INTRODUCTION

The Oonopid fauna of India currently com prises 52 species (more than half endemic) in 15 genera (^{Tiwari et al., 2021}), of which Brignolia Dumitrescu & Georgescu, 1983 is the most di verse, with 16 species (WSC, 2023). The known diversity in this country is the product of revi sionary work carried out in recent decades in the frame of the Goblin Spider PBI Project (e.g., ^{Baehr & Ubick, 2010}; ^{Platnick et al. 2011}, 2012; ^{Grismado et al., 2014}). However, the territorial extension of India and the numerous regions that have not been surveyed suggest a more ex tensive diversity. In addition, some taxonomic entities require to be reassessed in systematic studies (Tiwari et al., 2021: 8341).

In the Goblin Spider PBI Project context, I examined some museums collections and found new species herein proposed. They belong to the oonopid genera Paramolotra Tong & Li (from West Bengal) and Aprusia Simon (from Tamil Nadu).

Paramolotra, which had recently been de scribed on base of two species from Tibet, in China (^{Cheng et al., 2021}), is a genus of hard-bodied oonopids recognizable by the conical pro trusions on the basal anterior face of chelicerae, by the large bulb fused with the cymbium, and the complex elements at the embolar region (Cheng et al., 2021:56).

The genus Aprusia is a group whose most re markable diversity (seven out of eight species) is known from Sri Lanka (^{Ranasinghe & Benjamin, 2018}). The known distribution of this genus ap pears as fitting in the Western Ghats-Sri Lanka hotspot of biodiversity, where it seems to be en demic. Nevertheless, the unity of the two com ponents of that region (southwest of mainland India and Sri Lanka) has recently been ques tioned (^{Bossuyt et al., 2004}; ^{Gunawardene et al., 2007}).

Aprusia comprises goblin spiders with ab dominal scutae, similar to Ischnothyreus Simon and Camptoscaphiella Caporiacco, although less sclerotized (see, for example, ^{Baehr & Ubick, 2010}; ^{Edward & Harvey, 2014}). They also differ in the genitalia, particulary the male palps, with the bulb fused to the cymbium and a tiny, slightly sclerotized embolus.

When I studied specimens housed in the California Academy of Sciences, San Francisco, I found two morphospecies in a same vial labeled as the same species. One of them was assigned to the species A. kerala^{Grismado & Deeleman, 2011}, previously known from males collected in neighboring Kerala, and a new morphospecies that I present in this work.

^{Ranasinghe & Benjamin (2018}) presented an identification key and a cladistic analysis of the genus Aprusia. These authors propose three unambiguous synapomorphies for this genus: smooth male endites, two prolateral spines on the femur I, and copulatory opening inconspicu ous, located in the epigastric furrow. All know females for this genus have a narrow anterior receptacle, apparently without lumen, connect ed with muscle bunches, for what presumably serves as sites for muscle attachment (as apo demes) rather than sperm storage. The female specimens studied here showed some unique fea tures compared with the previously known spe cies, e.g., paired seminal receptacles present in A. kerala, and the absence of scutal ridges in the new morphospecies. On the other hand, Paramolotra and Aprusia have the basal article of the ante rior lateral spinnerets entire (not crossed by a diagonal membranous area), and the males have the sperm pore exposed; then, both genera could be included in the “higher gamasomorphines” sensu^{Grismado et al. (2014}: 7).

In the present contribution, I describe two new species belong to Paramolotra and Aprusia genera, describe for the first time the female of Aprusia kerala, and increase the knowledge of Indian goblin spider fauna.

MATERIAL AND METHODS

Specimens are deposited in the following col lections: Museum d’histoire naturelle de la Ville de Geneve (MHNG, Peter Schwendinger) and California Academy of Sciences (CAS, Lauren Esposito). The descriptions were generated auto matically from the Species Descriptive Database of the oonopid Planetary Biodiversity Inventory project. Concerning the terminology of the male genitalic structures, I reinterpreted them in light of my own observations, as explained below (see the taxonomic section). Because the right palp of the Paramolotra specimen was accidentally sepa rated, the images were generated on it and then digitally inverted to facilitate comparison with the other species. Female genitalia were observed in clove oil. Drawings were made with a camera lucida mounted on an Olympus BH-2 compound microscope. Photographs of the preserved speci mens were taken with a Leica DFC 290 digital camera mounted on a Leica M165 C stereoscopic microscope, and the focal planes were aligned with Helicon Focus 4.62.2. All measurements are in millimeters. Due to the scarcity of museum specimens, I avoided using irreversible tech niques-such as scanning electron microscope-to document the animals’ anatomy fully. When descriptions of new species are made through a single specimen (holotype), to preserve it, genita lia dissection was not performed. In some cases, I provide approximate coordinates (denoted by “ca.”) calculated with Google Earth (http://earth.google.com) from label data.

Abbreviations: ar, anterior receptacle; bp, ba sal protrusion; co, copulatory opening; e, embo lus; la, lateral apodeme; pp, posterior protrusion; r, ridge; sr, seminal receptacles.

TAXONOMY

Family Oonopidae ^{Simon, 1890}

Genus Paramolotra Tong & Li, 2021

Paramolotra Tong & Li, 2021 in ^{Cheng et al., 2021}: 56, (type species Paramolotra pome Tong & Li, by original designation).

Remarks. The authors of the genus labeled the structures of the male embolar region naming them by unspecific positional criteria: for exam ple, “basal protrusion”, “anterior protrusion”, “posterior protrusion”, “dorsal protrusion”, “an terior membrane”, and “posterior membrane” (Cheng et al.: figs. 2, 5). However, in the cleared preparation of P. bengalensis (Fig. 1), I located the ejaculatory opening in the structure that seems to be equivalent to the «anterior protrusion» (at least in P. metok, see Chen et al., 2021: fig. 5 C, E-F). Then, I interpret this sclerite as the em bolus, and for the remaining closer structures, I follow the terminology of Chen et al. (2021) only for easier comparisons.

Paramolotra bengalensis, new species

LSID: urn:lsid:zoobank.org:act:AADBB1E4-96 D1-41F9-AD4E-33B4BD2B128E

(Figs. 1-3, 7)

Fig. 1 Paramolotra bengalensis, new species, holotype male. (A) right palp, cleared (inverted), prolateral; (B) detail of the embolar region, dorsal; (C) same prolateral; (D) same retrolateral. Abbreviations: bp, basal protrusion; e, embolus; pp, posterior protrusion. 

Fig. 2 Paramolotra bengalensis, new species, holotype male. (A) habitus dorsal; (B) same, ventral; (C) same lateral; (D) carapace, dorsal view; (E) same, ventral; (F) same, lateral. 

Fig. 3 Paramolotra bengalensis, new species, holotype male. (A) carapace anterior; (B) right palp (inverted) dorsal; (C) same prolateral; (D) same retrolateral. 

Diagnosis. The male of P. bengalensis is distin guished by the orange color of the sclerotized parts integuments (Fig. 2A-F), brown or yellow brown in the Chinese congeners P. pome Tong & Li and P. metok Tong & Li, (^{Cheng et al. 2021}: figs. 1, 3-4, 6), and by the details of the embolar region: P. bengalensis has the posterior protru sion located retrolaterally. In addition, the embo lus is slender, with a flattened and expanded tip curved backwardly, and the ejaculatory opening on its prolateral side (Figs. 1B-D).

Description. Male (holotype). Total length 1.88. Cephalothorax: Carapace orange, without any pattern, pyriform in dorsal view, pars cephali ca strongly elevated in lateral view, anteriorly narrowed more or less 0.5 times its maximum width, with rounded posterolateral corners, pos terior margin not bulging below posterior rim, anterolateral corners without extension or pro jections, posterolateral surface without spikes, surface of carapace smooth, thorax without de pressions, fovea absent; lateral margin straight, rebordered, without denticles; setae absent, the grabrous carapace is probably due by the loss of the setae, because some few bases are visible on the pars cephalica, clypeus and margins. Clypeus margin unmodified, sinuous in front view, ver tical in lateral view, high, ALE separated from edge of carapace by more than their diameter. Chilum absent. Eyes six, well developed, all sub equal, all eyes circular; posterior eye row straight from above, procurved from front; ALE separat ed by less than their radius, ALE-PLE separated by less than ALE radius, PME touching for less than half their length, PLE-PME separated by less than PME radius. Sternum orange, longer than wide, uniform, not fused to carapace, with radial furrows between coxae I-II, II-III, III-IV (furrows consisting in aligned large pits), ster num covered with small round pits, microsculp ture covering entire surface, anterior margin unmodified, posterior margin not extending pos teriorly of coxae IV, anterior corner unmodified, distance between coxae approximately equal; setae sparse, light, needle-like, evenly scattered, originating from surface. Mouthparts: chelicerae straight, with prominent, small, basal process; fangs without tooth-like projections, directed medially, tip unmodified; setae light, needle-like; paturon inner margin with pairs of enlarged se tae, distal region abruptly narrowed. Labium tri angular, not fused to sternum, anterior margin indented at middle, same as sternum in sclero tization, chelicerae, endites and labium orange. Endites distally not excavated, unmodified, same as sternum in sclerotization, with serrula present in single row. Abdomen: ovoid, rounded posteriorly. Book lung covers large, elliptical, without setae, anterolateral edge unmodified. Posterior spiracles not connected by groove. Pedicel tube medium, ribbed, scutum extend ing far dorsal of pedicel. Dorsal scutum strongly sclerotized, orange, without color pattern, cover ing full length of abdomen, no soft tissue visible from above, not fused to epigastric scutum, sur face smooth. Epigastric scutum strongly sclero tized, orange, surrounding pedicel, not protrud ing. Postepigastric scutum strongly sclerotized, orange, almost semicircular, covering about 2/3 of abdominal length, fused to epigastric scutum, anterior margin unmodified. Spinneret scutum absent. Dorsum setae present, light, needle-like. Epigastric area setae uniform, light, needle-like. Postepigastric area setae light, needle-like. Basal article of the ALS entire (not crossed by a diago nal membranous area). Colulus absent. Legs: pale orange, without color pattern; femur IV not thickened, same size as femora I-III, patella plus tibia I shorter than carapace, tibia I unmodified. Leg spination (only surfaces bearing spines list ed): leg I: tibia: v2-2-2-2-0, metatarsus: v2-2, leg II: tibia: v2-2-2-2-0, metatarsus: v2-2. Tarsus I to IV without inferior claw. Genitalia: Epigastric region with sperm pore small, circular, situated between anterior and posterior spiracles. Palp (Figs. 3B─D) very large, not strongly sclerotized, yellowish, proximal segments pale orange; trochanter normal size, unmodified; femur normal size, one to two times as long as trochanter, at taching to patella basally; patella shorter than femur, not enlarged; bulb nearly reniform, but tapering apically, cymbium completely fused with bulb, no seam visible, but discernible by their numerose setae; embolus with apical ori gin, flattened, and upwardly curved, with the ejaculatory opening on its prolateral side (Fig. 1B); at the distal part, there are two paraembolic elements, the prolateral one (“posterior protru sion”) is flattened, the retrolateral one (“basal protrusion”) looks hyaline, bearing a ventral fur row and with a rounded posterior tip pointing to backwards (Fig. 1B-D).

Female unknown.

Type material. Male holotype from India: West Bengal: Darjeeling: 13km north of Ghoom (way to Bijanbari), ca. N 27° 3’56”, E 88°15’29.83”, leaf litter, 1500m, Oct. 15, 1978, Besuchet, C. and Löbl, I., PBI_OON 15680, deposited at MHNG.

Etymology. The specific epithet refers to the Indian state where the type specimen was col lected.

Distribution. Only known from the type locality.

Genus Aprusia^{Simon, 1893}

Aprusia^{Simon, 1893}a: 295, (type species, by monotypy Aprusia strenuus Simon).

Aprusia kerala^{Grismado & Deeleman, 2011}

(Figs. 4, 6A─E, 7)

Fig. 4 Aprusia kerala, Grismado & Deeleman, female CASENT 9038464 (vch CJG-1964). (A) habitus dorsal; (B) same ventral; (C) same lateral; (D) carapace, dorsal view; (E) same ventral; (F) same anterior; (G) epigastric region ventral. 

Fig. 6 Aprusia spp. female genitalia; (A-E) A. kerala Grismado & Deeleman CASENT 9038464 (A) vch CJG-1964; (B-E) vch CJG-1983; (F) A. rothorum, new species, holotype. (A-C, E-F), cleared, ventral view; (D), cleared, dorsal view; (E) detail of right half of the fig. B compared at the same scale with the embolus of the male holotype (taken from Grismado et al., 2011). Abbreviations: ar, anterior receptacle; co?, copulatory opening; la, lateral apodeme; r, ridge; sr, seminal receptacles. 

Fig. 7 Distribution of the species here studied: Paramolotra bengalensis, new species (circle); Aprusia spp. (triangles). 

Aprusia kerala^{Grismado & Deeleman, 2011}, in Grismado et al., 2011: 18. Male holotype and male paratype from India: Kerala: Cardamom Hills, Muttapatti near Munnar, ca. N 9°52′, E 77°09′, 1700 m, Nov. 24, 1972, C. Besuchet and I. Löbl (deposited in MHNG, PBI_OON 12360, not reexamined).

Diagnosis. Female is recognizable by the re markable presence of paired seminal receptacles (Figs. 4G, 6A─D).

Remarks. The two females were not collected with the males. Still, they are matched by geo graphical proximity (ca. 56 km apart) and simi lar morphology: the holotype male’s low clypeus, the ocular configuration, and spine patterns (^{Grismado et al., 2011}: figs. 46-52) fit with these two females rather than the other sympatric spe cies. Thus, the other female is proposed to be long to a different new species (A. rothorum, see below).

Description. Female (CASENT 9038464, voucher CJG-1964). Total length 2.68. Cephalothorax: Carapace pale orange, without any pattern, broadly oval in dorsal view, pars cephalica strongly elevated in lateral view, ante riorly narrowed less to 0.50 times its maximum width, with rounded posterolateral corners, surface of elevated portion of pars cephalica smooth, sides finely reticulate, fovea absent; lateral margin straight, smooth, non-marginal pars cephalica setae mostly lost (bases remain); non-marginal pars thoracica setae mostly locat ed on the posterior slope; marginal setae light, needle-like. Clypeus margin unmodified, curved downwards in front view, vertical in lateral view, low, ALE separated from edge of carapace by less than their radius, median projection absent; se tae present, light, needle-like. Chilum absent. Eyes six, on dark pigment, well developed, all subequal, all eyes oval; posterior eye row straight from above, procurved from front; ALE sepa rated by less than their radius, ALE-PLE sepa rated by less than ALE radius, PME touching for less than half their length, PLE-PME sepa rated by less than PME radius. Sternum longer than wide, pale orange, uniform, not fused to carapace, without radial furrows between coxae, surface smooth, without pits, anterior margin unmodified, posterior margin extending posteri orly beyond anterior edges of coxae IV as single extension, anterior corner unmodified, distance between coxae approximately equal, extensions of pre-coxal triangles present, lateral margins with narrow extensions between coxae; setae sparse, light, needle-like, evenly scattered, origi nating from surface. Mouthparts: Chelicerae, endites and labium pale orange. Chelicerae straight, anterior face unmodified; without teeth on both promargin and retromargin; fangs with out tooth-like projections, directed posteriorly, without prominent basal process; setae light, needle-like, evenly scattered; paturon inner mar gin with pairs of enlarged setae. Labium wide, not fused to sternum, anterior margin indented at middle, same as sternum in sclerotization; subdistal portion with two needlelike setae. Endites relatively short, anteriorly directed, par allel, same as sternum in sclerotization. Female palp claws absent; spines absent; tarsus unmodi fied. Abdomen: ovoid, rounded posteriorly; dor sum soft portions white, without color pattern. Book lung covers large, elliptical, without setae. Posterior spiracles not connected by groove. Pedicel tube short, unmodified, scuto-pedicel re gion unmodified, scutum extending far dorsal of pedicel. Dorsal scutum weakly sclerotized, pale orange, without color pattern, covering less than 1/2 of abdomen length, more than 1/2 to most of abdomen width, fused to epigastric scutum, surface smooth. Epigastric scutum weakly scle rotized, surrounding pedicel, not protruding. Postepigastric scutum slightly more sclerotized, pale orange, larger than in congeners, semicir cular, covering about 1/4 of abdominal length, fused to epigastric scutum, anterior margin un modified. Spinneret scutum absent. Abdominal setae, light, needle-like. Basal article of the ALS entire (not crossed by a diagonal membranous area). Colulus represented only by setae. Legs: pale orange, without color pattern; femur IV not thickened, same size as femora I-III, patella plus tibia I near as long as carapace. Leg spina tion (only surfaces bearing spines listed,): leg I: femur: pv0-0-1-1-1-0, rv0-0-1-0-1-0, tibia: pv1-1-1-1-1-0, rv0-0-1-1-1-1-0, metatarsus: v2-2-0, leg II: femur: pv0-0-1-1-0-0, rv0-0-1-0-1-0, tibia: pv1-1-1-1-1-0, rv0-0-1-1-1-1-0, metatarsus: v2-2-0, leg III: tibia: dp1-1, dr1-1, vp0-1-1, metatar sus: dp1-1, dr1-1,pv0-1-1,v0-1-0, leg IV: patella: d0-2, tibia: pd1-0, p0-1, v1-0-1, metatarsus dp1-1, dr1-0, v0-1-2. Tarsi III and IV with a pair of claw like setae. Genitalia: Epigastric scutum with two short, transversal ridges; epigastric furrow very narrow compared with the other known species of the genus; anterior receptacle very thin and short, arising from a short transversal sclerite; lateral apodemes thick, parallel, and posteriorly directed; two rounded seminal receptacles deep orange (Figs. 4G, 6A-D), surrounded by mem branous tissue (Fig. 6D), two very small and nar row slits (hardly visible with light microscope) in the postepigastric scutum, at the level to the seminal receptacles, apparently related to the darkened regions, that are interpreted as a in vaginations or orifices for sperm ingression (co ? in Fig. 6A-C, E).

Material examined. India: Tamil Nadu: Kodaikanal, native cloud forest by falls. 10° 15´N, 77° 31´E. 2 January 1990. V. & B. Roth colls. (two females, CASENT 9038464 [APRUS DU 138], PBI_OON 00036125, vouchers vch CJG-1964 and CJG-1983).

Distribution. Southwestern India (Kerala and Tamil Nadu states).

Comments. The double receptacular condition is, of course, very surprising. It is a feature not observed in all the superfamily Dysderoidea. However, since the posterior receptacle of all Aprusia species is fragile and is usually destroyed despite extreme care during dissection (see ^{Grismado et al., 2011}: figs. 44 and 45), I cannot discard that the two rounded, dark receptacular elements found in A. kerala could be internal ele ments of a more inclusive, unpaired, thin walled, posterior receptacle, presumably broken during preparation. But the presence of two narrow slits apparently connected with the seminal recepta cles is suggestive, and forces to consider the pos sibility that they could be paired copulatory ori fices. A comparation of size of the male embolus with the presumably copulatory openings (Fig. 6E) suggest that this is a reasonably interpre tation. Unfortunately, the scarcity of specimens avoids trying addicional explorations, as SEM (one female was dissected, and the other one was only observed clearing the entire abdomen; Fig. 6A). The only other possible interpretation of this unusual paired structure is that they could be extreme modifications of internal apodemes or other unique structures related with the semi circular sclerotizations of other congeneric spe cies. Again, we need to find additional specimens for more exhaustive study.

If the receptacular invaginations are, in fact, orifices for sperm ingression, this would be a drastic morphological change, which would make them functional entelegyne spiders. It is un doubtedly a promising genus to study in greater detail in the future because it may be a case of radical reorganization of the genitalia not only for Oonopidae, but also for the entire superfami ly Dysderoidea.

Aprusia rothorum, new species

LSID: urn:lsid:zoobank.org:act:FEC3EB88-94 DC-487A-B67A-4C9D8F652BDF

(Figs. 5, 6F, 7)

Fig. 5 Aprusia rothorum, new species, female holotype. (A) habitus dorsal; (B) same ventral; (C) same lateral; (D) carapace, dorsal view; (E) same ventral; (F) same, anterior; (G) epigastric region, ventral. 

Diagnosis. Female of A. rothorum resembles to those of A. veddah, A. kataragama, and A. ko slandensis (^{Grismado et al., 2011}: figs. 44─45; ^{Ranasinghe & Benjamin, 2018}: fig. 2c) in having the anterior receptacle shorter than the lateral apodemes, but it is clearly distinguishable by the apodemes, more separated, and by lacking epi gastric ridges (Figs. 5G, 6F).

Description. Female (holotype). Total length 3.16. Cephalothorax: Carapace pale orange, without any pattern, broadly oval in dorsal view, pars cephalica slightly elevated in lateral view, anteriorly narrowed to less than 0.50 times its maximum width, with rounded posterolateral corners, surface smooth, fovea absent, lateral margin straight, smooth; setae light, needle-like. Clypeus margin straight in front view, vertical in lateral view, high, ALE separated from edge of carapace by more than their radius, median pro jection absent, cuticle traslucent, cheliceral bas es visible, setae absent. Chilum absent. Eyes six, well developed, ALE-ALE and PLE-PME almost touching. All eyes on dark pigment, ALE largest, circular, PME circular, PLE oval; posterior eye row procurved from front; ALE-PLE separated by less than ALE radius, PME touching through out most of their length. Sternum longer than wide, pale orange, uniform, not fused to carapace, surface smooth, without pits, anterior margin unmodified, posterior margin extending posteri orly beyond anterior edges of coxae IV, distance between coxae approximately equal, extensions of pre-coxal triangles present, setae sparse, light, needle-like, evenly scattered, originating from surface. Mouthparts: Chelicerae, endites and la bium pale orange. Chelicerae straight, anterior face unmodified; without teeth on both promar gin and retromargin; fangs directed posteriorly; setae light, needle-like, evenly scattered; paturon inner margin with pairs of enlarged setae. Labium rectangular, not fused to sternum, ante rior margin indented at middle, with many setae on anterior margin. Endites: serrula present in single row, anteromedian tip unmodified. Female palp claw absent; spines absent. Abdomen: ovoid, rounded posteriorly; dorsum soft portions white, without color pattern. Book lung covers large, elliptical, without setae. Posterior spiracles con nected by groove. Pedicel tube short, unmodified, scutum extending far dorsal of pedicel. Dorsal scutum weakly sclerotized, pale orange, without color pattern, covering less than 1/2 of abdo men, more than 1/2 to most of abdomen width, not fused to epigastric scutum, surface smooth. Epigastric scutum weakly sclerotized, not sur rounding pedicel, without lateral joints and without ridges. Postepigastric scutum weakly sclerotized, pale orange, short, only around epi gastric furrow, not fused to epigastric scutum, anterior margin amply procurved. Spinneret scutum absent. Abdominal setae light, needle-like. Basal article of the ALS entire (not crossed by a diagonal membranous area). Colulus repre sented only by setae. Legs: pale orange, without color pattern; patella plus tibia I near as long as carapace. Leg spination (only surfaces bearing spines listed): leg I: femur: pv0-0-1-1-1-1, rv0-0-1-1-1-0, tibia: V2-2-2-2-0, metatarsus: V2-2 (the retrolateral row slightly displaced to distal), leg II: femur: pv0-0-0-1-1-0, rv1-1-1-0-1-0, tibia: V2-2-2-2-0, metatarsus: V2-2 (the retrolateral row slightly displaced to distal), leg III: tibia: dp1-1/0, dr1-0, pv0-0-1, rv0-0-1, metatarsus: d2-2/0, leg IV: patella: d2, tibia: d1(very small)/0-0-0, dp1-1, dr1-1, pv0-0-1, rv0-0-1, metatarsus: dp1-0-1, dr1-0-0, v0-0-2. Tarsi I to IV without inferior claw. Genitalia: epigastric region semicircular, widely open (Figs. 5G, 6F): anterior receptaculum very narrow, posteriorly arising from a thin semicir cular transverse sclerite, and barely surpassing the anterior margin of the epigastric furrow; two lateral, relatively short apodemes reaching to the level of the posterior border of the postepigastric scutum; there is no evidence of any additional re ceptacle (Fig. 6F).

Male unknown.

Type material. Female holotype from India: Tamil Nadu: Kodaikanal, native cloud forest by falls. 10° 15´N, 77° 31´E. 2 January 1990. V. & B. Roth colls. (CAS TYPE 20372, voucher CJG-1965).

Etymology. The specific epithet is a patronymic in honor of Vincent and Barbara Roth, collectors of the holotype.

Distribution. Only known from the type local ity.