Zoosyst. Evol. 99 (1) 2023, 209-216 | DOI 10.3897/zse.99.99285 > PENSUFT. yee BERLIN A new species of the genus Si/er Simon, 1889 (Araneae, Salticidae, Chrysillini) from India John T. D. Caleb’, Ayush Parag*, Aniruddha Datta-Roy* 1 Department of Anatomy, Saveetha Medical College & Hospital, Saveetha Institute of Medical and Technical Sciences, Chennai 602503, Tamil Nadu, India. 2 School of Biological Sciences, National Institute of Science Education and Research, An OCC of Homi Bhabha National Institute, Khordha, Jatni 752050, Odisha, India https://zoobank. org/E4E17325-0928-4F F 9-83 D-67EAEB 964128 Corresponding author: John T.D. Caleb (caleb87woodgate@gmail.com) Academic editor: Danilo Harms # Received 26 December 2022 @ Accepted 27 February 2023 Published 9 March 2023 Abstract A new chrysilline jumping spider species belonging to the genus Si/er Simon, 1889 is described from Odisha, India. Detailed mor- phological descriptions, illustrations of the male palp and female genitalia and phylogenetic relationships of the new Si/er species are presented. Phylogenetic analysis reveals that the new species is sister to a clade of predominantly Southeast Asian Si/er species. Furthermore, the results indicate the presence of multiple cryptic species masquerading as S. semiglaucus sensu lato. We also briefly discuss some unique behavioural observations on the newly-described species. Key Words Jumping spider, NISER, Odisha, phylogeny, species description, taxonomy Introduction The chrysilline jumping spider genus Si/er Simon, 1889 presently includes 11 described species distributed from southern to eastern Asia (WSC 2023). Members of the ge- nus are conspicuously coloured with iridescent scales (Hill 2009; Baba 2010; Kulkarni and Joseph 2015) and some Species are reported to be myrmecophagous (Jackson et al. 1998; Nelson et al. 2004; Touyama et al. 2008). Siler semi- glaucus (Simon, 1901) a widespread species distributed across tropical Asia is the only representative known from India until now (Kulkarni and Joseph 2015; Sen et al. 2015; Roy et al. 2016; Dhali et al. 2017; Tyagi et al. 2019; Caleb and Sankaran 2023). During recent surveys, we discovered a population of an undescribed species of the genus Si/er from Odisha State, India. Members of the newly-discov- ered population had distinct morphological characters dif- ferent from those of S. semiglaucus. Phylogenetic analysis using the Maximum Likelihood (ML) approach corrobo- rates the morphological findings. In this paper, we describe this newly-discovered population as a new species. Materials and methods Specimens were hand collected and preserved in 70% ethanol. Images of live specimens were captured using a Nikon D500 camera with AF-S Micro Nikkor 60 mm macro lens. Morphological examination was carried out under a Leica SAPO stereomicroscope and photomicro- graphs were taken with a Leica MC190 HD camera and processed with the Leica Application Suite (LAS) ver- sion 4.13. The male left palp was detached and exam- ined in detail and photographed. The temporary mount of the epigyne was studied under a Leica DM3000 LED compound microscope and photographed with a Leica MC190 HD camera. The line drawings were prepared using the GNU Image Manipulation Program (GIMP) (Montesanto 2015). The GPS coordinates of the col- lection localities were taken with the help of a Garmin GPSMAP 66S (Datum WGS 84). The map was prepared using QGIS (v.3.12.3) software. Body and eye measure- ments follow Zabka (1991). Leg measurements are given as follows: total (femur, patella, tibia, metatarsus, tarsus). Copyright Caleb, J.T.D. et al. 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. 210 All measurements are in millimetres (mm). The studied Specimens are deposited in the Southern Regional Cen- tre, Zoological Survey of India (ZSIC), Chennai, Tamil Nadu, India. For phylogenetic analysis, the total genomic DNA was extracted from a complete specimen (collected apart from the deposited type material), using a Qiagen™ Blood and Tissue kit. A region of the mitochondrial CO-I was PCR amplified (see Table 1 for primer details and thermocy- cler conditions). The PCR product was purified using a QIAGEN PCR purification kit and was sent for sequenc- ing at Heredity Life Sciences Pvt. Ltd., Bhubaneswar. The generated sequence was aligned with other Siler species (see Table 2 for details) in MEGA V.5.2 (Tamura et al. 2011) using MUSCLE (Edgar 2004). The alignment was uploaded on the IQ-Tree online por- tal (Trifinopoulos et al. 2016; http://igtree.cibiv.univie. ac.at/) to estimate the best model of sequence evolu- tion using a Bayesian Information Criterion (BIC) and subsequently generated a Maximum Likelihood tree. The branch support was estimated using 1000 ultrafast bootstrap replicates. Two species of the genus Myrma- rachne were used as outgroups. Abbreviations used in the text: AER — anterior eye row; ALE — anterior lateral eye; AME — anterior median eye; EFL - eye field length; PER — posterior eye row; PLE — posterior lateral eye; PME — posterior median eye; RTA — retrolateral tibial apophysis. Species description Genus Siler Simon, 1889 Type species. Si/er cupreus Simon, 1889. Siler niser sp. nov. https://zoobank.org/08C8AFAD-A46EF-4330-99E3-28336877D588 Figs 1A-F, 2A-L, 3A-D, 4A, B, 5 Type material. Holotype ¢ (ZSIC-I/SP 40) from INDIA: Odisha, Bhubaneswar, Khordha, Jatani, NISER cam- pus (20.16861°N, 85.68493°E), 46 m as.l., 06.v.2021, leg. A. Parag. Paratypes: 4 29 (ZSIC-I/SP 41 to 44), data same as holotype, 05.v.2021. Caleb, J.1.D. et al.: New Siler species from India Etymology. The specific epithet is an acronym derived after the type locality, National Institute of Science Ed- ucation and Research (NISER) campus from where the specimens were collected. The name is treated as a noun in apposition. Suggested common name. Glossy jumping spider. Diagnosis. Si/er niser sp. nov. resembles Siler semiglaucus (Simon, 1901) in general morphology and colour pattern (cf. Fig. 1A, B, D, E with images la, b in Kulkarni and Joseph (2015)), but can be easily dis- tinguished by the morphology of the copulatory organs which are rather most similar to S. cupreus Simon, 1889 and S. severus (Simon, 1901): male palp with short beak- like embolus (slender and needle-like in S. cupreus and S. severus);, relatively smaller RTA (smallest of all con- geners, remaining below the small retrolateral tegular lobe); RTA directed anteriad in ventral view, with point- ed tip and broad base in retrolateral view, ventral margin vertical, dorsal margin gradually sloping, gently curved (RTA directed retrolaterally in ventral view, thick, rela- tively long reaching beyond the retrolateral tegular lobe and curved in S. cupreus and S. severus) (cf. Figs 2E, F, 3A, B with illustrations on pg. 133, 135 in Proszynski (1984), figs 246, 247 in Bohdanowicz and Proszynski (1987) and figs 12, 13 in Proszynski (1985)); females can be recognised by the short and bent copulatory ducts and globular spermathecae separated by more than their ra- dius (copulatory ducts relatively longer, almost straight and copulatory openings closely placed along the median in S. cupreus) (cf. Figs 2K, L, 3C, D with illustrations on pg. 134, 135 in Proszynski (1984) and figs 249, 251 in Bohdanowicz and Proészynski (1987)). Description. Male (based on holotype ZSI-I/SP-40, co- louration in alcohol) (Fig. 2A—F): total length 4.10; cara- pace 1.97 long, 1.50 wide; abdomen 2.13 long, 1.19 wide. Carapace greenish-brown; lateral sides with broad red patch; outer rim of carapace lined by blue scales (Fig. 2A). Anterior eyes surrounded by greyish-white orbital setae. Clypeus brown, ‘cheeks’ covered with bluish hairs (Fig. 2D). Eye measurements: AME 0.38, ALE 0.21, PME 0.07, PLE 0.19; AER 1.21; PER 1.34; EFL 0.95. Clypeus height 0.05. Sternum oval, yellowish-brown, covered with hairs; labium and maxillae brown (Fig. 2B). Chelicerae brown with two promarginal teeth and one fissident retromarginal tooth. Abdomen with an anchor-shaped red marking in the anterior three quarters; four grey spots composed of metallic Table 1. Thermocycler condition and details of primer used for PCR reaction. Primer name Thermocycler conditions Reference LCO-1490 (forward primer) Initial denaturation: 94 °C, 3 min Denaturation: 94 °C, 30s Annealing: 45 °C, 30s Extension: 72 °C, 90s Final extension: 72 °C, 7 min Bork RJ (2015) Primer efficacy in the DNA barcoding of spiders. Honours Program Theses. 169. https://scholarworks.uni.edu/hpt/169 HCO-700ME (reverse primer) __ Initial denaturation: 94 °C, 3 min Denaturation: 94 °C, 30s Annealing: 45 °C, 30s Extension: 72 °C, 90s Final extension: 72 °C, 7 min zse.pensoft.net Bork RJ (2015) Primer efficacy in the DNA barcoding of spiders. Honours Program Theses. 169. https://scholarworks.uni.edu/hpt/169 Zoosyst. Evol. 99 (1) 2023, 209-216 A Ziel Figure 1. General habitus of Siler niser sp. nov. A. Male habitus, dorsal view; B. Same, lateral view; C. Same, front view; D. Female habitus, dorsal view; E. Same, lateral view; F. Same, front view. scales present in the anterior portion; lateral and posterior half dark grey, composed of iridescent scales; venter brown, covered with iridescent scales; light brown below the epi- gastric region (Fig. 2A, B). Spinnerets yellowish-brown. Legs yellow with longitudinal brown streaks; leg I robust, brown, femur and patella with ventral fringe of dense black hairs, tibia with both ventral and dorsal fringe (Fig. 2A—C). Leg measurements: I 4.87 (1.64, 0.73, 1.08, 0.87, 0.55); II 3.45 (1.08, 0.54, 0.72, 0.68, 0.43); III 3.97 (1.20, 0.52, 0.80, 0.93, 0.52); TV 5.36 (1.63, 0.62, 1.18, 1.36, 0.57). Leg for- mula 4132. Palp as shown in Figs 2E, F, 3A, B. Female (ZSI-I/SP-41, colouration in alcohol) (Fig. 2G—-L). Total length 5.31; carapace 2.24 long, 1.69 wide; abdomen 3.07 long, 2.05 wide. Carapace brown; covered with black hairs; lateral margins covered with reddish hairs, outer rim lined by bluish-white hairs (Fig. 2G). Anterior eyes surrounded by greyish-white or- bital setae. Clypeus brown, ‘cheeks’ covered with bluish hairs below ALEs (Fig. 2J). Eye measurements: AME 0.40, ALE 0.24, PME 0.09, PLE 0.23; AER 1.33; PER 1.50; EFL 1.04. Clypeus height 0.02. Sternum oval, yel- low; labium and maxillae brown, maxillae apically paler (Fig. 2H). Chelicerae brown with two promarginal teeth and one fissident retromarginal tooth. Abdomen grey- ish-black with a median transverse black patch: venter yellow, lateral and posterior region brown (Fig. 2G—I). Spinnerets brown. Legs brownish-yellow with longitudi- nal black stripes (Fig. 2G, I). Leg measurements: I 4.30 (1.45, 0.77, 0.85, 0.76, 0.47); II 3.62 (1.18, 0.61, 0.69, Table 2. Accession numbers of samples used for the phyloge- netic analysis. Species name Accession number Siler semiglaucus KY888770 Siler semiglaucus MK392837 Siler collingwoodi LC485246 Siler cupreus LC485236 Siler ruber LC485239 Siler niser sp. nov. 0Q553823 Myrmarachne formicaria MZ626812 Myrmarachne robusta MK154679 0.69, 0.45); III 4.13 (1.25, 0.62, 0.79, 0.99, 0.48); [IV 5.59 (1.71, 0.72, 1.21, 1.40, 0.55). Leg formula 4132. Epigyne and vulva as shown in Fig. 2K, L, 3C, D. Variation. Total length of females ranges from 5.31 to 6.88 (n =4). Colour in life. Male. Carapace covered with a mixture of bluish and greenish scales; blue scales extending below the ALEs and running along the rim; lateral sides covered with a thick band of red scales (Fig. 1A—C). Legs yellow brown with a few interspersed iridescent scales and dark brown longitudinal stripes; leg I dark brown with white proximal half of metatarsi and white tarsi. Palps brown, cymbium, tibia and patella covered with white hairs. Abdomen covered with iridescent scales along anterior, lateral and posterior regions; anterior half with a distinct pattern composed of a reddish anchor-shaped mark and four bright blue spots (Fig. 1A—C). zse.pensoft.net 212 Caleb, J.T.D. et al.: New Siler species from India Figure 2. Somatic and genital morphology of Siler niser sp. nov. A-F. Holotype male (ZSI-I/SP 40) A. Dorsal view; B. Ventral view; C. Lateral view; D. Front view; E. Left male palp, ventral view; F. Same, retrolateral view. G—L. Female paratype (ZSI-I/ SP 41); G. Dorsal view; H. Ventral view; I. Lateral view; J. Front view; K. Epigyne, ventral view; L. Vulva, dorsal view. Scale bars: 2 mm (C, I); 1 mm (A, B, G, H, J); 0.5 mm (D); 0.2 mm (E, F, L); 0.1 mm (K). zse.pensoft.net Zoosyst. Evol. 99 (1) 2023, 209-216 213 Figure 3. Genital morphology of Siler niser sp. nov. A. Left male palp of holotype (ZSI-I/SP 40), ventral view; B. Same, retrolateral view; C. Epigyne of paratype (ZSI-I/SP 41), ventral view; D. Vulva, dorsal view. Scale bars: 0.2 mm (A, B, D); 0.1 mm (C). Female. Carapace covered with white and grey scales; lateral margins covered with reddish scales; outer rim covered with bluish-white scales. Legs yellow-brown with longitudinal dark brown and white stripes. Abdomen greyish with a similar, but incomplete pattern as seen in male; a median transverse red band sandwiched between two blue bands; anterior blue band discontinuous medi- ally, posterior band broad and continuous (Fig. 1D, E). In darker females, the red band is completely black, this perhaps occurring in older females when the scales are lost (Fig. 1F). Natural history. Specimens were found in acute vi- cinity of the ant Camponotus compressus. Individuals were myrmecophagic like Siler semiglaucus (confirmed through visual observations). Both sexes waved the first pair of legs in the air, perhaps mimicking the antennae of the ants (Fig. 1B, F). This behaviour was observed both in the presence as well as in the absence of ants. Individ- uals were extremely agile and moved to the underside of leaves as soon as they detected any external movement. Specimens were collected from Hyptis suaveolens and Lannea coromandelica seedling foliage, both about 2 feet (ca. 60 cm) above the ground, during the month of May. The average humidity during the collection period was 63% (range: 62% to 64%). Threat status. The individuals of Siler niser sp. nov. were collected from a grassland-shrubland type habitat (Fig. 4A, B). These biomes are under threat as they are often labelled as unproductive and the biodiversity they harbour remains poorly documented (Nerlekar et al. 2022). Since Siler niser sp. nov. is known only from its type locality (Odisha, India), other possible locations of its occurrence are yet unknown. Due to lack of distribu- tional data especially in grasslands where surveys are most often ignored, we categorise Siler niser sp. nov. as data deficient. Distribution. Known only from the type locality (Odisha, India) (Fig. 5). Phylogeny. The best model of sequence evolution according to BIC was TRN+I+G. The obtained ML tree (Fig. 6) was rooted using the branch leading to the two outgroup taxa. The new species was sister to a clade con- sisting of S. collingwoodi, S. cupreus and S. ruber, albeit with low bootstrap support. The uncorrected p-distance within Siler species ranged from 1.3% to 7.4%. The new species was also morphologically different from all other congeners with a suite of morphological characters as described above. The members of the pop- ulation we collected from Odisha State, India share char- acters like the body covered with iridescent scales, abdo- men with transverse banding pattern and spots, tibia I in males with typical dense bottle brush-like setae, male palp with elongated bulbus and flattened, spatulate RTA, female with simple rounded epigyne and short copulatory ducts zse.pensoft.net 214 A Siler niser sp. nov. © Siler semiglaucus 65.0°E 70.0°E I | Caleb, J.T.D. et al.: New Siler species from India B Figure 5. Map showing the current distribution of the genus Si/er in India. (Proszynski 1985; Zabka 1985) with congeners, but are distinct both morphologically and phylogenetically. There- fore, we use the general lineage concept in order to delin- eate this population as a new species (de Quiroz 1998). Discussion We used multiple lines of evidence to establish the new- ly-found population as a new species. The morphological diagnosis provides clear evidence for the distinction of this zse.pensoft.net new population of Si/er species. Given the known distri- bution of jumping spiders of the genus Si/er, we expected a sister relationship between our new species and the only known congener from India — Siler semiglaucus. Surpris- ingly, our phylogenetic analysis retrieved S. niser sp. nov. as sister to a clade containing the Southeast Asian lin- eages namely — S. collingwoodi, S. cupreus and S. ruber. We included two published sequences of S. semiglaucus (Specimen K Y888770 from Sri Lanka, Specimen 392837 from India) (Kanesharatnam and Benjamin 2019; Tyagi et al. 2019). These specimens, however, were retrieved as Zoosyst. Evol. 99 (1) 2023, 209-216 0.04 21D Siler semiglaucus (KY 888770) Siler semiglaucus (MK392837) Siler niser sp.nov. Siler collingwoodi (LC485246) Siler cupreus (LC485236) Siler ruber (LC485239) Myrmarachne formicaria (MZ626812) Myrmarachne robusta (MK154679) Figure 6. Maximum Likelihood tree of the genus Si/er using an unpartitioned partial CO-1 gene. The numbers near the nodes rep- resent branch support values. sister lineages, albeit with low bootstrap support. Although S. semiglaucus 1s considered a widespread species, such low bootstrap support indicates a species complex encompass- ing multiple independently evolving lineages that require further investigation. In addition, our findings suggest that the low number of species observed tn the genus Si/er may be an artefact of inadequate sampling. A taxonomic revision of jumping spiders of the genus Si/er is thus warranted. The behaviour of individuals of S. niser sp. nov. is es- pecially intriguing. Both sexes bob their abdomens and wave their front pair of legs to produce a type of behaviour termed as “antennal illusion”. This type of behaviour is commonly observed in various species of ant-mimicking spiders (Ceccarelli 2008; Shamble et al. 2017). Although S. niser sp. nov. bears no visual resemblance to an ant, it may utilise such behavioural adaptations to infiltrate ant colonies, as ants are their preferred prey (pers. obs. AP). Moreover, they may bear a chemical resemblance to the ants, which may also enable them to stay in close vicini- ty with the ants despite bearing no morphological resem- blance (Uma et al. 2013). These traits may also grant pro- tection from predators, which is the most commonly cited explanation for the evolution of such processes (Ceccarelli 2008; Uma et al. 2013; Shamble et al. 2017). Since such types of behaviour have been documented in S. semiglau- cus (Grob 2015), we suspect that the presence of the same mymercophagy is prevalent in other Si/er species (Jackson et al. 1998: Nelson et al. 2004; Touyama et al. 2008; Grob 2015). We recommend that the evolution and functional ecology of such behaviour in the brilliantly coloured spi- ders of the genus Si/er are investigated in the future. Acknowledgements We are grateful to Dr K.A. Subramanian, Offi- cer-in-Charge, SRC, ZSI for kindly allowing JC to use the microscope facility. Thanks to Mr. Shubhang Srikanth Acharya (NISER Bhubaneswar) for the help and support during the fieldwork. ADR and AP thank the Department of Atomic Energy and NISER Bhubaneswar for their fa- cilities and support. 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