Hippasa Simon, 1885

SANKARAN, PRADEEP M. & CALEB, JOHN T. D., 2023, Notes on Indian wolf spiders: II. Genus Hippasa Simon, 1885 (Araneae: Lycosidae Hippasinae), Zootaxa 5230 (2), pp. 101-152 : 102-104

Hippasa Simon, 1885

Diagnosis

[Modified after Alderweireldt & Jocqué (2005) and Wang et al. (2015)]. The genus Hippasa can be distinguished from other lycosid genera by the following combination of features: slender body with chevron-like markings on the dorsum of the opisthosoma ( Figs 2A, C , 3B , 4A , 5A , 6A , 10A–B , 12A , 14A , 17A , 19A , 22A , 24A, G , 27A, E , 33A , 34A ), sternum usually with a mid-longitudinal black band ( Figs 2D , 3D–E , 5E , 6E , 17B , 19B , 20B , 27B, F , 30C ), bi-segmented posterior spinnerets with elongated basal segments ( Figs 2B , 3B , 4A , 5F , 6F , 19D , 20D , 24D, J , 27D, H , 30E ), male pedipalp with hooked synembolus ( Figs 7D , 8C , 19H , 21C , 25D , 26C , 28D , 29C ), bifid tegular apophysis ( Figs 8A , 21A–B , 26A , 29A ), and slender embolus ( Figs 7D , 8C , 19H , 21C , 25D , 26C , 28D , 29C ), epigyne thickly covered with bushy setae ( Figs 7E , 14C , 17D , 20E , 25E , 28E , 30F ), epigyne composed of a median and paired lateral plates, with or without posterior scape or median atrium leading to internal hood ( Figs 7F–G , 8D–E , 11D–E , 13C–D , 14C , 15A–B , 17E , 18A–B , 20F–G , 21D–F , 25F , 26D–E , 28F , 29D–E ), and vulva with or without accessory glands, and slender and slightly to strongly curved spermathecal stalks bearing small spermathecal heads ( Figs 8E , 14D , 18B , 21F , 26E , 29E , 31B ). Moreover, Hippasa species can be identified in the field by their behaviour of spinning funnel-like webs ( Fig. 1E–H ).

Type species. Hippasa agelenoides ( Simon, 1884) , by subsequent designation ( Simon 1885).

Remarks. The type species of Hippasa , H. agelenoides ( Simon 1898; Alderweireldt & Jocqué 2005; World Spider Catalog 2022) was originally described under Pirata Sundevall, 1833 based on five immature female syntypes collected in Minhla, southern-central Myanmar ( Simon 1884). The details of the type material of H. agelenoides provided by Alderweireldt & Jocqué (2005) seem to be wrong. According to them, the types of H. agelenoides in the MNHN collection consist of one female and two subadult males that were collected from the North Guzerath in Myanmar / Burma. The locality ‘North Guzerath’ is not in Myanmar, but it indicates the northern part of the Indian State of Gujarat (see also Simon 1897). It is worth mentioning that even though Alderweireldt & Jocqué (2005) examined an adult female of H. agelenoides from Simon’s collection, they never illustrate its genitalia, and the epigyne of the type of this species is known only from textual description. Of the five immature syntypes, three are currently deposited in MSNG, Genova (M. Tavano, pers. comm., Fig. 3A–G ), while the remaining two are deposited in MNHN, Paris (E. Leguin, pers. comm., Fig. 2A–E ) (see also Tikader & Malhotra 1980). While erecting Hippasa, Simon (1885) transferred Lycosa greenalliae Blackwall, 1867 to Hippasa , which was also described based on two immature specimens ( Blackwall 1867), and put it as its generotype. In his paper, Simon (1885) mentioned the name H. greenalliae in block letters, which indicates that this name is valid and has priority over P. agelenoides , which was presented in small letters under the former one. Similar way of presentation of synonymies can also be seen in the case of other species mentioned in the same paper (for example, see Palpimanus gibbulus Dufour, 1820 ). However, in a later publication ( Simon 1898), he mentioned that H. agelenoides was the type species of Hippasa , indicating that Simon recognised H. agelenoides to be valid and thus separated it from the synonymy of H. greenalliae . He also mentioned that H. agelenoides is widespread in India, Indochina and Malaysia.

Though Simon (1885) redescribed H. greenalliae , it is apparent that his redescription was based not on topotype material, but instead on specimens collected in Kollegal (=Collegall, formerly part of Coimbatore District of Tamil Nadu State; presently part of Chamarajanagara District of Karnataka State, southern India) and Ramanathapuram/ Ramnad in the southern Indian State of Tamil Nadu. These collecting localities are more than 2,200 Km away from the type locality of H. greenalliae , which is located somewhere in northern India (either in Agra, Meerut, or New Delhi / Delhi — Blackwall 1867; Thorell 1887). Tikader & Malhotra (1980) erroneously located the type locality of H. greenalliae in Tamil Nadu, which was the collecting locality of specimens described by Simon (1885). Since the types of H. greenalliae are immatures ( Blackwall 1867), Simon’s identification of H. greenalliae and the Tikader & Malhotra’s (1980) redescription based on Simon’s material may be wrong. The illustrations done by the latter authors seem similar to that of H. pantherina (compare Tikader & Malhotra 1980: figs 73–74 to Figs 30F , 31D–E ), which was synonymised with H. greenalliae by the same authors. This is also applicable to the subsequent redescriptions of H. greenalliae , which were probably based on the description and illustrations of Simon (1885) and Tikader & Malhotra (1980) ( Karsch 1892; Barrion & Litsinger 1994; Biswas & Raychaudhuri 2007; Sen et al. 2015; Dhali et al. 2017; Caleb 2020). Thus, the species redescribed under the name ‘ H. greenalliae ’ is in fact H. pantherina and the identity of H. greenalliae is questionable due to the immature stage of the syntypes. It may be considered as nomen dubium until adult specimens from the type locality can be examined. As a result, the synonymy of H. pantherina with H. greenalliae as proposed by Tikader & Malhotra (1980) is no longer acceptable and the former is herein revalidated. This in turn challenges the validity of the ‘ H. greenalliae ’ species-group proposed by Wang et al. (2015), which should be reconsidered as the H. pantherina species-group.

Thorell (1887) provided textual description of a male specimen from the type locality of P. agelenoides and based on this, he transferred P. agelenoides to Diapontia Keyserling, 1877 . This decision reversed a few years later, when Simon (1898: 326) transferred the species back to Hippasa . Even though Thorell (1887) was not sure about the conspecificity between H. agelenoides and H. greenalliae , he considered that both H. agelenoides and ‘ H. greenalliae ’ described by Simon (1885) from Tamil Nadu were identical with D. agelenoides . Even though the label found inside the type vial of P. agelenoides deposited in MSNG, and which was probably written by Thorell (1887), mentioned that ‘ P. agelenoides is equal to H. greenalliae ’, Thorell’s interpretation regarding the conspecificity of both ‘ H. greenalliae ’ described by Simon (1885) from Tamil Nadu and H. agelenoides is wrong due to the uncertain identity of the former species.

Of the 10 valid Hippasa species found in India (see below), H. agelenoides , H. madraspatana and H. valiveruensis are smaller in body size (TL <9), whereas other species are larger (TL> 10). The smaller species have less developed tarsal scopulae that is well-developed in larger species.

Distribution. Africa, Asia ( World Spider Catalog 2022).

Key to Indian species of Hippasa

Species considered as nomina dubia and nomen nudum are excluded from the key. Males of H. himalayensis , H. loundesi and H. valiveruensis are also excluded as they are unknown. Details of the males of H. deserticola and H. holmerae were taken from Wang et al. (2015) and Marusik & Nadolny (2021), and that of the female of H. olivacea from Tikader & Malhotra (1980).

1a Males...............................................................................................2

1b Females............................................................................................. 7

2a Body less than 9 mm long.............................................................................. 3

2b Body more than 10 mm long.............................................................................4

3a Retrolateral lamellate process of tegulum well-developed, anterior arm of tegular apophysis short and broad towards distally, embolus lacks ventral membranous sheath-like extension ( Fig. 8A, C )................................ H. agelenoides

3b Retrolateral lamellate process of tegulum less developed, anterior arm of tegular apophysis long and in same width along entire length, embolus with ventral membranous sheath-like extension ( Fig. 26A, C )........................ H. madraspatana

4a Anterior and mesal arms of tegular apohysis closely spaced.................................................... 5

4b Anterior and mesal arms of tegular apohysis widely spaced.................................................... 6

5a Tegular apophysis with strongly curved anterior and broad mesal arms, synembolus less curved, embolus less curved with widened tip ( Marusik & Nadolny 2021: fig. 2A, E–H).............................................. H. deserticola

5b Tegular apophysis with straight anterior and narrow mesal arms, synembolus strongly curved, strongly curved embolus without widened tip ( Wang et al. 2015: fig. 3A)........................................................... H. holmerae

6a Anterior arm of tegular apophysis short, conductor small, synembolus strongly curved, embolus with smooth distal curvature ( Fig. 21A, C )................................................................................. H. lycosina

6b Anterior arm of tegular apophysis long, conductor large, synembolus less strongly curved, embolus with strong distal curvature ( Fig. 29A, C )............................................................................... H. pantherina

7a Body less than 9 mm long...............................................................................8

7b Body more than 10 mm long........................................................................... 10

8a Median plate of epigyne with atrium leading to internal hood ( Fig. 8D–E )............................. H. agelenoides

8b Median plate of epigyne without atrium................................................................... 9

9a Epigyne with broadly triangular median plate, accessory glands with stalks, spermathecae oval ( Fig. 26D–E ).................................................................................................. .. H. madraspatana

9b Epigyne with narrow, M-shaped median plate, accessory glands without stalks, spermathecae globular ( Fig. 31A–B )............................................................................................. .. H. valiveruensis

10a Median plate of epigyne with atrium leading to internal hood................................................. 11

10b Median plate of epigyne without atrium.................................................................. 13

11a Median plate of epigyne posteriorly with a median scape ( Fig. 15A )................................... .. H. holmerae

11b Median plate of epigyne without scape................................................................... 12

12a Median plate of epigyne well-developed, accessory glands present, spermathecae globular ( Fig. 18A–B )........ H. loundesi

12b Median plate of epigyne poorly developed, accessory glands absent, spermathecae elongate-ovoid ( Fig. 29D–E ).................................................................................................... H. pantherina

13a Median plate of epigyne with small atrium and a posteromedian scape ( Figs 11D–E , 13C–D )............... H. deserticola

13b Median plate of epigyne with large atrium and without posteromedian scape......................................14

14a Atrium of epigynal median plate transversely elliptical ( Tikader & Malhotra 1980: fig. 84)................... H. olivacea

14b Atrium of epigynal median plate triangular.................................................................15

15a Accessory glands with stalks, spermathecae peanut-shaped ( Fig. 14D )............................... H. himalayensis

15b Accessory glands without stalks, spermathecae globular ( Fig. 21F )...................................... H. lycosina

Figures

FIGURE 1. Field photographs of Hippasa spp. and their webs. A–B, F Hippasa agelenoides (Simon, 1884): A male; B female; F web. C–D, H Hippasa madraspatana Gravely, 1924: C male; D female; H web. E Hippasa pantherina Pocock, 1899, female. G web of Hippasa lycosina Pocock, 1900. Figures not to scale. Photo credits: A, F–G Pradeep M. Sankaran, B–D, H John T. D. Caleb, E, Jimmy Paul.

FIGURE 2. Hippasa agelenoides (Simon, 1884), syntype females.A–B female 1: A habitus, dorsal view; B same, ventral view. C–D female 2: C habitus, dorsal view; D same, ventral view. E labels from syntype bottle. Scale bars: A–B, 0.5 mm; C–D, 1 mm. © MNHN, Paris.

FIGURE 3. Hippasa agelenoides (Simon, 1884), syntype females.A, D female 1: A habitus, dorsal view; D same, ventral view. B, E female 2: B habitus, dorsal view; E same, ventral view. C, F female 3: C habitus, dorsal view; F same, ventral view. G labels from syntype bottle. Figures not to scale. © MSNG, Genova.

FIGURE 4. Hippasa agelenoides (Simon, 1884) male, labeled as Diapontia agelenoides. A habitus, dorsal view. B same, dorsoretrolateral view. C–D left pedipalp of the same: C ventral view; D retrolateral view. E labels from the bottle. Figures not to scale. © MSNG, Genova.

FIGURE 5. Hippasa agelenoides (Simon, 1884), male (ADSH595032). A habitus, dorsal view. B same, ventral view. C same, retrolateral view. D same, frontal view. E prosoma showing endites, labium, sternum, ventral view. F spinnerets, ventral view. G–H left chelicera: G ventroretrolateral view showing teeth and stridulatory file; H prolateral view showing stridulatory file. I left tarsus IV showing reduced scopula, retrolateral view. Scale bars: A–C, 2 mm; D–E, I, 1 mm; F, 0.5 mm; G–H, 0.2 mm.

FIGURE 6. Hippasa agelenoides (Simon, 1884), female (ADSH595032).A habitus, dorsal view. B same, ventral view. C same, retrolateral view. D same, frontal view. E prosoma showing endites, labium, sternum, ventral view. F spinnerets, ventral view. G–H left chelicera: G ventral view showing teeth; H retrolateral view showing stridulatory file. I left tarsus IV showing reduced scopula, prolateral view. Scale bars: A–C, 2 mm; D–E, 1 mm; F–I, 0.5 mm.

FIGURE 7. Hippasa agelenoides (Simon, 1884), male and female genitalia (ADSH595032). A–D male left pedipalp: A prolateral view; B ventral view; C retrolateral view; D embolic division, ventral view. E–G female genitalia: E epigyne intact, ventral view; F same, after clearing in KOH and removing bushy setae, ventral view; G vulva, dorsal view. Scale bars: A–G, 0.2 mm.

FIGURE 8. Hippasa agelenoides (Simon, 1884), male and female genitalia (ADSH595032). A–C male left pedipalp: A ventral view; B retrolateral view; C embolic division, ventral view. D–E female genitalia: D epigyne ventral view; E vulva, dorsal view. Abbreviations:AGaccessory gland;AT anteriorarmof tegularapophysis; Cconductor;E embolus; EBembolic base; FDfertilization duct; H hood; LEP lateral epigynal plate; LT lamellate process of tegulum; MEP median epigynal plate; MT mesal arm of tegular apophysis; Pa palea; RTP retrolateral process of tegulum; S spermatheca; Se synembolus; SD sperm duct; SS spermathecal stalk; ST subtegulum; T tegulum; TA tegular apophysis; TG tegular groove; TS tegular stalk. Scale bars: A–E, 0.2 mm.

FIGURE 10. Hippasa deserticola Simon, 1889, types of Hippasa charamaensis Gajbe, 2004.A holotype female habitus, dorsal view. B allotype male habitus, dorsal view. C labels from type bottle. Scale bars: A–B, 2 mm.

FIGURE 11. Hippasa deserticola Simon, 1889, genitalia of types of Hippasa charamaensis Gajbe, 2004.A–C left pedipalp of allotype male: A prolateroventral view; B ventral view; C retrolateral view. D–G holotype female genitalia: D epigyne, ventral view; E same, oblique ventral view; F vulva, dorsal view; G same, oblique dorsal view. Scale bars: A–C, 0.5 mm; D–G, 0.2 mm.

FIGURE 12. Hippasa deserticola Simon, 1889, types of Hippasa madhuae Tikader & Malhotra, 1980. A holotype female habitus (NZC-ZSI 4665/18), dorsal view. B allotype male habitus (NZC-ZSI 4666/18), dorsal view. C–D labels from type bottle. Scale bars: A–B, 2 mm.

FIGURE 13. Hippasa deserticola Simon, 1889, genitalia of types of Hippasa madhuae Tikader & Malhotra, 1980 (NZC-ZSI 4665/18 and 4666/18). A–B left pedipalp of allotype male: A prolateroventral view; B ventral view. C–D holotype female genitalia: C epigyne, oblique ventral view; D vulva, oblique dorsal view. Scale bars: A–B, 0.5 mm; C–D, 0.2 mm.

FIGURE 14. Hippasa himalayensis Gravely, 1924, lectotype female (NZC-ZSI 2268/18). A habitus, dorsal view. B same, ventral view. C epigyne, ventral view. D vulva, dorsal view. E labels from type bottle. Scale bars: A–B, 2 mm; C, 0.2 mm; D, 0.5 mm.

FIGURE 15. Hippasa holmerae Thorell, 1895, genitalia of a topotype female specimen deposited in ZSI (NZC-ZSI 2393/17). A epigyne, ventral view. B vulva, dorsal view. C labels from the bottle. Scale bars: A–B, 0.2 mm.

FIGURE 17. Hippasa loundesi Gravely, 1924, female (ADSH5950252). A habitus, dorsal view. B prosoma showing sternum, ventral view. C left chelicera, ventral view. D epigyne intact, ventral view; E same, after clearing in KOH and removing bushy setae, ventral view; F vulva, dorsal view. Scale bars: A–B, 2 mm; C, 1 mm; D, 0.2 mm; E–F, 0.5 mm.

FIGURE 18. Hippasa loundesi Gravely, 1924, genitalia of female (ADSH5950252). A epigyne, ventral view. B vulva, dorsal view. Abbreviations: AG accessory gland; FD fertilization duct; LEP lateral epigynal plate; MEP median epigynal plate; S spermatheca; SS spermathecal stalk. Scale bars: A–B, 0.5 mm.

FIGURE 19. Hippasa lycosina Pocock, 1900, male (ADSH595026). A habitus, dorsal view. B prosoma showing sternum, ventral view. C left chelicera, ventral view. D spinnerets, ventral view. E–H left pedipalp: E prolateral; F ventral; G retrolateral; H embolic division, ventral view. Scale bars: A–B, 2 mm; C, E–G, 0.5 mm; D, 1 mm; H, 0.2 mm.

FIGURE 20. Hippasa lycosina Pocock, 1900, female (ADSH595026). A habitus, dorsoretrolateral view. B prosoma showing sternum, ventral view. C left chelicera, ventral view. D spinnerets, ventral view. E intact epigyne, ventral view; F same, after clearing in KOH and removing bushy setae, ventral view; G vulva, dorsal view. Scale bars: A–B, 2 mm; C–D, 1 mm; E–G, 0.2 mm.

FIGURE 21. Hippasa lycosina Pocock, 1900, male and female genitalia (ADSH595026). A–C male left pedipalp: A ventral view; B retrolateral view; C embolic division, ventral view. D–E female genitalia: D epigyne, ventral view; E vulva, dorsal view; F same without left spermatheca showing accessory gland, dorsal view. Abbreviations: AG accessory gland; AT anterior arm of tegular apophysis; C conductor; E embolus; EB embolic base; FD fertilization duct; H hood; LEP lateral epigynal plate; MEP median epigynal plate; MT mesal arm of tegular apophysis; Pa palea; RTP retrolateral process of tegulum; S spermatheca; Se synembolus; SD sperm duct; SS spermathecal stalk; ST subtegulum; T tegulum; TA tegular apophysis. Scale bars: A–B, 0.5 mm; C–F, 0.2 mm.

FIGURE 22. Hippasa lycosina Pocock, 1900, allotype male of Hippasa mahabaleshwarensis Tikader & Malhotra, 1980 (NZC- ZSI 4651/18). A habitus, dorsal view. B–C left pedipalp: B ventral view; C retrolateral view. D label from the allotype tube. Scale bars: A, 2 mm; B–C, 1 mm.

FIGURE 24. Hippasa madraspatana Gravely, 1924, male (A–F) and female (G–L) (ADSH5950281). A, G habitus, dorsal view. B, H same, frontal view. C, I prosoma showing endites, labium, sternum, ventral view. D, J spinnerets, ventral view. E, K left chelicera, retrolateral view. F, L left tarsus IV showing reduced scopula, lateral view. Scale bars: A, G, 2 mm; B–C, F, H–I, 1 mm; D–E, J, L, 0.5 mm; K, 0.2 mm.

FIGURE 25. Hippasa madraspatana Gravely, 1924, male and female genitalia (ADSH5950281). A–C male left pedipalp, D male right pedipalp: A prolateral view; B ventral view; C retrolateral view; D embolic division, ventral view. E–G female genitalia: E intact epigyne, ventral view; F same, after clearing in KOH and removing bushy setae, ventral view; G vulva, dorsal view. Scale bars: A–C, 0.5 mm; D–G, 0.2 mm.

FIGURE 26. Hippasa madraspatana Gravely, 1924, male and female genitalia (ADSH5950281). A–B male left pedipalp, C male right pedipalp: A ventral view; B retrolateral view; C embolic division, ventral view. D–E female genitalia: D epigyne, ventral view; E vulva, dorsal view. Abbreviations: AG accessory gland; AT anterior arm of tegular apophysis; C conductor; E embolus; EB embolic base; FD fertilization duct; LEP lateral epigynal plate; LT lamellate process of tegulum; MEP median epigynal plate; MS membranous sheath; MT mesal arm of tegular apophysis; Pa palea; RTP retrolateral process of tegulum; S spermatheca; Se synembolus; SD sperm duct; SS spermathecal stalk; ST subtegulum; T tegulum; TA tegular apophysis; TG tegular groove; TS tegular stalk. Scale bars: A–B, 0.5 mm; C–E, 0.2 mm.

FIGURE 27. Hippasa pantherina Pocock, 1899, male (A–D) (ADSH59502) and female (E–H) (ADSH59503). A, E habitus, dorsal view. B, F prosoma showing endites, labium, sternum, ventral view. C, G left chelicera, ventral view. D, H spinnerets, ventral view. Scale bars: A–B, D–F, H, 2 mm; C, G, 1 mm.

FIGURE 28. Hippasa pantherina Pocock, 1899, male (ADSH59502) and female (ADSH59503) genitalia. A–D male left pedipalp: A prolateral view; B ventral view; C retrolateral view; D embolic division, ventral view. E–G female genitalia: E intact epigyne, ventral view; F same, after clearing in KOH and removing bushy setae, ventral view; G vulva, dorsal view. Scale bars: A–C, E–G, 0.5 mm; D, 0.2 mm.

FIGURE 29. Hippasa pantherina Pocock, 1899, male (ADSH59502) and female (ADSH59503) genitalia. A–C male left pedipalp: A ventral view; B retrolateral view; C embolic division, ventral view. D–E female genitalia: D epigyne, ventral view; E vulva, dorsal view. Abbreviations: AT anterior arm of tegular apophysis; C conductor; E embolus; EB embolic base; FD fertilization duct; LEP lateral epigynal plate; LT lamellate process of tegulum; MEP median epigynal plate; MT mesal arm of tegular apophysis; Pa palea; RTP retrolateral process of tegulum; S spermatheca; Se synembolus; SD sperm duct; SS spermathecal stalk; ST subtegulum; T tegulum; TA tegular apophysis; TG tegular groove. Scale bars: A–B, D–E, 0.5 mm; C, 0.2 mm.

FIGURE 30. Hippasa valiveruensis Patel & Reddy, 1993, female (ADSH595035). A habitus, dorsal view. B prosoma, dorsal view. C prosoma showing sternum, ventral view. D left chelicera, ventroretrolateral view. E spinnerets, ventral view. F intact epigyne, ventral view; G same, after clearing in KOH and removing bushy setae, ventral view; H vulva, dorsal view. Scale bars: A, 2 mm; B, E, 0.5 mm; C, 1 mm; D, F–H, 0.2 mm.

FIGURE 31. Hippasa valiveruensis Patel & Reddy, 1993, genitalia of female (ADSH595035). A epigyne, ventral view. B vulva, dorsal view.Abbreviations:AG accessory gland; FD fertilization duct; LEP lateral epigynal plate; MEP median epigynal plate; S spermatheca; SS spermathecal stalk. Scale bars: A–B, 0.2 mm.

FIGURE 33. Hippasa hansae Gajbe & Gajbe, 1999, holotype female (NZC-ZSI). A habitus, dorsal view. B same, ventral view. C tissue from the epigynal area, dorsal view. D label from type bottle. Scale bars: A–B, 2 mm; C, 0.5 mm.

FIGURE 34. Hippasa wigglesworthi Gajbe & Gajbe, 1999, holotype female (NZC-ZSI).A habitus, dorsal view. B opisthosoma, ventral view. C piece of tegument over the epigynal area, dorsal view. D label from type bottle. Scale bars: A, 2 mm; B–C, 1 mm.