated global phylogeny, the Sri Lankan taxa exist in five distinct clades (Fig. 3), reflecting both multiple colonization and localized radiations within the island. 1. Brignolia and Opopaea clade The genus Brignolia is well represented in all analyses including sixteen species for the combined analysis (Fig. 3). A well-supported Brignolia + Opopaea clade was recovered. However, neither genera are monophyletic. Brignolia ondaatjei is sister to Sri Lankan Opopaea spinosiscorona (Fig. 3). This group is well supported. Further, B. ambigua, B. carlmulleri and B. shyami formed a separate group. B. ambigua appeared as a sister to the other two species. This group is sister to B. meemure and B. sinharaja. B. ratnapura together with B. parumpunctata formed the next group. In the 18S+28S concatenated analysis, all Brignolia species together with Opopaea species are sister to O. apicalis from the Galapagos, Ecuador (Fig. 3). However, Opopaea from Sri Lanka form a group separate from Australian, South American and African Opopaea. Thus, this study suggests that both genera need to be redefined in morphological terms. 2. Xestaspis clade The character combination that defines Xestaspis is the reddish brown, well-sclerotised scuta, spineless legs, sternum with radial furrows and the presence of a palpal bulb that is not fused with the cymbium (Eichenberger et al. 2012). Xestaspis is included for the first time in a phylogenetic analysis of goblin spiders. It forms a well-supported clade (Fig. 3). Four species were included Xestaspis kandy, X. nuwaraeliya, X. padaviya and X. pophami. Here, X. padaviya is sister to all other Xestaspis species. Xestaspis is sister to Prethopalpus tropicus of Australia (Fig. 3). 3. Silhouettella and Pelicinus clade The genera Silhouettella together with Pelicinus form sister groups in the combined analysis (Fig. 3). Both genera have quite similar secondary genital organs. However, Pelicinus differs by having conspicuously longer and slender spineless legs and large, oval-shaped operculae (Saaristo 2001). This clade consists of P. marmoratus, P. snooky and P. tumpy. They are sister to Silhouettella snippy and S. saaristoi. Evolutionary Systematics 9 2025, 259–270 evolsyst.pensoft.net 263 Figure 3. Phylogenetic relationships of Oonopidae and related families based on concatenated 18S+28S sequence data (2782 bp), including 154 taxa (142 ingroup), species of Orsolobidae, Dysderidae, Segestriidae included as outgroups. Numbers on branches indicate bootstrap proportions > 40% from 1000 replicates. Sri Lankan endemics are indicated with an asterisk (*). evolsyst.pensoft.net U. G. S. L. Ranasinghe & Suresh. P. Benjamin: Sri Lankan goblin spiders 264 4. Aprusia, Camptoscaphiella and Ischnothyreus clade The South Asian goblin spider genus Aprusia is characterised by a unique combination of characters, such as the relatively small to medium-sized dorsal scutum, the presence of strong spines on leg I and II, the anteriorly directed receptaculum and the male palpal bulb fused to cymbium and a small, slightly sclerotised embolus (Grismado et al. 2011). According to the combined analysis Aprusia and Ischnothyreus together with Camptoscaphiella form a well-supported clade (Fig. 3). However, both Aprusia and Ischnothyreus are paraphyletic (only a single species of Camptoscaphiella is included in the current study). Aprusia sp. 2 is sister to Ischnothyreus peltifer from the Galapagos, albeit with low support. Further, Aprusia vestigator (Simon 1893) is sister to Aprusia sp. 1 (IFS_Oon_189). This clade is recovered as sister to Sri Lankan Camptoscaphiella simoni. 5. Orchestina clade The soft-bodied goblin spider genus Orchestina includes canopy-dwelling jumpers that are easily identified by the elongated and enlarged fourth femora, H-shaped eye arrangement and enlarged male palpal tibia (Saaristo 2001; Henrard and Jocqué 2012). Three specimens form a well-separated branch. In the global analyses, it is represented by 19 specimens from different continents (Africa, Asia and South America). The three Sri Lankan Orchestina form a well-supported clade which is sister to the Congolese Orchestina saaristoi in all three analyses (Fig. 3, Suppl. materials 2, 3, fig. 2 in Ranasinghe and Benjamin 2025). Tree-based species delimitation (Poisson tree processes model (Zhang et al. 2013)) for only Sri Lankan species for their individual 18S, 28S, and combined 18S+28S trees showed above five distinct clustering pattern (shown as boxes in Fig. 4). Figure 4. Tree-based Poisson tree processes model of species delimitation for Sri Lankan oonopids for their individual 18S, 28S, and combined 18S+28S trees. Boxes indicate agreement between molecular species delimitation method and species assignment. Evolutionary Systematics 9 2025, 259–270 evolsyst.pensoft.net 265 Discussion The selection of genes for this study was guided by two main considerations. First, although we attempted to amplify fragments of 18S, 28S, 16S, CO1 and H3 for the sampled species, the most consistent and successful results were obtained for the ribosomal RNA genes 18S and 28S. Second, we prioritised compatibility with the existing global phylogeny of Busschere et al. (2014), necessitating the use of these same markers. However, the 18S and 28S genes evolve relatively slowly and are highly conserved across species, making them well-suited for examining deep evolutionary relationships (Koh et al. 2023; Xing et al. 2025; Fig. 4). Although strong support for the monophyly of Brignolia + Opopaea has been previously reported (Busschere et al. 2014), both were recovered here as paraphyletic. This result contrasts with the current morphological hypothesis that both genera are monophyletic. The presence of the dorsal depression (‘‘fenestra’’) on the palp was previously considered synapomorphic for both genera (Platnick and Dupérré 2009a). However, Opopaea differ from Brignolia by the presence of a distinct enlarged male palpal patella. Females of Brignolia and Opopaea are very similar in having a dorsal abdominal scutum that covers most of the abdomen. However, female Brignolia have a simple or twisted posterior genitalic tube that is absent in females of Opopaea (Platnick et al. 2011; Platnick et al. 2012a). In this study, B. ratnapura and B. parumpunctata formed a well-supported group. Platnick et al. (2011) stated that B. ratnapura might be the sister species of the pantropical B. parumpunctata, based on their genitalic structures. The two species share a rounded dorsal protrusion on the male palpal bulb (figs 12, 280 in Platnick et al. 2011) and a highly twisted posterior tube within the female genitalia (figs 71, 294 in Platnick et al. 2011). B. carlmulleri and B. shyami is recovered as sister species in the combined analysis. The presence of a triangular projection on the base of the endite (figs 4c, 17a, b in Ranasinghe and Benjamin 2016a), dorsally directed palpal tip and the rounded palpal tibia in males (figs 6a, 18a in Ranasinghe and Benjamin 2016a) are characters shared in both species. Further, males of Opopaea, Prethopalpus, Camptoscaphiella and Malagiella have a greatly enlarged palpal patella that originates sub-basally or medially from the femur (Platnick and Dupérré 2009; Baehr 2011; Baehr et al. 2013). The later genus was not included in this analysis. Aprusia and Ischnothyreus together with Camptoscaphiella form a well-supported clade in the 18S and the 18S+ 28S phylogeny (Fig. 3, Suppl. materials 2, 3). However, Aprusia and Ischnothyreus are both paraphyletic and needs further taxonomic attention. Aprusia resembles Ischnothyreus and Camptoscaphiella in eye arrangement, the shape of the abdominal scutum and the strong spination on legs I and II (Grismado et al. 2011). However, they differ in their copulatory organs. The copulatory orifice opens on the postepigastric plate in Camptoscaphiella and Ischnothyreus, while the opening is inconspicuous and located presumably in the epigastric furrow in Aprusia (Baehr and Ubick 2010; Grismado et al. 2011; Platnick et al. 2012a). All analyses recovered a well-supported monophyletic Orchestina (same as in Busschere et al. 2014). Sri Lankan Orchestina species form a well-supported clade, sister to O. saaristoi from the Congo, whereas two specimens from Singapore form a different clade with O. macrofoliata Henrard & Jocqué, 2012 and O. fractipes Henrard & Jocqué, 2012 from the Congo. Sri Lankan Orchestina specimens are morphologically similar to O. saaristoi (figs 573–605, 607 in Henrard and Jocqué 2012; Ranasinghe and Benjamin 2025). The genus Silhouettella is paraphyletic in the 18S as well as the 18S+28S combined phylogeny (Fig. 3, Suppl. material 2). In the combined analysis, Silhouettella snippy and S. saaristoi from Sri Lanka group together, while Silhouettella sp. from Madagascar is placed as sister to Farqua sp. from South Africa. However, S. tiggy from Sri Lanka is placed as sister to Silhouettella loricatula from France. According to the 18S and 18S+28S phylogeny, Pelicinus formed a well-supported clade (Fig. 3, Suppl. material 2). Two species of Pelicinus from Sri Lanka (Pelicinus marmoratus and P. snooky) group with Pelicinus sp. 65 from Australia. Previous morphological studies group Pelicinus, Silhouettella, Lionneta and Grymeus together and consider them members of the Pelicinus group (Burger 2011; Platnick et al. 2012b). Shared features include the “squiggled” shape of the globular appendix, the pore plate in front of the posterior receptaculum and the details of the anterior “paddle like sclerite” in the female genitalia (Burger 2011; Platnick et al. 2012b). The Dysderina-group is a large morphologically defined group within Oonopidae, and several of these species form numerous well-supported groups within the Dysderina clade (Platnick and Dupérré 2011a, b). These phylogenetic relationships are consistent with those of Busschere et al. (2014). The addition of new data does not seem to have any effect (but see Graybeal 1998; Zwickl and Hillis 2002; Heath et al. 2008). However, as in previous studies, even this small, geographically restricted sample is not recovered as monophyletic. According to the results of the combined analysis, the hypothesis that Oonopidae is monophyletic is supported. Further, many oonopid genera constitute natural groups. Further, most soft-bodied genera are basal within the Oonopidae. Busschere et al. (2014) suggested that a low degree of body sclerotisation is a plesiomorphic trait, with a higher degree of body sclerotization being considered a derived trait among Oonopidae. This hypothesis is supported by the present results. In addition, these results confirm the hypothesis that Orchestininae and Sulsulinae are basal oonopids. However, inclusion of absent genera in future studies will help obtain better resolution and form a definite conclusion. Considering the distribution of 45 Sri Lankan species, 38 are endemic (Ranasinghe 2017). Among them, few showed island-wide distribution, while most showed restricted distribution and were found in only one, two or evolsyst.pensoft.net U. G. S. L. Ranasinghe & Suresh. P. Benjamin: Sri Lankan goblin spiders 266 three sites among more than hundred sampling localities. B. ratnapura, B. parumpunctata, B. ambigua, X. kandy, X. padaviya, A. vestigator, O. spinosiscorona showed widespread distribution, while soft bodied O. manicata was reported only from the island’s central montane wet evergreen forests. Certain species were found only in confined areas such as B. ondaatjei is known only from two sites in the Badulla District (L 50, L 30; 990–1280 m). B. carlmulleri is known only from Hantane and Knuckles mountain ranges in the Kandy District (L 43, 44; 585–1240 m). X. pophami is reported only at two sites (L 17, L 9) in the Matale District and X. nuwaraeliya is reported at three sites (L 53, L 54, L 55; >1700 m) in the Nuwara Eliya District. A. vankhedei is recorded from Udawattakele FR and Dunumadalawa FR, secondary forests located within Kandy City (L 1, L 27; 580–600 m). A. koslandensis was found in a small forest patch between Koslanda-Beragala, Namunukula and Bandarawela (L 52, L 51, L 67; 1000–1800 m). S. saaristoi Labugama FR, Kalugala FR (L 61, L 41; 40–200 m). Meanwhile, most species were found only the type locality: A. rawanaellensis (Rawana Ella), B. meemure (L 29; 636 m), B. shyami (L 28), Cavisternum bom (Mandaitivu FR), S. snippy (L 17), S. tiggy (Corbett′s Gap), P. tumpy (Mihinthale L 48), P. snooky (Ethagala FR, L 10), Ischnothyreus chippy (L 52) (see fig. 1 Ranasinghe and Benjamin 2016a, figs 6, 13 Ranasinghe and Benjamin 2016b, fig. 11 Ranasinghe and Benjamin 2018 a; fig. 21 Ranasinghe and Benjamin 2018b; fig. 5 Ranasinghe and Benjamin 2025). It is premature to comment on the origin of Sri Lanka’s goblin spider diversity as the Indian fauna is not well known. However, this study suggests that both ecology and evolution may be contributing factors in their evolution. Considering phylogenetic analysis and species distribution patterns, we can hypothesise that most of this diversity is generated through within-island speciation. This is seen in clades such as Aprusia, Brignolia and Xestaspis, which consist of closely related assemblages of more than two endemic species (Ranasinghe and Benjamin 2016a, b, 2018a, b, 2025). These species are narrow endemics with very restricted distribution and are not expected to be found outside of Sri Lanka. Aprusia, Brignolia and Xestaspis are morphologically revised, and Indian species are known and differ from Sri Lankan species (Baehr and Ubick 2010; Grismado et al. 2011; Platnick et al. 2011; Grismado 2023). Acknowledgements This study was funded by the National Institute of Fundamental Studies, Kandy. Special thanks to N. Athukorala for support in the field. Thanks to H. Sandamali, S. Batuwita, C. Clayton, N. Kanesharatnam, I. Sadunika all of NIFS for collecting some of the described materials. We are grateful to Norman Platnick, Darrell Ubick, Yvonne Kranz-Baltensperger and Arnaud Henrard for identification of some genera during the study. 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