A review of the Indian species formerly assigned to the genus Storena Walckenaer, 1805 (Araneae: Zodariidae) with the description of a new genus

Indian spider species currently assigned to Storena Walckenaer, 1805 are revised mostly based on the type material available in the National Zoological Collection, Zoological Survey of India, Kolkata. A new genus, Laminion gen. nov. is proposed to include four species; three are transferred from St orena: Laminion arakuensis (Patel & Reddy, 1989) gen. et comb. nov., Laminion birenifer (Gravely, 1921) gen. et comb. nov. and Laminion debasrae (Biswas & Biswas, 1992) gen. et comb. nov., whereas the fourth species is from Suffasia Jocqué, 1991: Laminion gujaratensis (Tikader & Patel, 1975) gen. et comb. nov. The species Storena tikaderi Patel & Reddy, 1989 syn. nov. is synonymised with L. birenifer gen. et comb. nov. Storena dibangensis Biswas & Biswas, 2006 and Storena indica Tikader & Patel, 1975 are transferred to Mallinella Strand, 1906. All the type material examined are imaged and redescribed. In addition, images of the type material of Storenomorpha joyaus (Tikader, 1970) are presented.


Introduction
The zodariid genus Storena Walckenaer, 1805 currently consists of 61 nominal species distributed across Africa, Australia, Australasia, Neotropical and Southeast Asia (World Spider Catalog 2020). The genus is particularly rich in Australia with more than half of the described species (41 species) and is relatively well revised (Jocqué & Baehr 1992; World Spider Catalog 2020). Species described from other parts of the globe are poorly known and currently the African, Australasian and one Southeast Asian representative (probably other species also) are considered to be misidentifi ed (Bosmans & Van Hove 1986;Dankittipakul et al. 2012;World Spider Catalog 2020). Storena is most probably an endemic Australian genus (Jocqué & Baehr 1992;Baehr & Jocqué 1994) and representatives of this genus from other countries need to be revised as well. The genus currently has six representatives in India: Storena arakuensis Patel & Reddy, 1989, Storena birenifer Gravely, 1921, Storena debasrae Biswas & Biswas, 1992, Storena dibangensis Biswas & Biswas, 2006, Storena indica Tikader & Patel, 1975 and Storena tikaderi Patel & Reddy, 1989(Caleb & Sankaran 2020World Spider Catalog 2020). One species, Storena gujaratensis Tikader & Patel, 1975 has already been transferred to Suffasia Jocqué, 1991(Solanki et al. 2018). All the Indian species of Storena are known only from their original descriptions, which lack clear diagnostic illustrations (see Gravely 1921;Tikader & Patel 1975;Patel & Reddy 1989;Biswas & Biswas 1992, 2006, making their identifi cation very diffi cult. Because of inadequate descriptions, Dankitt ipakul et al. (2012) considered all Indian Storena species as incertae sedis. Therefore, we revise the Indian species of Storena based on the type material available in the National Zoological Collection, Zoological Survey of India, Kolkata.

Material and methods
The specimens were studied under a Leica EZ4 HD stereo microscope. All measurements are in millimetres (mm). Lengths of pedipalp/palp and leg segments are given as: total (femur, patella, tibia, metatarsus (except for pedipalp/palp), tarsus). The micrographic images were taken with a Leica DFC500 digital camera attached to a Leica M205A stereo microscope with the software package Leica Application Suite (LAS, ver. 3.8) for stacking images taken at different focal planes.
CHELICERAE. Covered with fi ne hairs dorsally, with disto-mesal group of hairs; cheliceral margins without teeth, but with denticles; lateral condyles prominent, moderately elevated. Fangs short and stout.
LEGS. Covered with fi ne hairs, with elongated metatarsi and tarsi, femora and patellae with few spines, tibiae and metatarsi with numerous spines, mostly on posterior ones, metatarsi bear brush of distoventral setae distinctly denser on posterior ones, tarsi lined with paired ventral spinules; leg formula 4132.

Justifi cation of the synonymy
The species S. tikaderi was described on the basis of male and female specimens collected in Andhra Pradesh (Patel & Reddy 1989). Altho ugh we did not examine the types of this species, the habitus and the female genitalia illustrated for this species show similarities with that of L. birenifer gen. et comb. nov., including paired, reniform markings on anterior opisthosoma and the shape of median and lateral plates of epigynum (compare Figs 1B, 3D with Patel & Reddy 1989: fi g. 2a, c). Moreover, the type localities of both species are only about 320 km apart along the eastern Coast, which refl ects a wide distributional range of L. birenifer gen. et comb. nov. Based on these observations, we propose to consider S. tikaderi as a junior synonym of L. birenifer gen. et comb. nov.

Remarks
The ZSI collection has two glass tubes for S. birenifer. First tube labeled as "Types" (2914/H2) contains one male and one female specimens in good condition, with intact genitalia. Second tube labeled as "cotype" (2915/H2) contains one male specimen in good condition. We were unable to trace out the types of S. tikaderi in the arachnid collection of ZSI, even though the authors mentioned that the types would be deposited here (Patel & Reddy 1989). (Patel & Reddy, 1989) gen. et comb. nov. Fig. 9 Storena arakuensis Patel & Reddy, 1989: 223, fi g. 1a-h.

Male
Unknown.

Justifi cation of the transfer
Patel & Reddy (1989) described this species on the basis of male and female specimens collected in Andhra Pradesh. Although we did not examine the types of this species, the female genitalia illustrated for this species clearly show the presence of sclerotised lateral and median epigynal plates (Patel & Reddy 1989: fi g. 1c), which allows the inclusion of this species in Laminion gen. nov.

Remarks
We were unable to trace out the types of S. arakuensis in the arachnid collection of ZSI, even though the authors mentioned that the types would be deposited here (Patel & Reddy 1989). (Biswas & Biswas, 1992 )

Male
Unknown.

Remarks
The ZSI collection has one glass tube for this species, which is labeled as "holotype" (no register number) containing one female specimen in fairly good condition, with broken legs and chelicerae and almost detached opisthosoma. The same tube has a small glass vial containing the dissected genitalia. The label mentions the year of collection as "1987".

Remarks
The ZSI collection has one glass tube for this species, which is labeled as "holotype" (no register number) containing one male specimen in good condition. Its left pedipalp found removed and kept in the same tube.

Male
Unknown.

Justifi cation of the transfer
This species was described on the basis of one female specimen collected in Arunachal Pradesh. Although we did not examine the type of this species, the internal female genitalia illustrated for this species (Biswas & Biswas 2006: fi g. 3) clearly matches with that of Mallinella Strand, 1906 in its general appearance. Based on this observation, we propose to transfer S. dibangensis to Mallinella. The vulva illustrated for this species suggests a possible synonymy of this species with Mallinella redimita (Simon, 1905); however, confi rmation requires the re-examination of type or topotype materials of M. dibangensis comb. nov.

Remarks
We were unable to trace out the type of S. dibangensis in the arachnid collection of ZSI. It may either be lost or misplaced elsewhere in the collection. (Tikader & Patel, 1975) comb. nov.
Measurements of paratype female (Fig. 10B): body length 7.95. Carapace length 4.17, width 2.60. Opisthosoma length 3.78, width 2.80. Genitalia as in Fig. 11A-B. Justifi cation of the transfer Tikader & Patel (1975) described this species on the basis of male and female specimens collected in Gujarat. Detailed study of the types of this species indicates that it has diagnostic features of Mallinella   Fig. 11. Mallinella indica (Tikader & Patel, 1975) comb. nov., ♂♀, paratype and allotype of Storena indica Tikader & Patel, 1975 as illustrated in Dankittipakul et al. (2012). Based on this observation, we propose to transfer S. indica to Mallinella.

Remarks
The ZSI collection has three glass tubes for this species. First tube labeled as "holotype" (5354/18) contains one female specimen in good condition, with intact genitalia. Second tube labeled as "paratype" (5355/18) contains one female specimen in good condition, with detached opisthosoma. Third tube labeled as "allotype" (5356/18) contains one male specimen in good condition. The same tube contains another label mentioning that the collecting locality of holotype, paratype and allotype as Pavagadh with register number 5354/18.

Note
We examined the holotype, paratypes and allotype of S. joyaus (Fig. 12A-E) and fo und that these were all subadults, thereby, the morphology of its female genitalia remains yet unknown. Jocqué & Bosmans (1989) des cribed the male of this species based on non-type material deposited in BMNH, London, that were collected in Sikkim. Without referring this, Sen et al. (2015) claimed that they had provided the fi rst description of the male of S. joyaus. It is surprising that, even though they had two female specimens at hand, they never attempted to illustrate its genitalia, instead illustrated the male genitalia that was already well illustrated (Jocqué & Bosmans 1989: fi gs 11-12).

Remarks
The ZSI collection has three glass tubes for this species. First tube labeled as "holotype" (no register number) contains one subadult female specimen in good condition, with intact genitalia. Second tube labeled as "paratype" (3232/18) contains four subadult female and one juvenile specimens in good condition. Third tube labeled as "allotype" (3231/18) contains one subadult male specimen in good condition.  (Tikader, 1970), ♂♀, holotype and allotype of Homalonychus joyaus Tikader, 1970

Discussion
The zodariid spiders in India are currently represented by 11 genera and 31 species (Caleb & Sankaran 2020). Like many other poorly represented spider families known in India, the zodariid spiders are also poorly known compared to the intensely studied fauna of Southeast Asia (e.g., see Dankittipakul et al. 2012). Species described from India are mostly based on descriptions that are poor and insuffi cient, which make their proper identifi cation often impossible. Most often, the majority of these species are misplaced and misidentifi ed (e.g., see Sankaran et al. 2019). All these demand revisions of known Indian zodariid species in order to facilitate their identifi cation and help to aptly classify unknown / undescribed taxa. In the present work, we transferred all Indian species of Storena into the right genera including a new genus, as Storena occurs only in Australia. It not only resolves the taxonomic dilemma (in part) existing in Indian Zodariidae, but the description of the new genus highlights the need for extensive surveys to get a clear picture of the actual diversity of this group in the country.