Taxonomic review of the ant genus Lepisiota Santschi, 1926 (Hymenoptera: Formicidae: Formicinae) from India

Abstract

The species-rank taxonomy of the genus Lepisiota Santschi, 1926 in India is revised. Thirteen species are recognized, with two described as new: L. layla sp. n. and L. mayri sp. n. The three previously infraspecific taxa L. integra stat. nov., L. pulchella stat. rev. and L. wroughtonii stat. rev. are elevated to species rank. Four species or subspecies are
excluded from the Indian Lepisiota fauna: L. capensis (Mayr, 1862), L. frauenfeldi (Mayr, 1855), L. rothneyi watsonii (Forel, 1894), and L. simplex (Forel, 1892) for issues related to previous doubtful distribution or species misidentifcation. An identification key to the worker caste of Indian Lepisiota species is provided.

Full text

Received: 14.12.2020 Corresponding editor: R. Peters
Accepted: 08.06.2021 Published: 25.06.2021
Bonn zoological Bulletin 70 (2): 227–245 ISSN 2190–7307
2021 · Wachkoo A.A. et al. http://www.zoologicalbulletin.de
https://doi.org/10.20363/BZB-2021.70.2.227
INTRODUCTION
The formicine ant genus Lepisiota Santschi, 1926 con-
tains 138 species and subspecies, including two that are
newly described here (Sharaf et al. 2020; Bolton 2021),
and is widespread in the grasslands, savannahs or wood-
lands of the Afrotropical, Indomalayan, and Palearctic
regions (Brown 2000; Hita Garcia et al. 2013). The tax-
onomy of the genus is in a dreadful condition with the
lack of any revisionary studies for most of the zoogeo-
graphical regions of the World (Sharaf et al. 2020). How-
ever, a number of authors have published isolated species
descriptions and updated, taxonomic revisions for several
regional faunas (Sharaf et al. 2020; Bolton 2021).
The Lepisiota fauna of India is poorly known. Most of
the species and infraspecic taxa are poorly dened and
have been treated under different genera. Apart from the
major contributions in the late nineteenth and the early
twentieth century (Forel 1892, 1894, 1895, 1902a; Bing-
ham 1903), and subsequent isolated treatments (Muker-
jee 1930; Bharti 2002), the fauna has been neglected tax-
onomically.
In this study, we revise the species-rank taxonomy of
the ant genus Lepisiota for India. We describe the two
new species L. layla and L. mayri and recognize the three
poorly dened sympatrically distributed infraspecic
taxa integra, pulchella and wroughtonii, at species rank.
The data reported herein represent the rst deep insight
of the Indian Lepisiota with the hope of correcting some
of the taxonomic neglect that has plagued the Indian For-
micinae (Bharti & Wachkoo 2012, 2014a, b; Wachkoo &
Bharti 2015a, b).
MATERIAL AND METHODS
The taxonomic study was conducted on a Nikon SMZ
1500 stereoscope. For digital images, MP Evolution dig-
ital camera was used on the same microscope with Au-
to-Montage (Syncroscopy, Division of Synoptics, Ltd.)
software. Later, images were cleaned with Adobe Pho-
toshop CS5.
Institutional abbreviations
MHNG = Museum of Natural History, Geneva,
Switzerland
MSNG = Natural History Museum, Genoa, Italy
NHMUK = Natural History Museum, London, UK
PUAC = Punjabi University Patiala Ant Collection,
Punjab, India
Abstract. The species-rank taxonomy of the genus Lepisiota Santschi, 1926 in India is revised. Thirteen species are re-
cognized, with two described as new: L. layla sp. n. and L. mayri sp. n. The three previously infraspecic taxa L. integra
stat. nov., L. pulchella stat. rev. and L. wroughtonii stat. rev. are elevated to species rank. Four species or subspecies are
excluded from the Indian Lepisiota fauna: L. capensis (Mayr, 1862), L. frauenfeldi (Mayr, 1855), L. rothneyi watsonii
(Forel, 1894), and L. simplex (Forel, 1892) for issues related to previous doubtful distribution or species misidentifcation.
An identication key to the worker caste of Indian Lepisiota species is provided.
Keywords. Ants, Formicinae, key, Lepisiota, new species, new status, revived status, India.
Research article
urn:lsid:zoobank.org:pub:165FB2AA-6B1D-446A-9191-EACFB18FD732
Taxonomic review of the ant genus Lepisiota Santschi, 1926
(Hymenoptera: Formicidae: Formicinae) from India
Aijaz Ahmad Wachkoo1, Himender Bharti2 & Shahid Ali Akbar3, *
1
Department of Zoology, Government Degree College, Shopian, Jammu and Kashmir, 192303 India
2
Department of Zoology and Environmental Sciences, Punjabi University, Patiala, Punjab, 147002 India
3 Division of Plant Protection; Department of Entomology, Central Institute of Temperate Horticulture, Srinagar, Jammu and
Kashmir, 191132 India
*
Corresponding author: Email: kingakbarali@gmail.com
1 urn:lsid:zoobank.org:author:6F19EB1F-5DDC-4722-BBD3-F75C29F901D9
2 urn:lsid:zoobank.org:author:5CFEBC9B-3CA9-4459-83A6-6D7B61B984B7
3 urn:lsid:zoobank.org:author:5A0AC4C2-B427-43AD-840E-7BB4F2565A8B

Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
Aijaz Ahmad Wachkoo et al.228
ZSI = Zoological Survey of India, Kolkata,
India
Holotype and paratypes of both new species, and other
newly collected material have been deposited in PUAC.
One paratype of both new species will be deposited at
NHMUK. Morphological terminology for measurements
(given in millimeters) and indices found below follow
Wachkoo & Bharti (2014a, b) and Akbar et al. (2017).
Abbreviations for morphological terms and indices
used in the text
CI = Cephalic index: HW/HL × 100
EL = Maximum length of eye as measured normally
in oblique view of the head to show full surface
of eye
HL = Maximum length of head in full-face view,
measured in straight line from the anterior most
point of the median clypeal margin to a line
drawn across the posterior margin
HW = Maximum width of head in full-face view
(excluding the portion of eyes that extends past
the lateral margins of the head)
ML = Mesosomal length measured from the anterior
surface of the pronotum proper (excluding the
collar) to the posteriormost point of the propo-
deal lobes
PFL = Maximum length of the profemur from its
margin with the trochanter to its margin with the
tibia
PFW = Maximum width of the profemur
PnW = Maximum width of the pronotum in dorsal view
SI = Scape index: SL/HW × 100
SL = Maximum length of the antennal scape
excluding the basal neck and condyle
REL = Relative eye length index: EL/HL × 100
SYSTEMATIC ACCOUNTS
Genus Lepisiota Santschi, 1926
Type species: Plagiolepis rothneyi Forel, 1894
Worker diagnosis. Antenna 11-segmented; eyes
well-developed, ocelli frequently present; propodeum
armed with a pair of spines, teeth, or tubercles; petiole a
scale-like with the dorsal margin bispinose, bidentate or
emarginated; acidopore well-developed (Bolton 1994).
Synoptic list of Indian species of Lepisiota
Lepisiota annandalei (Mukerjee, 1930)
Lepisiota bipartita (Smith, 1861)
Lepisiota fergusoni (Forel, 1895)
Lepisiota integra (Forel, 1894) stat. nov.
Lepisiota layla sp. n.
Lepisiota lunaris (Emery, 1893)
Lepisiota mayri sp. n.
Lepisiota modesta (Forel, 1894)
Lepisiota opaca (Forel, 1892)
Lepisiota pulchella (Forel, 1892) stat. rev.
Lepisiota rothneyi (Forel, 1894)
Lepisiota sericea (Forel, 1892)
Lepisiota wroughtonii (Forel, 1902) stat. rev.
Lepisiota annandalei (Mukerjee, 1930) (Figs 1–3)
Acantholepis annandalei Mukerjee, 1930: 156, g. 4.
Syntype workers, Shimla [Simla], Himachal Pradesh,
India [ZSI].
Lepisiota annandalei (Mukerjee); Bolton 1995: 226; rst
combination in Lepisiota.
Worker measurements. HL 0.50–0.54; HW 0.46–
0.52; EL 0.16–0.18; SL 0.48–0.56; PnW 0.31–0.35;
ML 0.60–0.68; PFL 0.38–0.43; PFW 0.11–0.13 mm. In-
dices: CI 91–95; SI 102–108; REL 29–33 (n = 14).
Description. Head subquadrate; longer than wide,
distinctly wider posteriorly than in front; lateral margins
convex, posterior margin transverse, posterolateral cor-
ners rounded; clypeus subcarinate in the middle; ante-
rior clypeal margin complete and convex; eyes broadly
oval, weakly convex, placed at the middle-line of head,
covering one-third of lateral cephalic margin; three small
ocelli present; antennal scape surpassing posterior head
margin by about one-third its length.
In lateral view promesonotum dome like, convex;
metanotum distinctly lower than promesonotum but al-
most as high as propodeum; mesometanotum demarcat-
ed, constricted; metanotal area distinct; propodeal spines
reduced to tubercles; propodeal declivity steep.
Petiole upright with angular sides, dorsally without
distinct teeth or spines, shallowly emarginate; apical cor-
ners with acute angles pointing upward.
Overall body smooth and shiny.
Gaster covered with short, abundant, erect setae; head
and pronotum with only a few setae; pubescence on body
uniformly very ne and sparse; antennal funiculus with
appressed to subdecumbent pubescence.
Color black; antenna, mandible and tarsi light brown.
Distribution and habitat. India (Himachal Pradesh,
Jammu and Kashmir, Punjab, Sikkim, Uttarakhand, West
Bengal) (Bharti et al. 2016; present study). Although,
infrequent in collections this species seems widespread
across India; workers were mostly hand collected from
tree trunks and honey baits.
Remarks. This species has largely been misidentied
as Afrotropical L. simplex (Forel, 1892) due to its su-
percial resemblance with the latter. The small to medi-
um-sized workers can be distinguished from the some-
what similar L. lunaris by their smooth and shiny body,
a dorsally emarginate petiole and few setae on the prono-

Review of the ant genus Lepisiota from India
Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
229
tum; L. lunaris has a relatively dull and microreticulate
body, a bispinose petiole and several pairs of setae scat-
tered on the mesosoma including the propodeum.
Material examined. India: Himachal Pradesh,
Andretta, 32.0744° N, 76.5856° E, 940 m, 6 ♀♀,
12.vi.2010; Renuka, 30.6083° N, 77.4615° E, 600 m,
3 ♀♀, 8.v.2009; Terrace, 31.9234° N, 75.9294° E,
420 m, 2 ♀♀, 11.x.2008. Jammu and Kashmir, Mans-
ar, 32.6979° N, 75.1489° E, 690 m, 3 ♀♀, 13.vii.2009.
Punjab, Dharampur, 31.8420° N, 75.9132° E, 450 m,
5 ♀♀, 14.x.2008. Uttarakhand, Dakpathar, 30.4966° N,
77.8004° E, 750 m, 5 ♀♀, 20.viii.2009; Forest Rese-
arch Institute, 30.3416° N, 77.9903° E, 640 m, 6 ♀♀,
1.x.2008; Mussoorie, 30.4606° N, 78.0521° E, 1820 m,
4 ♀♀, 9.viii.2009; Selaqui, 30.3720° N, 77.8605° E,
670 m, 3 ♀♀, 7.viii.2009, Aijaz A. Wachkoo leg.
Lepisiota bipartita (Smith, 1861) (Figs 4–6)
Formica bipartita Smith, 1861: 33. Syntype workers,
Lebanon; Israel, Holy Land [NHMUK]. [Images of
CASENT0903167 Syntype worker examined].
Acantholepis bipartita (Smith); Roger 1863: 11; rst
combination in Acantholepis.
Acantholepis bipartita (Smith); Mayr 1863: 394 as junior
synonym of Acantholepis frauenfeldi.
Acantholepis frauenfeldi var. bipartita (Smith); André
1882: 211 queen described and revived from synony-
my as variety of frauenfeldi.
Acantholepis frauenfeldi bipartita (Smith); Emery 1891:
16; Emery 1898: 126; Forel 1902b: 155; Ruzsky 1905:
463; Santschi 1917: 47; variety/subspecies of frauen-
feldi.
Acantholepis frauenfeldi var. bipartita (Smith); Kara-
vaiev 1911: 44; male described.
Lepisiota frauenfeldi subsp. bipartita (Smith); Baroni
Urbani et al. 1992: 303; rst combination in Lepisiota.
Lepisiota bipartita (Smith); Collingwood & Agosti 1996:
365; status as species.
Worker measurements. HL 0.66–0.80; HW 0.57–
0.71; EL 0.19–0.25; SL 0.98–1.09; PnW 0.36–0.53;
ML 0.97–1.06; PFL 0.75–0.98; PFW 0.14–0.20 mm. In-
dices: CI 84–88; SI 148–171; REL 28–31 (n = 15).
Description. Head subrectangular; longer than wide,
equally wide anteroposteriorly; lateral and posterior
margins convex, posterolateral corners rounded; clypeus
medially subcarinate to carinate; anterior clypeal margin
complete and convex; eyes subglobulose, convex, pro-
jecting beyond cephalic lateral margins, covering one-
third of lateral cephalic margin, placed at posterior half
of head; three ocelli present; antenna long, scape surpass-
ing posterior head margin by about half its length.
In lateral view pronotum convex, mesometanotum
strongly constricted, lower than pronotum and propo-
deum, giving mesosoma a dumbbell shape; mesometa-
Figs 1–3. Worker; Lepisiota annandalei (Mukerjee, 1930). 1.
Head, frontal view. 2. Body, lateral view. 3. Body, dorsal view.
1
2
3

Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
Aijaz Ahmad Wachkoo et al.230
notum demarcated; metanotal area distinct; propodeum
armed with a pair of teeth diverging posteriorly; propo-
deal declivity steep.
Petiole upright with distinctly angular sides, dorsally
emarginate, apical corners teeth like pointing upward.
Head and mesosoma feebly shining; gaster relatively
more shiny; overall body weakly microreticulate, howev-
er, sculpturing is effaced in some specimens.
Body covered with erect setae on head, pronotum and
gaster; setae on gaster usually restricted to venter and
segmental margins; pubescence very ne and sparse,
most visible on head; almost absent on mesosoma and
gaster; antennal funiculus with appressed to decumbent
pubescence.
Head and gaster dark brown; mesosoma reddish brown;
antenna and tarsi brown.
Distribution and habitat. Greece, India (Andhra
Pradesh, Gujarat, Haryana, Himachal Pradesh, Jammu
and Kashmir, Karnataka, Kerala, Maharashtra, Megha-
laya, Punjab, Rajasthan, Uttarakhand, West Bengal),
Iran, Israel, Lebanon (Bharti et al. 2016; Dad et al. 2019;
Sharaf et al. 2020). This species is widespread across In-
dia. The specimens were collected by hand, from honey
baits and by beating vegetation.
Remarks. Lepisiota bipartita is a medium to large-
sized ant. The main feature that will enable it to be dis-
tinguished from L. integra, the species with which it is
most likely to be confused, is the shiny bicolored body,
whereas the latter species has a dull and uniformly red-
dish-brown body.
Material examined. India: Himachal Pradesh: Andret-
ta, 32.0744° N, 76.5856° E, 940 m, 1 ♀, 11.vi.2010, 1 ♀,
27.viii.2009. Jammu and Kashmir: Manda, 32.7496° N,
74.8673° E, 500 m, 1 ♀, 15.vii.2009, Aijaz A. Wachkoo
leg.
Lepisiota fergusoni (Forel, 1895) (Figs 7–9)
Acantholepis fergusoni Forel, 1895: 459. Syntype work-
ers, Thiruvananthapuram [Travancore], Kerala, In-
dia [MHNG]. [Images of CASENT0909883 Syntype
worker examined].
Acantholepis fergusoni Forel; Bingham 1903: 319.
Lepisiota fergusoni (Forel); Bolton 1995: 227; rst com-
bination in Lepisiota.
Worker measurements. HL 0.69–0.72; HW 0.65–
0.66; EL 0.17–0.19; SL 0.69–0.70; PnW 0.42–0.45;
ML 0.95–1.05; PFL 0.59–0.61; PFW 0.16–0.17 mm. In-
dices: CI 91–94; SI 105–107; REL 24–26 (n = 5).
Description. Head subquadrate; slightly longer than
wide, wider posteriorly than in front; lateral margins
convex, posterior margin convex, with rounded postero-
lateral corners; clypeus carinate in the middle; anterior
clypeal margin complete and convex; eyes oval, weakly
convex, placed at the middle-line of head, covering about
one-third of lateral cephalic margin; three small ocelli
Figs 4–6. Worker; Lepisiota bipartita (Smith, 1861). 4. Head,
frontal view. 5. Body, lateral view. 6. Body, dorsal view.
4
5
6

Review of the ant genus Lepisiota from India
Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
231
present; antennal scape surpassing posterior head margin
by about one-third its length.
In lateral view promesonotum convex, metanotum low,
almost straight; pronotum at above; mesometanotum
demarcated; metanotal area distinct; mesometanotum
constricted; propodeum armed with a pair of diverging
blunt spines directed backward; propodeal declivity
steep.
Petiole upright with angular sides, dorsally without
distinct teeth or spines, shallowly emarginate; apical cor-
ners with acute angles pointing upward.
Head distinctly reticulate-striate, opaque; mesosoma
dorsally ne microreticulate, overall opaque, laterally
relatively shiny; gaster polished smooth and shining;
propleuron relatively smoother, remainder of lateral
mesosoma longitudinally striate; mesometanotal suture
cross-ribbed.
Body abundantly covered with erect setae; pubescence
very ne and sparse; antennal funiculus with subdecum-
bent to suberect pubescence.
Head reddish-brown, mesosoma and petiole red-
dish-yellow, gaster reddish-brown with a light red-
dish-yellow patch anteriorly on rst gastral tergite.
Distribution and habitat. India (Karnataka, Kerala,
West Bengal), Sri Lanka (Bharti et al. 2016; Dias et al.
2020). This species is infrequent in collections and seems
to be restricted to the coastal plains of India. The speci-
mens were collected in a leaf litter sample.
Remarks. Lepisiota fergusoni is a medium-sized ant.
The reticulate-striate head and blunt propodeal spines
directed backward allow distinction from the somewhat
similar L. pulchella which has a reticulate-punctate head
and pointed propodeal spines directed upward.
Material examined. India: Kerala: Periyar Tiger Re-
serve, Thanikkudy, 9.4997° N, 77.2665° E, 1003 m,
5 ♀♀, 15.x.2011, Shahid A. Akbar leg.
Lepisiota integra (Forel, 1894) stat. nov. (Figs 10–12)
Acantholepis frauenfeldi var. integra Forel, 1894: 411.
Syntype workers, Dharamshala, Himachal Pradesh, In-
dia [MHNG]. [Images of CASENT0909888 Syntype
worker examined].
Acantholepis frauenfeldi integra Forel; Forel 1906: 86;
Santschi 1917: 44; race/stirps of frauenfeldi.
Lepisiota frauenfeldi var. integra (Forel); Bolton 1995:
227;  rst combination in Lepisiota.
Worker measurements. HL 0.66–0.75; HW 0.57–
0.66; EL 0.20–0.22; SL 0.82–0.95; PnW 0.39–0.49;
ML 1.00–1.12; PFL 0.66–0.77; PFW 0.13–0.16 mm. In-
dices: CI 84–88; SI 140–151; REL 27–30 (n = 10).
Description. Head subrectangular; longer than wide,
almost equally wide anteroposteriorly; lateral and poste-
rior margins convex, posterolateral corners rounded; cly-
peus medially subcarinate; anterior clypeal margin com-
plete and convex; eyes subglobulose, convex, projecting
beyond cephalic lateral margins, covering one-third of
lateral cephalic margin, placed at posterior half of head;
three ocelli present; antennal scape surpassing posterior
head margin by about half its length.
In lateral view pronotum convex, mesometanotum
strongly constricted, lower than pronotum and propo-
Figs 7–9. Syntype worker (AntWeb: CASENT0909883); Lepi-
siota fergusoni (Forel, 1895). 7. Head, frontal view. 8. Body,
lateral view. 9. Body, dorsal view. (Photograph courtesy of
Zach Lieberman).
7
8
9

Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
Aijaz Ahmad Wachkoo et al.232
deum, giving mesosoma a dumbbell shape; mesometa-
notum demarcated; metanotal area distinct; propodeum
armed with a pair of teeth diverging posteriorly; propo-
deal declivity steep.
Petiole upright, with distinctly angular sides, dorsum
emarginate with teeth-like apical corners.
Overall body dull and opaque, relatively densely and
evenly microreticulate.
Body covered with erect setae on head and pronotum;
gastral setae restricted to apical segments; pubescence
relatively more as in Lepisiota bipartita and Lepisiota
sericea, ne and sparse, most visible on head and gaster;
sparser on mesosoma; antennal funiculus with appressed
to decumbent pubescence.
Color uniformly reddish-brown.
Distribution and habitat. India (Himachal Pradesh,
Jammu and Kashmir, Madhya Pradesh, Meghalaya, Pun-
jab, Uttarakhand), Pakistan (Bharti et al. 2016; Rasheed
et al. 2019). This species seems to be general in distri-
bution in India; it was collected in non-forest as well as
forest habitats. Workers were mostly collected by beat-
ing vegetation, Winkler’s extractor, hand collecting, from
honey baits and in pitfall traps.
Remarks. Lepisiota integra is raised here to species
rank, originally described by Forel (1894) as an infra spe-
cies taxon frauenfeldi var. integra. This is a medium to
large-sized ant with a dull and uniformly reddish-brown
body and a dorsally emarginate petiole, whereas L. frau-
enfeldi (Mayr, 1855) is a relatively shiny, paler species
with mesosoma, legs and antennae orange, distinctly
contrasting the brown head and gaster, and a dorsally
dentate petiole.
Material examined. India: Himachal Pradesh:
Baijnath, 32.0527° N, 76.6500° E, 1125 m, 9 ♀♀,
17.vi.2010; Bakhra, 31.4087° N, 76.4327° E, 650 m,
21 ♀♀, 7.x.2008; Ghamrur, 31.6620° N, 76.0601° E,
460 m, 9 ♀♀, 1.vi.2009; Ghatti, 31.9300° N, 75.9302° E,
425 m, 1 ♀, 12.x.2008; Guraldhar, 31.6670° N,
76.4684° E, 660 m, 15 ♀♀, 2.vi.2009; Jogi Panga,
31.5408° N, 76.3161° E, 600 m, 13 ♀♀, 9.x.2008; Re-
nuka, 30.6083° N, 77.4615° E, 600 m, 5 ♀♀, 8.v.2009;
Terrace, 31.9234° N, 75.9294° E, 420 m, 10 ♀♀,
24.v.2009, 20 ♀♀, 25.v.2009, 2 ♀♀, 9.vii.2010. Jammu
and Kashmir: Manda, 32.7496° N, 74.8673° E, 500 m,
8 ♀♀, 15.vii.2009; Mansar, 32.6979° N, 75.1489° E,
690 m, 4 ♀♀, 13.vii.2009. Punjab: Chohal, 31.6666° N,
76.0666° E, 450 m, 17 ♀♀, 11.x.2008. Uttarakhand: Fo-
rest Research Institute, 30.3416° N, 77.9903° E, 640 m,
3 ♀♀, 11.v.2009, 3 ♀♀, 12.v.2009; Ranger’s College,
30.3225° N, 78.0445° E, 660 m, 20 ♀♀, 22.v.2010,
9 ♀♀, 25.v.2010, Aijaz A. Wachkoo leg.
Lepisiota layla sp. n. (Figs 13–15)
urn:lsid:zoobank.org:act:4AF8C610-5E75-42B6-921F-E21DED00054E
Type material. Holotype worker. India, Himach-
al Pradesh, Kotla, 31.8821° N, 75.9963° E, 500 m,
22.x.2010. Paratype, 1 worker, same data as for holotype,
Aijaz A. Wachkoo leg.
Figs 10–12. Worker; Lepisiota integra (Forel, 1894) stat. nov.
10. Head, frontal view. 11. Body, lateral view. 12. Body, dorsal
view.
10
11
12

Review of the ant genus Lepisiota from India
Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
233
Worker measurements. HL 0.57–0.58; HW 0.57–
0.58; EL 0.18–0.19; SL 0.64–0.65; PW 0.40–0.41;
PFL 0.50–0.51; PFW 0.12–0.13; ML 0.86–0.87 mm. In-
dices: CI 100; SI 112; REL 33 (n = 2).
Description. Head subquadrate; as long as wide, as
wide posteriorly as in front; lateral margins convex, pos-
terior margin transverse with strongly rounded postero-
lateral corners; clypeus subcarinate in the middle; ante-
rior clypeal margin complete and convex; eyes broadly
oval, weakly convex, placed at the middle-line of head,
covering about one-third of lateral cephalic margin; three
small ocelli present; antennal scape surpassing posterior
head margin by about one-third its length.
In lateral view promesonotum convex, metanotum low,
almost straight; mesometanotum demarcated; metanotal
area distinct; mesometanotum constricted; propodeum
armed with a pair of posteriorly diverging thick blunt
spines; propodeal declivity steep.
Petiole upright with angular sides, dorsally without
distinct teeth or spines, shallowly emarginate; apical cor-
ners with acute angles pointing upward.
Overall body polished, smooth and shining; mesometa-
notal suture cross-ribbed; mesometapleuron longitudi-
nally striate.
Body abundantly covered with long, erect white setae;
scape and legs with relatively shorter erect setae than
on body; antennal funiculus with dense suberect pubes-
cence.
Color black; scape and tarsi yellow brown.
Distribution and habitat. India (Himachal Pradesh).
This species seems to be rare in India: the specimens
were found in the Shivalik range of Western Himalaya.
The specimens were collected by hand from the trunk of
an Acacia tree.
Etymology. The species epithet is an Arabic noun
meaning dark beauty, in reference to the shining black
color of this species.
Remarks. Lepisiota layla is a medium-sized ant. It is
supercially similar to L. lunaris and L. modesta, but its
body is abundantly covered with long, erect white setae
while L. lunaris and L. modesta have a body covered
with usual short erect setae.
Lepisiota lunaris (Emery, 1893) (Figs 16–18)
Acantholepis lunaris Emery, 1893a: 250, plate 6,
g. 12. Syntype workers, Colombo, Sri Lanka [Cey-
lon] [MSNG]. [Images of CASENT0905157 Syntype
worker examined].
Acantholepis capensis lunaris Emery; Forel 1895: 458;
Forel 1906: 86; Forel 1909: 395; Emery 1925: 24; Me-
nozzi 1939: 312; Chapman & Capco 1951: 209; race/
subspecies of capensis.
Acantholepis capensis subsp. lunaria Emery; Chap-
man & Capco 1951: 209; misspelled as lunaria.
Acantholepis lunaris Emery; Collingwood 1970: 379;
status as species.
Figs 13–15. Holotype worker; Lepisiota layla sp. n. 13. Head,
frontal view. 14. Body, lateral view. 15. Body, dorsal view.
13
14
15

Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
Aijaz Ahmad Wachkoo et al.234
Acantholepis lunaris Emery; Imai et al. 1984: 9; karyo-
type described.
Lepisiota capensis subsp. lunaris (Emery); Bolton 1995:
228; rst combination in Lepisiota.
Lepisiota capensis subsp. lunaris (Emery); Bolton 1995:
228; Bharti et al. 2016: 28; Bharti et al. 2017: 42; erro-
neously treated as subspecies of capensis.
Worker measurements. HL 0.56–0.63; HW 0.51–
0.57; EL 0.17–0.19; SL 0.62–0.66; PnW 0.34–0.42;
ML 0.74–0.86; PFL 0.48–0.53; PFW 0.12–0.14 mm. In-
dices: CI 89–91; SI 111–123; REL 29–31 (n = 9).
Description. Head subquadrate; slightly longer than
wide, wider posteriorly than in front; lateral and poste-
rior margins convex, posterolateral corners rounded; cly-
peus subcarinate in the middle; anterior clypeal margin
complete and convex; eyes broadly oval, weakly convex,
placed at the middle-line of head, covering one-third of
lateral cephalic margin; three small ocelli present; anten-
nal scape surpassing posterior head margin by about one-
third its length.
In lateral view promesonotum convex, metanotum low,
almost straight; mesometanotum demarcated; metanotal
area long and constricted; propodeum armed with a pair
of posteriorly diverging thick blunt spines; propodeal de-
clivity steep.
Petiole upright, with angular sides, dorsally emargin-
ate, armed with a pair of almost straight spines pointing
upward.
Head and mesosoma feebly shiny to subopaque; gas-
ter polished, smooth and shiny; overall head and meso-
somal dorsum weakly microreticulate; propleuron shiny;
mesonotum and lateral mesosoma longitudinally striate;
declivous face transversally striate.
Body covered with short, erect, abundant setae on head
and gaster, sparser on mesosoma; pubescence very ne
and sparse, most visible on head; almost absent on me-
sosoma and gaster; antennal funiculus with sparse ap-
pressed to decumbent pubescence.
Black; antenna, mandible and tarsi light brown.
Distribution and habitat. India (Himachal Pradesh,
Jammu and Kashmir), Nepal, Pakistan, Sri Lanka (Thapa
2000; Bharti et al. 2016; Rasheed et al. 2019; Dias et al.
2020). This species seems to be widespread across India,
although infrequent in the collections. The specimens
were collected both in forest and non-forest habitats,
mainly by hand collecting from the dry soil and honey
baits.
Remarks. This small to medium-sized species is sim-
ilar to many workers of the medium sized L. mayri but
the erect setae on the body are distinctly sparse, whereas
the latter species is covered with abundant erect setae and
very hairy overall.
Material examined. India: Himachal Pradesh: Bari,
31.6591° N, 76.5000° E, 520 m, 3 ♀♀, 6.vi.2009; Ghatti,
31.9300° N, 75.9302° E, 425 m, 3 ♀♀, 12.x.2008; Guga,
Figs 16–18. Worker; Lepisiota lunaris (Emery, 1893). 16. Head,
frontal view. 17. Body, lateral view. 18. Body, dorsal view.
16
17
18

Review of the ant genus Lepisiota from India
Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
235
31.6864° N, 76.1898° E, 600 m, 2 ♀♀, 22.x.2008; Jas-
sur, 32.2824° N, 75.8496° E, 520 m, 6 ♀♀, 6.vi.2009;
Kotla, 31.8821° N, 75.9963° E, 500 m, 5 ♀♀, 22.x.2008;
Palampur, 32.1109° N, 76.5430° E, 1200 m, 4 ♀♀,
18.vi.2010; Nagabari, 32.3004° N, 75.8901° E, 420 m,
3 ♀♀, 18.vi.2009; Terrace, 31.9234° N, 75.9294° E,
420 m, 4 ♀♀, 12.x.2009. Punjab: Dunera, 32.4443° N,
75.8900° E, 520 m, 3 ♀♀, 24.vii.2010, Aijaz A. Wach-
koo leg.
Lepisiota mayri sp. n. (Figs 19–21)
urn:lsid:zoobank.org:act:F9BBB1FB-4249-4BCB-9FCF-5128382CF9AC
Type material. Holotype worker. India, Himach-
al Pradesh, Andretta, 32.0744° N, 76.5856° E, 940 m,
21.vi.2010. Paratypes: 12 workers, same data as for
holotype; 7 workers, Himachal Pradesh, Baijnath,
32.0527° N, 76.6500° E, 1125 m, 17.vi.2010; 6 work-
ers, Himachal Pradesh, Kotla, 31.8821° N, 75.9963° E,
500 m, 13.x.2008; 2 workers, Himachal Pradesh, Nahan,
760 m, 30.5596° N, 77.2960° E, 27.viii.2009; 1 work-
er, Himachal Pradesh, Renuka, 30.6083° N, 77.4615° E,
600 m, 8.v.2009; 5 workers, Himachal Pradesh, Terrace,
31.9234° N, 75.9294° E, 420 m, 24.v.2009; 3 workers,
Uttarakhand, Forest Research Institute, 30.3416° N,
77.9903° E, 640 m, 2.ix.2009; 8 workers, Uttarakhand,
Rajaji Forest Area, 30.2483° N, 77.9878° E, 660 m,
21.v.2010; 1 worker, Uttarakhand, Selaqui, 30.3720° N,
77.8605° E, 670 m, 24.v.2010, Aijaz A. Wachkoo leg.
Worker measurements. HL 0.65–0.73; HW 0.60–
0.69; EL 0.20–0.23; SL 0.68–0.77; PnW 0.42–0.50;
ML 0.92–1.04; PFL 0.57–0.64; PFW 0.14–0.17 mm. In-
dices: CI 91–95; SI 111–116; REL 29–31 (n = 15).
Description. Head subquadrate; slightly longer than
wide, wider posteriorly than in front; lateral and posteri-
or margins convex, with rounded posterolateral corners;
clypeus carinate in the middle; anterior clypeal margin
complete and convex; eyes broadly oval, weakly convex,
placed at the middle-line of head, covering about one-
third of lateral cephalic margin; three small ocelli pres-
ent; antennal scape surpassing posterior head margin by
about one-third its length.
In lateral view promesonotum convex, metanotum low,
almost straight; mesometanotum demarcated; metanotal
area distinct; mesometanotum constricted; propodeum
armed with a pair of posteriorly diverging thick blunt
spines; propodeal declivity steep.
Petiole upright, with angular sides, dorsally emargin-
ate, armed with a pair of nearly straight spines pointing
upward.
Overall body shiny; head weakly microreticulate to
smooth without sculpture; pronotum, metanotum and
propodeal dorsum coarsely microreticulate; mesonotum
weakly microreticulate, mesometanotal suture cross-
ribbed; propleuron sometimes weakly longitudinally stri-
ate; mesometapleuron coarsely longitudinally striate.
Body abundantly covered with short, erect setae; scape
and legs with dense suberect pilosity; antennal funiculus
with dense subdecumbent to suberect pubescence.
Figs 19–21. Holotype worker; Lepisiota mayri sp. n. 19. Head,
frontal view. 20. Body, lateral view. 21. Body, dorsal view.
19
20
21

Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
Aijaz Ahmad Wachkoo et al.236
Black; scape and tarsi yellow brown.
Distribution and habitat. India (as L. capensis;
Arunachal Pradesh, Assam, Bihar, Goa, Haryana, Him-
achal Pradesh, Jammu and Kashmir, Jharkhand, Karnata-
ka, Kerala, Madhya Pradesh, Maharashtra, Manipur, Me-
ghalaya, Mizoram, Sikkim, Uttarakhand, West Bengal)
(Bharti et al. 2016; Dad et al. 2019). This species appears
to be widespread on the Indian subcontinent typically re-
ported as L. capensis; specimens were collected by hand,
beating vegetation and honey baiting.
Etymology. The species is named in honor of Gustav
Mayr, author of L. capensis to which it has been a hom-
onym.
Remarks. The species is similar to L. modesta with
which it shares its uniform black color and a medi-
um-sized body. The distinguishing features include a
combination of ne microreticulate sculpture on head
and mesosoma, prominent propodeal spines and abun-
dant erect setae on the body in L. mayri, versus a smooth
and shiny body, reduced propodeal spines, and sparse
erect setae in L. modesta.
Lepisiota modesta (Forel, 1894) (Figs 22–24)
Acantholepis modesta Forel, 1894: 412. Syntype work-
ers, Mussoorie, Uttarakhand, India [MHNG]. [Images
of CASENT0909892 Syntype worker examined].
Lepisiota modesta (Forel); Bolton 1995: 228; rst com-
bination in Lepisiota.
Worker measurements. HL 0.60–0.67; HW 0.52–
0.63; EL 0.16–0.20; SL 0.62–0.70; PnW 0.35–0.44;
ML 0.78–0.92; PFL 0.48–0.57; PFW 0.12–0.15 mm. In-
dices: CI 85–93; SI 110–125; REL 26–29 (n = 12).
Description. Head subquadrate; longer than wide,
wider posteriorly than in front; lateral margins convex,
posterior margin shallowly concave, with rounded pos-
terolateral corners; clypeus carinate in the middle; anteri-
or clypeal margin complete and convex; eyes oval, weak-
ly convex, placed at the middle-line of head, covering
about one-third of lateral cephalic margin; three small
ocelli present; antennal scape surpassing posterior head
margin by about one-third its length.
In lateral view promesonotum convex, metanotum low,
almost straight; mesometanotum demarcated; metanotal
area distinct; mesometanotum constricted; propodeal
spines reduced to tubercles; propodeal declivity steep.
Petiole upright with angular sides, dorsally emarginate,
armed with spines pointing upward.
Overall body shiny; mesometanotal suture cross-
ribbed; mesometapleuron coarsely longitudinally striate
weakly so, on lateropropodeum; propodeal declivity
transversally striate.
Body abundantly covered with erect setae; pubescence
on body uniformly very ne and sparse; antennal funicu-
lus with subdecumbent to suberect pubescence.
Color black; scape, mandible and tarsi yellow brown.
Distribution and habitat. India (Himachal Pradesh,
Jammu and Kashmir, Punjab, Uttarakhand) (Bharti et al.
2016; Wachkoo et al. 2020). Although this species is col-
lected infrequently it seems widespread across India. It
Figs 22–24. Worker; Lepisiota modesta (Forel, 1894). 22. Head,
frontal view. 23. Body, lateral view. 24. Body, dorsal view.
22
23
24

Review of the ant genus Lepisiota from India
Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
237
seems to have a preference for relatively higher altitudes
above 1200 m. Most of the specimens were collected un-
der stones and from honey baits.
Remarks. The medium-sized workers can be dis-
tinguished from the somewhat similar small to medi-
um-sized L. annandalei by a combination of abundant
erect setae on body, a transversally striate propodeal de-
clivity and a bispinose petiolar dorsum; L. annandalei
has sparse erect setae on the body, a smooth and shiny
propodeal declivity and an emarginate petiolar dorsum.
Material examined. India: Himachal Pradesh: Lwasa,
30.7394° N, 77.1528° E, 1200 m, 17 ♀♀, 27.viii.2009.
Jammu and Kashmir: Baderwah, 32.9832° N, 75.7097° E,
1700 m, 18 ♀♀, 30.vi.2009, 30 ♀♀, 1.vii.2009. Utta-
rakhand: Mussoorie, 30.4606° N, 78.0521° E, 1820 m,
37 ♀♀, 9.viii.2009, Aijaz A. Wachkoo leg.
Lepisiota opaca (Forel, 1892) (Figs 25–27)
Acantholepis opaca Forel, 1892: 43. Syntype workers,
Kanara, Karnataka, India [MHNG, MSNG]. [Imag-
es of CASENT0909893, CASENT0905158 Syntype
workers examined].
Acantholepis opaca Forel; Bingham 1903: 318.
Lepisiota opaca (Forel); Xu 1994: 235; rst combination
in Lepisiota.
Worker measurements. HL 0.63–0.70; HW 0.58–
0.65; EL 0.19–0.22; SL 0.64–0.76; PnW 0.42–0.47;
ML 0.79–1.00; PFL 0.53–0.61; PFW 0.14–0.16 mm. In-
dices: CI 91–93; SI 111–120; REL 29–32 (n = 8).
Description. Head subquadrate; slightly longer than
wide, wider posteriorly than in front; lateral margins con-
vex, posterior margin nearly transverse to convex, with
rounded posterolateral corners; clypeus carinate in the
middle; anterior clypeal margin complete and convex;
eyes oval, weakly convex, placed at the middle-line of
head, covering about one-third of lateral cephalic mar-
gin; three small ocelli present; antennal scape surpassing
posterior head margin by about one-third its length.
In lateral view promesonotum convex, metanotum low,
almost straight; pronotum at above; mesometanotum
demarcated; metanotal area distinct; mesometanotum
constricted; propodeum armed with a pair of posteriorly
diverging, upward directed sharp spines; propodeal de-
clivity steep.
Petiole upright, with angular sides, dorsally emargin-
ate, armed with a pair of almost straight spines pointing
upward.
Overall body smooth and shiny; malar space usually
coarsely microreticulate, remainder of head mostly shiny
with effaced sculpture; pronotum smooth and shiny, re-
mainder of mesosomal dorsal feebly microreticulate;
propleuron smooth and shiny, rest of lateral mesosoma
longitudinally striate; mesometanotal suture cross-
ribbed; propodeal declivity feebly transversally striate;
sculpturing almost entirely absent in some specimens.
Figs 25–27. Worker; Lepisiota opaca (Forel, 1892). 25. Head,
frontal view. 26. Body, lateral view. 27. Body, dorsal view.
25
26
27

Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
Aijaz Ahmad Wachkoo et al.238
Body abundantly covered with erect setae; pubescence
very ne and sparse, least on mesosoma; antennal funic-
ulus with subdecumbent to suberect pubescence.
Head reddish-brown, mesosoma and petiole red-
dish-yellow, gaster purplish-black, some specimens with
a light reddish-yellow patch anteriorly on rst gastral
tergite; antenna, mandible and legs reddish-brown.
Distribution and habitat. China, India (Arunachal
Pradesh, Assam, Himachal Pradesh, Goa, Jammu and
Kashmir, Karnataka, Kerala, Madhya Pradesh, Maha-
rashtra, Sikkim, West Bengal), Sri Lanka (Guénard &
Dunn 2012; Bharti et al. 2016; Sheikh et al. 2019; Dias
et al. 2020). This is one of the widespread species of its
genus and relatively common in India. The specimens
were mainly collected by beating vegetation.
Remarks. Lepisiota opaca is a medium-sized ant. It
supercially resembles L. fergusoni and L. pulchella
but can be separated from both by a smooth and shiny
body, whereas L. fergusoni and L. pulchella have dull and
coarsely sculptured bodies.
Material examined. India: Himachal Pradesh:
Andretta, 32.0744° N, 76.5856° E, 940 m, 16 ♀♀,
11.vi.2010; Chanaur, 32.0545° N, 75.6503° E, 600 m,
5 ♀♀, 12.vi.2009, 5 ♀♀, 20.x.2008; Ghatti, 31.9300° N,
75.9302° E, 425 m, 7 ♀♀, 12.x.2008, 6 ♀♀, 27.ix.2009,
3 ♀♀, 28.ix.2009; Guraldhar, 31.6670° N, 76.4684° E,
660 m, 1 ♀, 2.vi.2009, 2 ♀♀, 10.vi.2009; Khatiar,
32.0057° N, 75.9388° E, 450 m, 7 ♀♀, 18.x.2008, 1 ♀,
3.vi.2009; Kotla, 31.8821° N, 75.9963° E, 500 m, 8 ♀♀,
13.x.2008, 4 ♀♀, 22.x.2008, 5 ♀♀, 28.v.2009; Lwasa,
30.7394° N, 77.1528° E, 1200 m, 12 ♀♀, 7.viii.2009;
Nahan, 30.5596° N, 77.2960° E, 760 m, 13 ♀♀,
20.viii.2009, 9 ♀♀, 27.viii.2009; Siholi, 31.9456° N,
75.9949° E, 560 m, 1 ♀, 4.vi.2009; Terrace, 31.9234° N,
75.9294° E, 420 m, 36 ♀♀, 11.x.2008, 1 ♀, 21.x.2008,
4 ♀♀, 26.v.2009, 1 ♀, 13.vi.2009, 11 ♀♀, 24.ix.2009,
10 ♀♀, 25.ix.2009. Jammu and Kashmir: Manda,
32.7496° N, 74.8673° E, 500 m, 7 ♀♀, 15.vii.2009;
Mansar, 32.6979° N, 75.1489° E, 690 m, 1 ♀, 12.vii.2009,
1 ♀, 13.vii.2009; Samba, 32.5537° N, 75.1317° E,
390 m, 2 ♀♀, 11.vii.2009; Surinsar, 32.7009° N,
75.1512° E, 700 m, 2 ♀♀, 14.vii.2009. Punjab: Dharam-
pur, 31.8420° N, 75.9132° E, 450 m, 2 ♀♀, 14.x.2008;
Dunera, 32.4443° N, 75.8900° E, 520 m, 1 ♀, 23.vi.2009;
Thein Dam, 32.4426° N, 75.7305° E, 1 ♀, 24.vi.2009.
Uttarakhand: Dakpathar, 30.4966° N, 77.8004° E, 750 m,
1 ♀, 20.viii.2009; Forest Research Institute, 30.3416° N,
77.9903° E, 640 m, 1 ♀, 17.viii.2009; Mussoorie,
30.4606° N, 78.0521° E, 1820 m, 2 ♀♀, 13.vii.2009; Ra-
jaji Forest Area, 30.2483° N, 77.9878° E, 660 m, 39 ♀♀,
6.viii.2009, 4 ♀♀, 10.viii.2009, 1 ♀, 13.viii.2009; Sela-
qui, 30.3720° N, 77.8605° E, 670 m, 1 ♀, 7.viii.2009,
2 ♀♀, 24.v.2010, Aijaz A. Wachkoo leg.
Lepisiota pulchella (Forel, 1892) stat. rev. (Figs 28–30)
Acantholepis opaca r. pulchella Forel, 1892: 43. Syntype
workers, Pune [Poona], Maharashtra, India [MHNG].
[Images of CASENT0909894 Syntype worker exam-
ined].
Acantholepis pulchella Forel; Dalla Torre 1893: 172;
Bingham 1903: 318; Wu & Wang 1995: 129; Zhou
2001: 168; status as species.
Acantholepis opaca subsp. pulchella Forel; Emery
1893b: 172; Forel 1894: 414; Forel 1895: 458; Emery
1925: 27; Chapman & Capco 1951: 210; Bolton 1995:
228; Guénard & Dunn 2012: 34; Bharti et al. 2016: 28;
Bharti et al. 2017: 42; subspecies of opaca.
Lepisiota opaca subsp. pulchella (Forel); Bolton 1995:
228; rst combination in Lepisiota.
Worker measurements. HL 0.56–0.64; HW 0.51–
0.59; EL 0.17–0.20; SL 0.60–0.68; PnW 0.35–0.42;
ML 0.77–0.87; PFL 0.45–0.53; PFW 0.12–0.14 mm. In-
dices: CI 90–94; SI 111–119; REL 30–31 (n = 15).
Description. Head subquadrate; slightly longer than
wide, wider posteriorly than in front; lateral and poste-
rior margins convex, posterolateral corners rounded; cly-
peus subcarinate in the middle; anterior clypeal margin
complete and convex; eyes oval, weakly convex, placed
at the middle-line of head, covering about one-third of
lateral cephalic margin; three small ocelli present; anten-
nal scape surpassing posterior head margin by about one-
third its length.
In lateral view promesonotum convex, metanotum low,
feebly concave; mesometanotum demarcated; metanotal
area distinct; mesometanotum constricted; propodeum
armed with a pair of posteriorly diverging, upward di-
rected sharp spines; propodeal declivity steep.
Petiole upright, with angular sides, dorsally emargin-
ate, armed with a pair of divergent spines pointing up-
ward and outward.
Head and mesosomal dorsum opaque; gaster polished,
smooth and shining; head and mesosomal dorsum dis-
tinctly reticulate-punctate; propleuron relatively smooth
and shiny, remainder of lateral mesosoma longitudinally
striate; mesometanotal suture cross-ribbed; propodeal
declivity transversally striate.
Body abundantly covered with erect setae; pubescence
very ne and sparse, least on mesosoma; antennal funic-
ulus with subdecumbent to suberect pubescence.
Head brown, mesosoma reddish-brown to dark brown;
antenna, petiole and legs light to dark brown; gaster
black with a light reddish-yellow patch anteriorly on rst
gastral tergite.
Distribution and habitat. China, India (Karnataka,
Kerala, Maharashtra) (Forel 1894; Guénard & Dunn
2012; Dad et al. 2019). This species seems to be distrib-
uted in the Western Ghats and other parts of central In-
dia. Records from Arunachal Pradesh, Assam, Himachal
Pradesh, Jammu and Kashmir, Punjab, Sikkim, Uttara-

Review of the ant genus Lepisiota from India
Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
239
khand and West Bengal (Bharti et al. 2016) seem to be a
misidentication of L. opaca and are excluded until fur-
ther evaluation. The workers were mainly collected from
honey baits and by beating vegetation.
Remarks. This medium-sized ant originally described
by Forel (1892) as an infraspecic taxon opaca r. pul-
chella was raised to species rank by Bingham (1903).
Emery (1925) without any justication reverted it back to
subspecies. It is a valid species, resembling more closely
L. fergusoni than L. opaca and is therefore revived here
to species level. The main characters that enable distin-
guishing it from L. opaca are a combination of a dull and
opaque body with a distinctly reticulate-punctate head
and mesosoma. L. opaca is smooth and shiny with super-
cial reticulate sculpture.
Material examined. India: Karnataka: Gundlupet,
11.8132° N, 76.6858° E, 800 m, 8 ♀♀, 27.ix.2010. Ker-
ala: Salim Ali Bird Sanctuary, 10.0918° N, 76.7420° E,
118 m, 6 ♀♀, 10.x.2011, Shahid A. Akbar leg.
Lepisiota rothneyi (Forel, 1894) (Figs 31–33)
Plagiolepis rothneyi Forel, 1894: 415. Syntype workers,
Belgaum, Karnataka; Barrackpore [Barrakpore], West
Bengal, India [MHNG]. [Images of CASENT0909865,
CASENT0909866 Syntype workers examined].
Acantholepis rothneyi Forel; Santschi 1926: 15; rst
combination in Acantholepis.
Lepisiota rothneyi Forel; Bolton 1995: 228; rst combi-
nation in Lepisiota.
Worker measurements. HL 0.74–1.04; HW 0.71–
1.03; EL 0.20–0.28; SL 0.72–0.88; PnW 0.48–0.63;
ML 0.88–1.09; PFL 0.61–0.81; PFW 0.18–0.25 mm. In-
dices: CI 95–98; SI 98–104; REL 26–27 (n = 13).
Description. Head subquadrate; about as long as wide,
wider posteriorly than in front; lateral margins convex,
posterior margin gently convex, posterolateral corners
rounded; clypeus subcarinate in the middle; anterior
clypeal margin complete and convex; eyes relatively
small, broadly oval, weakly convex, placed at the mid-
dle-line of head, covering three-tenths of lateral cephalic
margin; three small ocelli present; antennal scape sur-
passing posterior head margin by about one-fourth its
length.
Mesosoma constricted in the middle, in lateral view
promesonotum dome like, convex; metanotum distinctly
lower than promesonotum but almost as high as propo-
deum; mesometanotum demarcated; metanotal area short
but distinct; propodeum unarmed without a pair of teeth
or spines; propodeal declivity steep.
Petiole upright, with smoothly curved sides, dorsally
rounded, without a pair of teeth or spines.
Body overall smooth and shiny. Setae restricted to cly-
peus, gastral venter and segmental margins; a few setae
near lateral ocelli usually present; body covered with
sparse pubescence; antennal funiculus with appressed to
decumbent pubescence.
Color brown to black; antenna, mandible and tarsi light
brown.
Distribution and habitat. Bangladesh, China, India
(Karnataka, Kerala, Orissa, Tamil Nadu, Uttarakhand,
West Bengal), Myanmar, Vietnam (Wheeler 1927; Han-
nan 2003; Zryanin 2011, Guénard & Dunn 2012; Bharti
et al. 2016). This species is widespread in India however,
it appears to be a forest ant, restricted to relatively un-
Figs 28–30. Worker; Lepisiota pulchella (Forel, 1892) stat. rev.
28. Head, frontal view. 29. Body, lateral view. 30. Body, dorsal
view.
28
29
30

Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
Aijaz Ahmad Wachkoo et al.240
disturbed areas. The workers were mostly hand collected
from tree trunks and by beating vegetation.
Remarks. The medium-sized ant closely resembles
L. wroughtonii, but can be separated from it by having
a sparsely pubescent and shiny body, lacking erect setae
on the mesosomal dorsum. In contrast, L. wroughtonii is
more pubescent with few short setae on the pronotum.
Material examined. India: Uttarakhand: Forest Re-
search Institute, 30.3416° N, 77.9903° E, 640 m, 6 ♀♀,
11.v.2009, 1 ♀, 13.v.2009, 1 ♀, 30.vii.2009, 2 ♀♀,
26.v.2010; Rajaji Forest Area, 660 m, 30.2483° N,
77.9878° E, 1 ♀, 5.viii.2009, 2 ♀♀, 6.viii.2009, 4 ♀♀,
11.viii.2009, 1 ♀, 6.ix.2009; Selaqui, 30.3720° N,
77.8605° E, 670 m, 4 ♀♀, 7.viii.2009, Aijaz A. Wachkoo
leg.
Lepisiota sericea (Forel, 1892) (Figs 34–36)
Acantholepis frauenfeldi var. sericea Forel, 1892: 41.
Syntype workers, Mussoorie, Uttarakhand, India
[MHNG]. [Images of CASENT0909885 Syntype
worker examined].
Acantholepis sericea Forel; Pisarski 1967: 408; status as
species.
Lepisiota sericea (Forel); Bolton 1995: 228; rst combi-
nation in Lepisiota.
Worker measurements. HL 0.66–0.84; HW 0.59–
0.74; EL 0.20–0.25; SL 0.99–1.08; PnW 0.36–0.52;
ML 1.02–1.27; PFL 0.77–0.97; PFW 0.14–0.18 mm. In-
dices: CI 85–90; SI 146–152; REL 29–30 (n = 6).
Description. Head subrectangular; longer than wide,
slightly narrowed anteriorly; lateral and posterior mar-
gins convex, posterolateral corners rounded; clypeus me-
dially subcarinate; anterior clypeal margin complete and
convex; eyes subglobulose, convex, projecting beyond
cephalic lateral margins, covering one-third of lateral
cephalic margin, placed at posterior half of head; three
ocelli present; antennal scape surpassing posterior head
margin by about half its length.
In lateral view pronotum convex, mesometanotum
strongly constricted, lower than pronotum and propo-
deum, giving mesosoma a dumbbell shape; mesometa-
notum demarcated; metanotal area distinct; propodeum
armed with a pair of teeth diverging posteriorly; propo-
deal declivity steep.
Petiole upright, with smoothly curved sides, dorsally
rounded and narrow, without teeth or spines.
Body dull, overall feebly microreticulate, gastral sculp-
turing even feebler.
Body covered with sparse erect setae on head, pro-
notum and gaster; setae on gaster usually restricted to
venter and segmental margins; pubescence very ne and
sparse, most visible on head and gaster; almost absent on
mesosoma; antennal funiculus with appressed to decum-
bent pubescence.
Color uniformly dark brown to black; antenna, mandi-
ble and tarsi brown.
Figs 31–33. Worker; Lepisiota rothneyi (Forel, 1894). 31. Head,
frontal view. 32. Body, lateral view. 33. Body, dorsal view.
31
32
33

Review of the ant genus Lepisiota from India
Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
241
Distribution and habitat. Afghanistan, India (Him-
achal Pradesh, Jammu and Kashmir, Maharashtra, Ut-
tar Pradesh, Uttarakhand), Iran, Pakistan, Turkmenistan
(Kuznetsov-Ugamsky 1929; Pisarski 1967; Ghahari et al.
2011; Bharti et al. 2016; Rasheed et al. 2020). This spe-
cies seems widespread in India. Specimens were hand
collected, under stones, from tree trunks and the ground.
Remarks. Lepisiota sericea is a medium to large-sized
ant which can be distinguished from the very similar spe-
cies L. integra by a combination of a dark brown body,
smoothly curved sides of the petiole and a narrow, round-
ed petiolar dorsum. L. integra has a reddish-brown body,
angular sides of the petiole and an emarginate petiolar
dorsum with teeth-like apical corners.
Material examined. Himachal Pradesh: Andretta,
32.0744° N, 76.5856° E, 940 m, 9 ♀♀, 21.vi.2010; Bai-
jnath, 32.0527° N, 76.6500° E, 1125 m, 6 ♀♀ , 17.vi.2010;
Lwasa, 30.7394° N, 77.1528° E, 1200 m, 6 ♀♀,
27.viii.2009; Palampur, 32.1109° N, 76.5430° E,1200 m,
7 ♀♀, 18.vi.2010. Jammu and Kashmir: Surinsar,
32.7009° N, 75.1512° E, 700 m, 10 ♀♀, 14.vii.2009. Ut-
tarakhand: Mussoorie, 30.4606° N, 78.0521° E, 1820 m,
7 ♀♀, 9.viii.2009, Aijaz A. Wachkoo leg.
Lepisiota wroughtonii (Forel, 1902) stat. rev. (Figs 37–
39)
Plagiolepis rothneyi r. wroughtonii Forel, 1902a: 292.
Syntype workers, Nilgiris, Tamil Nadu, India [MHNG].
[Images of CASENT0909869 Syntype worker exam-
ined].
Plagiolepis wroughtonii Forel; Bingham 1903: 321; sta-
tus as species.
Plagiolepis rothneyi subsp. wroughtonii Forel; Emery
1925: 23; subspecies of rothneyi.
Lepisiota rothneyi subsp. wroughtonii (Forel); Bolton
1995: 229; rst combination in Lepisiota.
Worker measurements. HL 0.60–0.75; HW 0.54–
0.69; EL 0.16–0.21; SL 0.60–0.73; PnW 0.36–0.48;
ML 0.76–0.96; PFL 0.49–0.62; PFW 0.16–0.20 mm. In-
dices: CI 82–93; SI 103–119; REL 26–28 (n = 13).
Description. Head subquadrate; longer than wide,
wider posteriorly than in front; lateral margins convex,
posterior margin gently convex, posterolateral corners
rounded; clypeus subcarinate in the middle; anterior
clypeal margin complete and convex; eyes relatively
small, broadly oval, weakly convex, placed at the mid-
dle line of head, covering three-tenths of lateral cephalic
margin; three small ocelli present; antennal scape sur-
passing posterior head margin by about one-fourth its
length.
Mesosoma constricted in the middle, in lateral view
promesonotum dome like, convex; metanotum distinctly
lower than promesonotum but almost as high as propo-
deum; mesometanotum demarcated; metanotal area short
but distinct; propodeum unarmed without pair of teeth or
spines; propodeal declivity steep.
Figs 34–36. Worker; Lepisiota sericea (Forel, 1892). 34. Head,
frontal view. 35. Body, lateral view. 36. Body, dorsal view.
34
35
36

Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
Aijaz Ahmad Wachkoo et al.242
Petiole upright, with smoothly curved sides, dorsally
rounded, without a pair of teeth or spines.
Body overall dull covered with relatively dense pu-
bescence. Setae restricted to clypeus, gastral venter and
segmental margins; few short erect setae on pronotum
and near lateral ocelli usually present; pubescence rela-
tively dense, more so on gaster; antennal funiculus with
appressed to decumbent pubescence.
Color black; antenna, mandible and tarsi light brown.
Distribution and habitat. China, India (Himachal
Pradesh, Kerala, Tamil Nadu, Uttarakhand, West Ben-
gal), Sri Lanka (Guénard & Dunn 2012; Bharti et al.
2016; Dias et al. 2020). This species appears widespread
in India and was mainly hand collected from tree trunks,
by beating vegetation, from honey baits and occasionally
in pitfall traps.
Remarks. This medium-sized ant was originally de-
scribed by Forel (1892) as an infraspecic taxon roth-
neyi r. wroughtonii and later elevated to species rank by
Bingham (1903). Emery (1925) reverted it back to sub-
species without any justication. Based on our examina-
tion, we consider it as a valid species and elevate it to
species rank here. The main characters that will enable
it to be distinguished from L. rothneyi, the species with
which it is most likely to be confused, are a combination
of a densely pubescent and dull body with few erect se-
tae on the pronotum, whereas L. rothneyi has a sparsely
pubescent and shiny body, lacking erect setae on the me-
sosomal dorsum.
Material examined. India: Himachal Pradesh: Khatiar,
32.0057° N, 75.9388° E, 450 m, 3 ♀♀, 18.x.2008; Poanta
Sahib, 30.4384° N, 77.6239° E, 420 m, 4 ♀♀, 11.v.2009.
Uttarakhand: Assan Barrage, 30.4417° N, 77.6754° E,
750 m, 3 ♀♀, 21.viii.2009; Forest Research Institute,
30.3416° N, 77.9903° E, 640 m, 3 ♀♀, 1.x.2008, 2 ♀♀,
12.v.2009, 2 ♀♀, 30.vii.2009, 2 ♀♀, 20.v.2010, 4 ♀♀,
26.v.2010; Rajaji Forest Area, 30.2483° N, 77.9878° E,
660 m, 4 ♀♀, 6.viii.2009, 4 ♀♀, 10.viii.2009, 3 ♀♀,
25.v.2010; Ranger’s College, 660 m, 30.3225° N,
78.0445° E, 8 ♀♀, 25.v.2010, 1 ♀, 27.v.2010; Selaqui,
30.3720° N, 77.8605° E, 670 m, 6 ♀♀, 24.v.2010, 10 ♀♀,
7.viii.2009, 1 ♀, 5.ix.2010, Aijaz A. Wachkoo leg.
Species excluded from the Indian fauna
The following species are excluded from the Indian Lepi-
siota fauna and are considered as doubtful records and/or
misidentications.
Lepisiota capensis (Mayr, 1862)
The nominal form of L. capensis (Mayr, 1862) does not
occur naturally in India, despite its implied presence by
the use of the name L. capensis reported by Bingham
(1903). The species referred to in the latter paper, with
abundant yellowish erect setae is clearly not L. capen-
sis, which has sparse dark (brown or black) erect setae
(see Mayr 1862, 1865; Arnold 1920) and is distributed
in southern Africa (not from the Himalayas through to
Northeastern Africa).
Bingham’s (1903) report was apparently based on an
erroneous reporting of the presence of L. capensis with
Figs 37–39. Worker; Lepisiota wroughtonii (Forel, 1902) stat.
rev. 37. Head, frontal view. 38. Body, lateral view. 39. Body,
dorsal view.
37
38
39

Review of the ant genus Lepisiota from India
Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
243
whitish setae in India and Sri Lanka (then Ceylon) by
Forel (1892, 1894), and the attribution of dense pilosity
to L. capensis served only to perpetuate this error until
today (Peter Hawkes & F. Hita Garcia, pers. comm.).
The Indian material currently and in the past attributed
to L. capensis in fact represents a very different species,
and is accordingly described as a new species under the
name L. mayri to minimize confusion by eliminating fu-
ture use of the name L. capensis associated with Indian
species.
Lepisiota frauenfeldi (Mayr, 1855)
Forel (1885) rst considered that L. frauenfeldi was Pale-
arctic in distribution and attributed Indian specimens
from Kolkata (then Calcutta), West Bengal to L. bipar-
tita. But, in his subsequent publications (Forel 1886,
1894), he argued that he had confused L. bipartita with
L. frauenfeldi and accordingly treated specimens from
Kolkata and Barrackpore as L. frauenfeldi. However,
he also mentioned its rare occurrence in India. Bingham
(1903), Rothney (1903), and subsequent Indian authors
until today continued with Forel’s (1886, 1894) unver-
ied reporting without further clarication (see Bharti
et al. 2016).
Lepisiota rothneyi watsonii (Forel, 1894)
Bharti et al. (2016) include distribution of this species
in India based on Wheeler (1921). However, Wheeler
(1921) does not mention its distribution in India and it is
therefore excluded from the known Indian fauna.
Lepisiota simplex (Forel, 1892)
This is an African species that seems unlikely to be found
in India. Forel (1894) rst reported this species from
Orissa based on of its supercial resemblance to the type
material from Somalia. Bingham (1903) and subsequent
Indian authors followed this view without any further ex-
amination of Indian specimens (see Bharti et al. 2016).
Based on our examination, Indian material attributed to
L. simplex belong mostly to L. annandalei and can be dis-
tinguished by few setae restricted to pronotum, whereas
former has several pairs of setae scattered on the meso-
soma including the propodeum.
Key to Indian species of Lepisiota (workers)
1. Antennal scape long, surpassing posterior margin of
head by half its length or more (Figs 5, 11, 35) ..... 2
– Antennal scape shorter, surpassing posterior margin
of head by a third of its length or less (Figs 2, 14,
23) .......................................................................... 4
2. Bicolored; head and gaster dark brown to black,
mesosoma reddish brown (Fig. 5) ............................
....................................................L. bipartita (Smith)
– Uniform reddish-brown or black-brown (Figs 11,
35) .......................................................................... 3
3. Reddish-brown; petiolar scale with distinctly angular
sides, dorsally emarginate with teeth-like apical
corners (Figs 11–12) ......L. integra (Forel) stat. nov.
– Black-brown; petiolar scale with smoothly curved
sides, dorsally rounded and narrow, without teeth
(Figs 35–36) ................................. L. sericea (Forel)
4. Bicolored; mesosoma paler than gaster, mainly or
entirely reddish (Figs 8, 26, 29) ............................. 5
– Whole body uniformly dark brown to black (Figs 2,
14, 38 ) ................................................................... 7
5. All parts of the body shining with supercial
microreticulate sculpture at most (Figs 26–27) ........
......................................................... L. opaca (Forel)
– Body sculpture coarse, general appearance opaque
(Figs 9, 30) ............................................................. 6
6. Head reticulate-striate; propodeal spines blunt,
directed backward; petiole dorsally emarginate
(Figs 7–9) ................................. L. fergusoni (Forel)
– Head reticulate-punctate; propodeal spines pointed,
directed upward; petiole dorsally bispinose (Figs 28–
30) ..............................L. pulchella (Forel) stat. rev.
7 Propodeal spines indistinct; gastral pilosity restricted
to few pairs of black setae on the posterior margin of
tergites (Figs 32, 38) .............................................. 8
– Propodeal spines well-developed in the form of two
broadly-based blunt tubercles, teeth or spines; whole
gastral dorsum covered with abundant pale setae
(Figs 2, 14, 17) ....................................................... 9
8. Pronotum without any setae; body sparsely
pubescent, shiny (Fig. 32) .......... L. rothneyi (Forel)
– Pronotum with few short setae; body fairly pubescent,
opaque (Fig. 38) .... L. wroughtonii (Forel) stat. rev.
9. Body abundantly covered with long, erect white
setae (Fig. 14) ..................................... L. layla sp. n.
– Body covered with usual short erect setae (Figs 17,
20, 23) .................................................................. 10
10. Petiole dorsally emarginate without distinct teeth
or spines; mesosomal setae (if any) restricted to
pronotum (Figs 2–3) ....... L. annandalei (Mukerjee)
– Petiole dorsally distinctly bispinose; setae present
across entire mesosomal dorsum (Figs 17, 20, 23) ..
............................................................................... 11
11. Head and mesosoma smooth and shining; propodeal
spines in the form of two broadly-based blunt
tubercles (Figs 22–23) ................ L. modesta (Forel)
– Head and mesosoma nely microreticulate and dull;
propodeal spines pointed (Figs 16, 20) ................ 12
12. Antennal scape with sparse appressed to decumbent
pubescence; mesosomal setae sparse; smaller species
(HW 0.51–0.57) (Figs 16–18) ....L. lunaris (Emery)
– Antennal scape with dense subdecumbent to suberect
pubescence; mesosomal setae abundant; larger
species (HW 0.60–0.69) (Figs 19–21) .....................
............................................................L. mayri sp. n.

Bonn zoological Bulletin 70 (2): 227–245 ©ZFMK
Aijaz Ahmad Wachkoo et al.244
Acknowledgements. Financial assistance rendered by the Min-
istry of Environment and Forests (Grant No. 14/10/2007-ERS/
RE), Govt. of India, New Delhi is gratefully acknowledged. We
are grateful to Peter Hawkes (University of Pretoria) and Fran-
cisco Hita García (Okinawa Institute of Science and Technolo-
gy) for their helpful comments and suggestions about the man-
uscript. We are also thankful to the teams of AntWeb (www.
antweb.org), antmaps (www.antmaps.org) and AntWiki (www.
antwiki.org) for their immensely useful work. We would also
like to thank the subject editor Ralph S. Peters for his valuable
input.
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