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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by J.-X. Zhang: 24 Dec. 2022; published: 23 Jan. 2023 391
Zootaxa 5230 (3): 391–400
https://www.mapress.com/zt/
Copyright © 2023 Magnolia Press Article
https://doi.org/10.11646/zootaxa.5230.3.8
http://zoobank.org/urn:lsid:zoobank.org:pub:FAA6E8C0-39FD-4EF3-8B4F-717E52950DB6
Deciphering mysteries: On the identity of five enigmatic jumping spiders from
Northeast India, China and Philippines (Araneae, Salticidae)
JOHN T.D. CALEB1,2
1Department of Anatomy, Saveetha Medical College & Hospital, Saveetha Institute of Medical and Technical Sciences, Chennai
602503, Tamil Nadu, India.
2Zoological Survey of India, Prani Vigyan Bhawan, New Alipore, Kolkata—700053, West Bengal, India.
�
caleb87woodgate@gmail.com; https://orcid.org/0000-0002-9471-9467
Abstract
The type specimens of salticid species from Northeast India described by B. K. Biswas and K. Biswas kept in the National
Zoological Collections, Kolkata were studied. Four new synonyms are recognized: Marpissa mizoramensis Biswas &
Biswas, 2007 syn. nov. and Evarcha optabilis (Fox, 1937) syn. nov. with Evarcha flavocincta (C.L. Koch, 1846), and
Phidippus tirapensis Biswas & Biswas, 2006 syn. nov. and Hyllus maskaranus Barrion & Litsinger, 1995 syn. nov. with
Hyllus diardi (Walckenaer, 1837). A new combination is proposed: Pancorius manipuriensis (Biswas & Biswas, 2007)
comb. nov. (ex. Marpissa). P. manipuriensis is illustrated for the first time and redescribed based on the type material.
Detailed digital illustrations of the examined type material and distributional map are also provided.
Key words: revision, taxonomy, type specimen, Evarcha, Hyllus, Marpissa, Pancorius
Introduction
The jumping spider diversity in India is currently represented by 296 species and 103 genera (Caleb & Sankaran
2022). The North-eastern region of India is part of the Indo-Burma biodiversity hotspot (Myers et al. 2000) and is
a treasure trove of diversity waiting to be fully explored. A total of 25 salticid species have been described/reported
from the region so far (Caleb 2019), of which 10 species have been added to the regional list in the last few years
(Kananbala et al. 2014; Basumatary et al. 2018; 2020a, b; 2021; Caleb et al. 2018; 2019b; 2020; Caleb & Acharya
2020). Three species from Manipur, Mizoram and Arunachal Pradesh states of India were described between 2004
to 2007 (Biswas & Biswas 2004; 2006; 2007), but their identity remained unverifiable due to very poor descriptions
and sketchy drawings. Like many Indian salticids which were originally misplaced in genera (e.g. Lyssomanes
Hentz, 1845, Phidippus C.L. Koch, 1846, Zygoballus Peckham & Peckham, 1885) not actually occurring in the Old
World (Prószyński 1990; Edwards 2004; Logunov 2004; Kaldari 2010), these three species were also misplaced.
All of Indian Marpissa and Phidippus are apparently misclassified and require revision. In an attempt to clarify the
taxonomic status of Indian salticids, especially those described by B. K. Tikader, G. L. Sadana, B. K. Biswas and K.
Biswas, several papers were published recently (Caleb et al. 2017; 2019a; 2020; 2021; 2022; Sudhin et al. 2020).
This paper is part of a continual effort to revise Indian salticids and deal with fauna from the North-eastern part of
India. Locating these type specimens, in particular those described by B. K. Biswas and K. Biwas from North-east
India have been challenging since they were not placed in the type material section but were inconspicuously shelved
among the general collections along with the survey material of the respective states (Manipur and Mizoram). In
addition, they were poorly labeled, further obscuring them from being easily recognized or found. Yet, the type
material of Phidippus tirapensis Biswas & Biswas, 2006 described from Arunachal Pradesh by Biswas & Biswas
(2006) was not found either in the type material section or the respective survey materials; it may have been
misplaced elsewhere or lost. The present work, therefore, attempts (1) to clarify the taxonomic status of three salticid
species from North-east India, one from China and one from the Philippines, and (2) to provide first illustrations and
redescription for Marpissa manipuriensis Biswas & Biswas, 2004.
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Material and methods
Morphological examination of the specimens was carried out under a Leica EZ4 HD stereomicroscope. Photographs
were taken by a Leica DFC500 HD camera attached to a Leica M205 A stereomicroscope equipped with Leica
Application Suite (LAS) version 3.8. The male left palp was dissected and examined, and the female genitalia were
dissected and cleared in 10% KOH for observation and photography. Distributional map was prepared using the
online mapping software SimpleMappr (Shorthouse 2010). Wherever the exact collection geographical coordinates
were unavailable on labels, approximate coordinates were obtained using Google Maps (https://maps.google.com).
All measurements are in millimeters (mm). The studied materials are kept in the National Zoological Collections,
Zoological Survey of India, Kolkata, India (NZC-ZSI).
Abbreviations used in the text: AER—anterior eye row; ALE—anterior lateral eye; AME—anterior median
eye; D—described; EFL—eye field length; JC—personal collection of John Caleb; PER—posterior eye row;
PLE—posterior lateral eye; PME—posterior median eye; RTA—retrolateral tibial apophysis.
Taxonomy
Evarcha flavocincta (C. L. Koch, 1846)
Figs 1–6, 25
Maevia flavocincta C. L. Koch, 1846: 74, fig. 1330 (D♀).
Evarcha flavocincta Żabka, 1985: 224, figs 187–196 (♂♀); Prószyński & Deeleman-Reinhold, 2010: 165, figs 71–72 (♀); Roy
et al., 2014: 380, figs 1–6 (♀); Dhali, Saha & Raychaudhuri, 2017: 29, figs 38–42, pl. 17 (♀); Prasad, Kumar & Tyagi,
2022: 1021, figs A–D (♀).
Plexippus optabilis Fox, 1937: 16, fig, 7 (D♀); holotype ♀ in the United States National Natural History Museum, Washington,
D.C., examined drawings of the holotype by J. Prószyński (1987); syn. nov.
Marpissa mizoramensis B.K. Biswas & K. Biswas, 2007: 458, figs 1–3 (D♀); holotype ♀ in NZC-ZSI, Kolkata, examined; syn.
nov.
For a complete list of taxonomic references refer the WSC (2022).
Type material. Marpissa mizoramensis Biswas & Biswas, 2007: Holotype ♀ (NZC-ZSI 5480/18) from INDIA,
Mizoram, Lunglei (22.8629°N, 92.7750°E), 962 m a.s.l., 25.03.1994, leg. Bijan Biswas.
Justification of synonymy. Marpissa mizoramensis Biswas & Biswas, 2007 was described from Lunglei,
Mizoram based on a female holotype (Biswas & Biswas 2007). Detailed examination of the holotype revealed
that the general body colour pattern and the sclerotized epigyne with large, comma shaped grooves separated by a
broad median septum; posterior margin with a pair of lateral pockets; complex and coiled internal duct system with
membraneous ducts running initially from the copulatory openings and gradually becoming sclerotized; anteriorly
placed spermathecae are identical with that of Evarcha flavocincta (cf. Figs 1–4 with figs 187–192 in Żabka (1985)).
The name Marpissa mizoramensis is therefore regarded as a junior synonym of E. flavocincta.
Evarcha optabilis (Fox, 1937) was described based on a female holotype collected from Suifu, Szechwan
Province (now Sichuan Province), China (Fox 1937). The illustrations of the genitalia of the holotype (Fox 1937;
Prószyński 1987) depict the epigyne with comma shaped grooves separated by a broad median septum; posterior
margin with a pair of lateral pockets; internal duct system apparently coiling and positioned anteriorly are identical
with that of Evarcha flavocincta (cf. fig. 7 in Fox (1937) and illustration in Prószyński (1987: 26) with figs 188 & 191
in Żcabka (1985) and Fig. 2 herein). It is thus safe to consider E. optabilis as a junior synonym of E. flavocincta.
Distribution. India, China to Indonesia (WSC 2022). India: Mizoram (Biswas & Biswas 2004: sub Marpissa
mizoramensis), Odisha (Prasad et al. 2022), West Bengal (Roy et al. 2014; Dhali, Saha & Raychaudhuri 2017; Tyagi
et al. 2019) (Fig. 25).
TAXONOMIC NOTES ON FIVE SALTICID SPECIES Zootaxa 5230 (3) © 2023 Magnolia Press · 393
FIGURES 1–6. Somatic and genitalia morphology of Marpissa mizoramensis Biswas & Biswas, 2007, holotype. 1—habitus,
dorsal view; 2—epigyne, ventral view; 3—ditto, dorsal view; 4—ditto, anterior view; 5, 6—labels. Scale bars: (1) 1 mm, (2–4)
0.2 mm.
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Hyllus diardi (Walckenaer, 1837)
Figs 7–12, 25
Attus diardi Walckenaer, 1837: 460 (D♀).
Hyllus diardi Simon, 1886: 139; Prószyński, 1984: 62 (♀); Żabka, 1985: 229, figs 217–220 (♀); Xiong, Liu & Zhang, 2017: 23,
figs 1A–G (♂♀); Basumatary et al., 2018: 35, figs 1–5 (♀); Logunov, 2021: 1032, figs 41–44 (♀).
Hyllus maskaranus Barrion & Litsinger, 1995: 65, figs 29a–f (♀); holotype in International Rice Research Institute (IRRI), the
Philippines, examined drawings of the holotype by J. Prószyński (Fig. 12); syn. nov.
Phidippus tirapensis B. K. Biswas & K. Biswas, 2006: 495, figs 4–6 (♀); holotype could not be traced in NZC-ZSI, Kolkata,
not examined; syn. nov.
For a complete list of taxonomic references refer the WSC (2022).
Type material. Phidippus tirapensis B. K. Biswas & K. Biswas, 2006: Holotype ♀ (NZC-ZSI 5536/18) from
INDIA, Arunachal Pradesh, Dist. Tirap, Hukanjuri, 36 km from Khonsa (27.1128°N, 95.4536°E), 234 m a.s.l.,
14.03.1990, leg. T.K. Pal.
Comparative material: Hyllus diardi, 1♀ (JC) from INDIA, Assam, Karimganj District, Bazaricherra
(24.4318°N, 92.3307°E), 51 m a.s.l., May 2016, leg. V. A. Ismavel.
Justification of synonymy of Phidippus tirapensis and Hyllus maskaranus. Phidippus tirapensis Biswas
& Biswas, 2006 was described based on a female holotype collected from Hukanjuri, Arunachal Pradesh. The
holotype could not be traced in the collections at NZC-ZSI, Kolkata. However, based on the original descriptions
and illustrations, the species is large sized (total length 17.5) comparable with 17.28 in Basumatary et al. (2018), with
broad cephalothorax and two hair tufts present in the anterior part of cephalothorax (Fig. 9); unidentate chelicerae
(see Biswas & Biswas 2006: 495); epigyne with two rounded copulatory openings (Fig. 10), wide lateral posterior
pockets; spermathecae wider anteriorly and narrow posteriorly (Fig. 11), all these characters support its placement
in the genus Hyllus C.L. Koch, 1846 and matches strikingly with H. diardi (cf. Figs 9–11 with Figs 7, 8, figs 1B–D
in Xiong et al. 2017 and figs 1–4 in Basumatary et al. 2018). Based on these observations, it is thus safe to consider
the species as a junior synonym of Hyllus diardi.
FIGURES 7–12. Somatic and genitalia morphology of Hyllus diardi (Walckenaer, 1837): 7–8 Hyllus diardi from Assam:
7—habitus, dorsal view; 8—epigyne, ventral view. 9–11 Phidippus tirapensis Biswas & Biswas, 2006: 9—habitus, dorsal view;
10—epigyne, ventral view; 11—ditto, dorsal view. 12 Hyllus maskaranus Barrion & Litsinger, 1995: 12—epigyne, ventral
view. Scale bars: (8) 0.2 mm, (7) 1 mm. Figs 9–11 original illustrations adopted from Biswas & Biswas (2006). Fig. 12. Original
quick drawing of the holotype by J. Prószyński in 2003 (adopted from Prószyński 2016).
TAXONOMIC NOTES ON FIVE SALTICID SPECIES Zootaxa 5230 (3) © 2023 Magnolia Press · 395
Hyllus maskaranus Barrion & Litsinger, 1995 was described based on a female holotype collected from
Mindanao Island, the Philippines. Based on detailed comparison of the original descriptions, illustrations and
further illustration of the female genitalia by Prószyński (2016) (Fig. 12), the species matches unambiguously with
Hyllus diardi. The general body colour pattern and the epigyne with two rounded copulatory openings, wide lateral
posterior pockets; spermathecae with wide anterior and narrow posterior chambers match with that of H. diardi (cf.
Fig. 12 and figs. 29a–f in Barrion & Litsinger 1995 with figs. 217–220 in Żabka 1985 and figs. 1B–D in Xiong et
al. 2017). H. maskaranus is thus considered as a junior synonym of H. diardi.
Distribution. India, Myanmar, Thailand, Laos, China to Indonesia, Philippines (WSC 2022; Barrion & Litsinger
1995: sub Hyllus maskaranus). India: Arunachal Pradesh (Biswas & Biswas 2006: sub Phidippus tirapensis), Assam
(Basumatary et al. 2018; present data) (Fig. 25).
Pancorius manipuriensis (Biswas & Biswas, 2004) comb. nov.
Figs 13–24, 25
Marpissa manipuriensis Biswas & Biswas, 2004: 27 (D♂♀), figs 1–3 (figures not published); holotype ♀, allotype ♂ and
paratype ♀, in NZC-ZSI, examined.
Type material. Marpissa manipurensis B.K. Biswas & K. Biswas, 2004: Holotype (NZC-ZSI) ♀ from INDIA,
Manipur, Ukhrul, Wino Bazar (25.1059°N, 94.3618°E), 1858 m a.s.l., 09.04.1992, leg. A.K. Sanyal & Party.
Paratypes (6♀) and allotypes (2♂) (NZC-ZSI), together with the holotype. Registration numbers: 5488-90/18.
Diagnosis. The male palp of Pancorius manipuriensis is similar to Pancorius submontanus Prószyński, 1992
and Pancorius lui Gan, Mi & Wang, 2022 in having a similar embolus, but can be easily distinguished by the long,
thumb-like posterior tegular lobe directed at 5’o clock position (broad U-shaped and directed at 6’o clock position
in P. submontanus and P. lui); RTA conical, bent sub-apically with the tip directed dorsad (tapering without any
bend near tip and directed apically in P. submontanus; directed apically in P. lui); palpal tibia longer than wide
(as long as wide in P. submontanus and P. lui) (cf. Figs 15, 16 with figs 1B, C in Gan, Mi & Wang 2022). The
female is most similar to P. submontanus in having a sclerotized plate and a pair of epigynal pockets in the anterior
half of epigyne but can be distinguished by the alignment of the copulatory openings which diverge posteriorly
(v-shaped, converging posteriorly in P. submontanus) and the copulatory ducts diverging towards postero-lateral
region (parallel in P. submontanus) (cf. Figs 21–24 with figs 119–124 in Prószyński 1992).
Description. Female (paratype, Figs 18–21): Total length 7.45, carapace 3.04 long, 2.51 wide; abdomen 4.41
long, 3.18 wide. AME 0.65, ALE 0.35, PME 0.07, PLE 0.35, AEW 2.05, PEW 2.10, EFL 1.35. Clypeus height 0.12.
Carapace reddish brown; eyes surrounded by black region (Fig. 18). Clypeus covered with white hairs. Chelicerae
brownish, unidentate. Endites, labium and sternum yellowish. Legs yellowish. Abdomen yellow-brown with a mid-
longitudinal convex lens-shaped yellow stripe outlined with brown margin and lateral sides covered with brown
spots and streaks (Fig. 18). Spinnerets yellow brown. Epigynal plate sclerotized anteriorly and posterior half appears
membranous and translucent; a pair of slit-like curved copulatory openings and a pair of blind pockets present in
anterior portion; copulatory ducts diverge toward postero-lateral sides leading to bulbous anterior spermathecal
chamber, posterior chamber large, globular; fertilization ducts present at anterior part of the posterior spermathecal
chamber (Figs 21–24).
Male (allotype, Figs 13–16): Total length 6.80, carapace 3.31 long, 2.77 wide; abdomen 3.49 long, 2.42 wide.
AME 0.68, ALE 0.38, PME 0.09, PLE 0.35, AEW 2.21, PEW 2.27, EFL 1.47. Clypeus height 0.16. Carapace
reddish brown with yellow patch behind the fovea and extending on the thoracic region; lateral sides of carapace
with thick band white hairs; eyes surrounded by black region (Fig. 13). Clypeus covered with a few white hairs.
Chelicerae reddish-brown, unidentate. Endites, labium and sternum yellowish brown. Leg I robust, reddish-brown;
legs II to IV yellowish. Abdomen brown with yellow spots and mid-dorsal chevron shaped markings (Fig. 13).
Spinnerets brown. Pedipalps brown covered with pale yellow hairs; embolus thick, long, arising from 9’o clock
position; bulb with well-developed conical, thumb-like posterior lobe; RTA thick with pointed tip, bent sub-apically
with tip directed dorsally (Figs 15, 16).
Remarks. Biswas & Biswas (2004) described Marpissa manipuriensis from Ukhrul, Manipur, based on the
female holotype, 6 female paratypes and 2 male allotypes. The original paper referred to illustrations (Biswas &
Biswas 2004: 28) but they were not apparently published in the same work or later publications. The identity of the
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FIGURES 13–17. Somatic morphology and copulatory organs of Pancorius manipuriensis (Biswas & Biswas, 2007) comb.
nov., allotype male: 13—habitus, dorsal view; 14—ditto, front view; 15—left male palp, ventral view; 16—ditto, retrolateral
view; 17—label. Scale bars: (15, 16) 0.2 mm, (13, 14) 2 mm.
TAXONOMIC NOTES ON FIVE SALTICID SPECIES Zootaxa 5230 (3) © 2023 Magnolia Press · 397
FIGURES 18–24. Somatic morphology and copulatory organs of Pancorius manipuriensis (Biswas & Biswas, 2007) comb.
nov., holotype and paratype: 18—paratype female habitus, dorsal view; 19—ditto, front view; 20—ditto, lateral view; 21—
epigyne, ventral view; 22—holotype female, epigyne, ventral view; 23—ditto, dorsal view; 24—ditto, dorsal view. Scale bars:
(18, 19, 20) 2 mm, (21–23) 0.2 mm, (24) 0.1 mm.
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species therefore remained unknown. Although the original publication mentions type designation and appropriate
registration numbers assigned, the vials containing the specimens did not have proper labeling. The label read,
1. Marpissa manipurae sp.n., 14.6.99 (date of identification), 8♀, 2♂ with the signature of Bijan Biswas (Fig.
17), thus indicating that they are the type material used for describing M. manipuriensis. A different spelling for
manipuriensis, ‘manipurae’ is used in the label which is similar to that used for M. mizoramensis (‘mizoae’) in the
original label (Fig. 5) and the figure legends in the publication (Biswas & Biswas 2007: 458). The authors seem
to have used the suffix ‘ae’ for both names—‘manipurae’ and ‘mizoae’ on the labels after initial identification but
eventually modified the names with the suffix ‘ensis’ while publishing the descriptions. We can therefore be sure
that these specimens are indeed the type material used by the authors to describe M. manipuriensis. The larger of two
males is used for description, and the epigyne and vulva already dissected from a female (probably the holotype) are
depicted in Figs 22–24, and one of the paratypes is described and illustrated (Figs 18–21).
Justification for transfer. The characters such as the unidentate chelicerae, abdomen pattern with mid-
longitudinal stripe, the anterior half of epigyne with a sclerotized plate, a pair of pockets and copulatory openings,
multi-chambered spermathecae, the simple male palp with erect embolus arising from the antero-lateral portion
of bulbus, posterior tegular process, single RTA, support its placement in Pancorius (Prószyński 1992; 2017;
Kanesharatnam & Benjamin 2021).
Distribution. Known only from the type locality in India (Manipur) (Biswas & Biswas, 2004) (Fig. 25).
FIGURE 25. Map showing the distribution of Evarcha flavocincta (circles), Hyllus diardi (squares) and Pancorius manipuriensis
(star) studied in this work.
Acknowledgements
I am grateful to Dr Kailash Chandra, former Director of the Zoological Survey of India, Kolkata for allowing
the examination of types and to Dr Shelley Acharya (ZSI, Kolkata) for her support and to Chandan Bera (ZSI,
Kolkata) for his curatorial assistance. I profoundly thank Prof. Dr. Jerzy Prószyński (Poland) for the generous
support throughout the last decade which has greatly enriched the work on Indian Salticids and for allowing me to
use his drawings. I thank Dr Vijay Anand Ismavel for collecting and sending the Hyllus specimen from Assam. The
constructive suggestions of the editor Dr Junxia Zhang (Hebei, China), Dr Dmitri Logunov (Manchester, UK) and
an anonymous reviewer refined the manuscript greatly.
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