Taxonomic studies on the ant genus Lepisiota Santschi, 1926 (Hymenoptera: Formicidae: Formicinae) in India, with description of four new species

ABSTRACT Taxonomic studies were carried out on the ant genus Lepisiota Santschi, 1926 in India, with the description of four new species, L. binghami sp. nov., L. pusaensis sp. nov., L. satpuraensis sp. nov., and L. wilsoni sp. nov., based on the worker caste. Redescription and new distribution records of five known species, L. annandalei (Mukerjee, 1930), L. bipartita (Smith, 1861), L. integra (Forel, 1894), L. layla Wachkoo, Bharti & Akbar, 2021, and L. pulchella (Forel, 1892), are provided. An identification key to the seventeen Indian species of Lepisiota based on the worker caste is presented. The male genitalia of L. bipartita are described in detail with the illustration of different morphological details. In addition, the DNA barcode of L. annandalei, L. bipartita, L. pulchella, and L. pusaensis sp. nov. is generated. Lepisiota binghami http://zoobank.org/urn:lsid:zoobank.org:act:8AC39CAD-C2B8-4B8C-93C5-F8E891DF1F47 Lepisiota pusaensis http://zoobank.org/urn:lsid:zoobank.org:act:6040B8D0-E6A4-4082-AD98-A1785C33BB96 Lepisiota satpuraensis http://zoobank.org/urn:lsid:zoobank.org:act:0CAEE697-D876-4A9D-B053-AF5FDDD54FD0 Lepisiota wilsoni http://zoobank.org/urn:lsid:zoobank.org:act:119BCDA7-C751-413F-8F53-9980194B6477


Introduction
Lepisiota was originally described by Santschi (1926) as a subgenus of Acantholepis Mayr, 1861. Later, Acantholepis Mayr, 1861 was found to be a junior homonym of Acantholepis Kroyer, 1846 (Pisces), and hence Lepisiota, being the first available replacement name, becomes a valid genus (Bolton 1995). Genus Lepisiota Santschi is a diverse genus of the subfamily Formicinae, comprising 139 species and subspecies (Bolton 2021;Wachkoo et al. 2021;Jaitrong et al. 2022), mainly from Afrotropical, Indomalaya, and Palearctic regions. In India, the taxonomy of the genus HL Head length: Maximum distance from the midpoint of anterior clypeal margin to the midpoint of posterior margin of head, measured in fullface view. HW Head width: Maximum width of the head behind eyes in full-face view. MML Mesometanotum length: Maximum length of mesometanotum in dorsal view. PH Petiole height: measured from petiole sternum to apex in profile view. PRW Pronotal width: Maximum pronotal width in dorsal view. SL Scape length: Maximum scape length excluding basal condyle and neck. TL Total length: Roughly measured from anterior clypeal margin to tip of the gaster. WL Weber's length: Diagonal length of mesosoma in profile view from the posteroventral margin of the propodeal lobe to anterior-most point of the pronotal slope, excluding neck. CI Cephalic index: HW/HL × 100 OI Ocular index: EL/HW × 100 REL Relative eye length index: EL/HL × 100 SI Scape index: SL/HW × 100

Male genitalia study
The apical part of the gaster including abdominal segment nine and genitalia of Lepisiota bipartita was detached and cleared with 10% KOH. Genitalia was dissected and studied under a stereomicroscope (Leica S8AP0). Multi-focused montage images were created using LEICA MC190 HD digital camera attached to the LEICA M205 C stereozoom automountage microscope. Morphological terminologies from Yamada and Eguchi (2016) and Boudinot (2013) were used for description.

Description (Worker): Head
Head slightly longer than broad, sides and posterior margin weakly convex, posterolateral corners rounded and head covered with appressed pubescence (Fig. 1C); posterior margin of head with 2-3 erect setae (including a pair of erect setae between lateral ocelli); palp formula 6,4 and third maxillary segment from base longest of all segments; mandible triangular with five teeth on masticatory margin, from apex third and fifth tooth are smaller than others; antennae 11 segmented, scape extending to posterior margin of head about 1/3 rd of its length, third antennal segment smaller than second and fourth segment, length of second segment is about equal to combined length of third and fourth segment, antennae covered with decumbent to suberect pubescence; antennal insertions touching posterior clypeal margin; clypeus dorsally convex, subcarinate medially, anterior clypeal margin convex, posterior clypeal margin with a pair of long erect setae, anterior margin with two pairs of long erect setae with a downwardly directed long median seta; compound eyes broadly oval, positioned at about midlength of head and covering about 1/3 rd of lateral cephalic margin; three ocelli present but faintly visible.
The patronymic name honours entomologist Charles Thomas Bingham for his pioneering extensive work on Indian Hymenoptera including ants.

Comments
Lepisiota binghami distinctly differs from other known species of the genus Lepisiota from India. It differs from the bicoloured species L. pulchella in the sculpture of body, distribution of setae on the body, and size. Lepisiota pulchella workers have reticulate-punctate sculpture on the dorsum of head and mesosoma with gaster microreticulate and subopaque while the head of L. binghami is faintly microreticulate and shiny, most mesosoma without reticulation, gaster smooth and shiny. Mesosoma of L. pulchella has abundant pilosity whereas L. binghami has very less pilosity. Lepisiota pulchella is comparatively larger (HL 0.58-0.64, WL 0.83-0.91) than L. binghami (HL 0.51-0.56, WL 0.69-0.76) (here we are not considering measurements of Wachkoo et al. (2021) as they might have also considered specimens with less prominent characters of L. pulchella as discussed in personal communication). Lepisiota binghami differs from another similar species L. chutimae in the sculpture of mesosoma and head, colouration of gaster, and petiole height. The dorsum of promesonotum is microreticulate and subopaque, the dorsum of metanotum and propodeal spines are reticulate in L. binghami whereas the dorsum of mesosoma is completely smooth in L. chutimae; dorsum of head is comparatively clear microreticulate with feeble striation below compound eyes in L. binghami whereas L. chutimae is about completely smooth. The first gastral segment is yellowish-brown to brown and the remaining part of the gaster is black in L. binghami whereas the gaster is completely dark brown in L. chutimae. The petiole of L. chutimae is higher (PH 0.33-0.35) than L. binghami (PH 0.24-0.28) (Jaitrong et al. 2022

Description (Worker): Head
Head slightly longer than broad, subquadrate, sides and posterior margin almost straight, posterolateral corners rounded and head sparsely covered with appressed pubescence (Fig. 2C); posterior margin of head with 4-5 erect setae (including a pair of erect setae between lateral ocelli) while three pairs on middle of head; palp formula 6,4 and third maxillary segment from base is the longest of all segments; mandible triangular with five teeth on masticatory margin and third tooth is smallest; mandible covered with decumbent to semi-erect hairs; antennae 11 segmented, scape extending to posterior margin of head about 1/3 rd of its length, third antennal segment is smaller than second and fourth segment, length of second segment is about equal to combined length of third and fourth segment; antennae densely covered with appressed to decumbent pubescence; clypeus dorsally convex, subcarinate medially, anterior clypeal margin convex, posterior clypeal margin having a pair of long erect setae while anterior margin with two pair of long erect setae with a downwardly directed long median seta; clypeus covered sparsely with decumbent pubescence; compound eyes large, broadly oval, convex and positioned at about midlength of head; ocelli three in number.

Mesosoma
Promesonotum convex in profile view, higher than metanotum while almost same height as propodeum ( Fig. 2A); propodeum armed with posteriorly directed, blunt spines; propodeal declivity slanting; the distal end of foretibia with pectinate spur and basitarsus with hairy notch; pronotum with more than 10 erect yellowish setae, mesonotum with two pairs of erect yellowish setae while metanotum with one pair of erect yellowish setae and propodeum with 2-3 pairs of erect yellowish setae ( Fig. 2A); mesosoma having sparse, appressed pubescence.

Metasoma
Petiole upright, dorsally bispinose, sides angular with the presence of petiolar spiracles at about mid-height of the petiole ( Fig. 2A); gastral segments with abundant yellowish erect setae mostly on posterior half except first gastral segment which is entirely covered with sparse yellowish erect setae; acidopore well-developed with the fringe of hairs.

Etymology
The species name refers to the type locality.

Comments
Lepisiota binghami differs from L. pusaensis based on the distribution of setae on body, size, and colouration. In L. pusaensis, the posterior margin of the head has 4-5 erect setae as compared to 2-3 erect setae in L. binghami. Mesosoma of L. pusaensis abundantly covered with erect setae while L. binghami have very less pilosity. L. pusaensis is a comparatively larger (HL 0.58-0.59, WL 0.83-0.85) species than L. binghami (HL 0.51-0.56, WL 0.69-0.76). The first gastral segment in L. binghami is yellowish-brown to brown while the remaining segments are black while L. pusaensis have completely dark gaster in most specimens. L. pusaensis distinctly differs from another similar species L. pulchella in the sculpture of the body. The head and pronotum of L. pulchella are reticulate-punctate whereas L. pusaensis has microreticulate sculpture. The gaster of L. pulchella is subopaque while shiny in L. pusaensis.
Lepisiota pusaensis was possibly misidentified as L. opaca in Wachkoo et al. (2021). Lepisiota pusaensis distinctly differs from L. opaca in the sculpture of the body. Lepisiota opaca is entirely dull with the reticulate-punctate sculpture on the head and mesosoma (Forel 1892) whereas the head of L. pusaensis is microreticulate and shiny; the dorsum of pronotum is microreticulate and subopaque, propleuron shiny.

Ecological and biological notes
Lepisiota pusaensis has been found at the base of the Ficus religiosa L. tree where they made a nest in soil with small entrance holes. The workers were slowly foraging on the tree trunk and ground around their nest. Workers are seen to be collecting dead dry insects and taking them to nest. Workers were also observed tending aphids on the grass, Bothriochloa sp. at IARI, New Delhi. Distribution in India: Delhi. Lepisiota satpuraensis sp. nov. (Fig. 3

Description (Worker): Head
Head slightly longer than broad, subquadrate, sides and posterior margin slightly convex, posterolateral corners rounded, head wider posteriorly than in front and covered with semi-erect comparatively long pubescence (Fig. 3C); four pairs of long erect setae on face of head from posterior margin of head to front of antennal insertions (including a pair of erect setae between lateral ocelli); palp formula 6,4 and third maxillary segment from base longest of all segments while sixth segment distinctly longer than fifth segment; mandible triangular with five teeth on masticatory margin, from apex third tooth smaller than second and fourth tooth; antennae 11 segmented, scape extending to posterior margin of head about 1/3 rd of its length, third antennal segment is smaller than second and fourth segment, length of second segment is about equal to combined length of third and fourth segment, antennae covered with semi-erect pubescence; antennal insertions touching posterior clypeal margin; clypeus dorsally convex, subcarinate medially, having numerous semi-erect yellowish pubescent hairs along with a pair of long erect setae on posterior margin of clypeus while anterior margin with two pair of long dark erect setae with a yellowish median seta; anterior clypeal margin weakly convex; compound eyes broadly oval, positioned at about midlength of head; ocelli three in number.

Mesosoma
Promesonotum convex in profile view, higher than metanotum while about same height as of propodeum (Fig. 3A); mesometathorax constricted; propodeum armed with broad blunt spines and propodeal declivity steep; fore tibiae with a pectinate spur at the distal end and basitarsus with a hairy notch; dorsum of the whole mesosoma with  A. HARSHANA AND D. DEY numerous erect setae (Fig. 3A); propleuron and legs covered with semi-erect pubescence while rest of mesosoma seemingly lacking pubescence.

Metasoma
Petiole upright, dorsally bispinose and emarginated, sides with small setae; first gastral segment fully covered with numerous yellowish erect setae while remaining gastral segments mostly covered on posterior half; acidopore well-developed and fringed with hairs.

Sculpture and colour
Head microreticulate and subopaque, the region below compound eyes weakly longitudinally striate (Fig. 3C); dorsum of mesosoma microreticulate to smooth and shiny (Fig. 3B), mesometapleuron rugose; gaster faintly microreticulate and shiny. Body bicoloured; head, mesosoma, and petiole are dark brown to reddish-brown while compound eyes and gaster are completely black.

Etymology
The species name refers to the Satpura Hills in Central India, where the type locality is located.

Comments
Lepisiota satpuraensis is a unique species of Indian Lepisiota fauna with the following combination of characters: body with comparatively long semi-erect pubescent hairs, numerous setae on the body, body size bigger than any other species reported from India having antennal scape surpassing posterior margin of head about 1/3 rd of its length. Lepisiota satpuraensis differs from the closest species L. fergusoni in the structure of petiole, body size, and body colour. Lepisiota satpuraensis is characterised by distinct bispinose petiole while L. fergusoni does not possess the distinct teeth or spines on the petiole. Lepisiota satpuraensis is larger (HL 0.80-0.83, HW 0.78-0.82) than L. fergusoni (HL 0.69-0.72, HW 0.65-0.66). In L. satpuraensis head, mesosoma, and petiole are dark brown to reddish-brown while the gaster is completely black whereas in L. fergusoni head is reddish-brown, mesosoma and petiole are reddish-yellow, and the gaster is reddish-brown with a light reddish-yellow patch anteriorly on first gastral tergite (Wachkoo et al. 2021). Distribution in India: Madhya Pradesh (Pachmarhi). Lepisiota wilsoni sp. nov. (Fig. 4)

Measurements and indices
Workers (

Description (Worker): Head
Head little longer than broad, sides weakly convex, posterior margin almost straight, posterolateral corners rounded and head covered with appressed pubescence (Fig. 4C); posterior margin of head with 4-5 erect setae (including a pair of erect setae between lateral ocelli); palp formula 6,4 and third maxillary segment from base longest of all segments, sixth segment distinctly longer than fifth segment; mandible with five teeth on masticatory margin, third tooth from apex smaller than fourth tooth, mandibular surface with fine long setae; antennae 11 segmented, scape extending to posterior margin of head about 1/3 rd of its length, antennal insertions touching posterior clypeal margin; antennae with decumbent to semi-erect pubescence; clypeus dorsally convex, subcarinate at middle, having appressed pubescence, posterior margin of clypeus with a pair of long erect setae while anterior margin with two pairs of long erect setae with a median seta; anterior clypeal margin convex; compound eyes broadly oval, placed at about mid-length of head; three ocelli present.

Mesosoma
Promesonotum convex in profile view, higher than metanotum and as high as propodeum (Fig. 4A); mesometathorax constricted; propodeal spines short, broad at the base and blunt at apex; propodeal declivity slanting; fore tibiae with a pectinate spur at the distal end and basitarsus with a hairy notch at the proximal end; pronotum with 5-9 erect yellowish setae, mesonotum with two pairs of erect yellowish setae, metanotum with a pair of erect yellowish setae and propodeum with 3-4 erect yellowish setae (Fig. 4A); mesosoma with very sparse appressed pubescence. Petiole dorsally bispinose, each spine with a single small erect seta; gaster covered with sparse yellowish erect setae and appressed pubescence; acidopore well-developed and fringed with hairs.

Etymology
The patronymic name honours Prof. Edward O. Wilson for his extensive contribution to myrmecology.

Description (Worker): Head
Head little longer than broad, sides slightly convex, posterior margin about straight, posterolateral corners rounded and head covered with sparse appressed pubescence (Fig. 5C); palp formula 6,4 and third maxillary segment from base longest of all segments; mandible with five teeth on masticatory margin, third and fourth tooth from apex smaller than other teeth; antennae 11 segmented, scape extending to posterior margin of head not more than 1/3 rd of its length; antennal insertions touching posterior clypeal margin and a pair of erect setae present little behind the antennal insertions; clypeus dorsally convex, having sparsely distributed appressed pubescence, posterior margin having a pair of long yellowish erect setae while anterior margin with two pairs of long yellowish erect setae with a downwardly directed long median seta, anterior clypeal margin weakly convex; eyes broadly oval, placed at about mid-length of head and covering about 1/3 rd lateral margin of head; three ocelli present with a pair of erect setae in between the lateral ocelli.

Metasoma
Petiole upright, without distinct teeth, emarginated dorsally while its sides are convex; gaster with sparsely distributed erect setae; metasoma covered with sparse appressed pubescence; acidopore well-developed with the fringe of hairs.

Sculpture and colour
Head, mesosoma, and metasoma are smooth and shining except for rugulose mesopleuron. Body uniformly black; mandible, antennae, trochanters, tarsi, fore tibiae, the distal and proximal end of mid and hind tibiae yellowish-brown to brown.

Comments
Lepisiota annandalei has smaller-sized workers, which we have found dwelling on different plants or flowers. It is the smallest among all the Lepisiota species in our collection from Central India and it is not so frequent in collections. Distribution in India: Delhi*, Himachal Pradesh, Jammu & Kashmir, Madhya Pradesh*, Punjab, Sikkim, Uttarakhand, West Bengal.

Description (Worker): Head
Head longer than broad, sides slightly convex, posterior margin almost straight, posterolateral corners rounded (Fig. 5F) and head covered with sparse appressed pubescence; palp formula 6,4 and third maxillary segment from base longest of all segments, fifth maxillary segment distinctly smaller than sixth segment; mandible with five teeth on masticatory margin, third tooth from apex is smallest while second tooth smaller than fourth tooth, mandibular surface having fine setae; antennae 11 segmented, scape extending to posterior margin of head about half its length, antennal insertions touching the posterior margin of clypeus; a pair of erect setae present little behind the antennal insertions, and a pair at about mid-length of head; clypeus dorsally convex, subcarinate medially, anterior clypeal margin convex, clypeus with appressed pubescence, posterior margin of clypeus with a pair of long erect setae while anterior margin with two pair of long erect setae along with a downwardly directed median seta; compound eyes broadly oval, convex and placed little behind mid-length of head; three ocelli present with a pair of erect setae in between lateral ocelli.

Mesosoma
Pronotum and about half of the mesonotum forming an arch in profile view (Fig. 5D); pronotum with 2-4 standing setae; mesometanotum strongly constricted (Fig. 5E) and lower than rest of mesosoma; propodeum armed with a pair of short spines and propodeal declivity slanting.

Metasoma
Petiole upright, dorsally emarginated, and apical corners with very short teeth, sides angular; gastral segments with erect setae on their distal margin and basal part of the first gastral sternum with 2-4 erect setae; acidopore well-developed with the fringe of hairs.

Sculpture and colour
The body is not smooth, with effaced sculpture and subopaque. Body bicoloured; head and gaster dark brown to black, mesosoma reddishbrown; antennae, mandible, distal segments of legs brownish.

Description (Male): Head
Head little longer than broad, sides mostly covered by compound eyes, posterior margin about straight, posterolateral corners rounded (Fig. 6A); palp formula 6,4 and third maxillary segment from base longest of all segments, fifth maxillary segment distinctly smaller than fourth and sixth segment; mandible with five teeth on masticatory margin, third tooth from apex is than fourth tooth, mandible with a pit at its base; antennae 12 segmented, scape extending to posterior margin of head about half its length, third antennal segment is about same length of second and fourth segment; antennal insertions touching the posterior margin of clypeus; clypeus dorsally convex, anterior clypeal margin weakly convex, posterior margin of clypeus with a pair of long erect setae while anterior margin with two pairs of long erect setae along with a downwardly directed median seta; compound eyes large, broadly oval, convex and covering more than half of sides of head; three large ocelli present with a pair of erect setae in between lateral ocelli.

Mesosoma
Pronotum strap-like, anteroventral, and hidden by mesoscutum dorsally; mesoscutum well developed, dorsally flat, and anterior margin rounded; mesoscutellum subtriangular in dorsal view and about the same height as mesoscutum; metanotum strip-like; propodeum lower, slightly convex in profile view; propodeal spiracles well developed, anterolateral in position, and its diameter about equal to the length of metanotum; fore tibiae with a pectinate spur at the distal end and basitarsus with a notch at the proximal end.

Metasoma
The petiole is short, without teeth, dorsally emarginated, and apical corners seemingly rounded; gastral segments with erect setae on their distal margin.

Sculpture and colour
Overall body microreticulate and shiny. Body colour brown to dark brown. Description of the male genitalia (Fig. 6C-H) Abdominal sternum nine wider than long, outline somewhat bat-shaped dorsally, spiculum (anterior apophysis of the sternum nine) long, anterior margins meeting basal to the spiculum at an obtuse angle, lateral margins sloping inwardly, posterior margin concave with weakly bulging medially, anterolateral corners well-produced and posterolateral corners with long setae (Fig. 6C). Genital capsule is longer than broad, cupula broader than long, and dorsally anterior margin of cupula concave (Fig. 6D). Basimere broad with outer surface convex; telomere attached ventrally and the inner side of basimere. In the lateral view, the telomere is a little longer than broad and posterior part with long setae (Fig. 6E). Volsella with setae on the ventral ridge. Digitus is broad and hooked ventrally in the lateral view (Fig. 6F). The lateral view shows penisvalva having more than 15 denticles on the ventral margin, posterior apex hooked ventrally, and foveae sparsely present on the surface (Fig. 6G). Valvura directed dorsoanterolaterally.

Description (Female)
Head little longer than broad, sides weakly convex, posterior margin about straight, and posterolateral corners rounded (Fig. 6B); palp formula 6,4; the third maxillary segment from base longest of all segments, and fifth maxillary segment distinctly smaller than the sixth segment; mandible with five teeth on masticatory margin; antennae 11 segmented, scape extending to posterior margin of head about half its length; antennal insertions touching the posterior margin of clypeus; clypeus dorsally convex, anterior clypeal margin weakly convex, posterior margin of clypeus with a pair of long erect setae while anterior margin with two pairs of long erect setae along with a downwardly directed median seta; compound eyes large, broadly oval, convex and covering more than 1/3 rd of sides of the head; three large ocelli present with a pair of erect setae in between lateral ocelli.

Mesosoma
Pronotum strap-like, anteroventral in position; mesoscutum well developed, dorsally flat, and anterior margin rounded; mesoscutellum subtriangular in dorsal view and about the same height as mesoscutum; metanotum striplike; propodeum lower, slightly convex in profile view; propodeal spiracles well developed, anterolateral in position; fore tibiae with a pectinate spur at the distal end and basitarsus with a notch at the proximal end.

Metasoma
Petiole upright, squamiform, with teeth on the apical corners; gastral segments with erect setae on the posterior margin except for first gastral sternum which has erect setae on the middle part; acidopore well developed and fringed with hairs.

Sculpture and colour
Body sculpture is rough and subopaque. Body colour brown.

Comments
This species is widely distributed in India and adapted well in urban localities. It is frequently encountered and therefore collected easily.

Description (Worker): Head
Head longer than broad, sides are slightly convex, posterior margin almost straight with rounded posterolateral corners, head covered with sparse appressed pubescence (Fig. 7C); palp formula 6,4 and the third segment of maxillary palp from base longest of all, fifth segment smaller than the sixth segment; mandible with five teeth on masticatory margin, the third tooth is smallest; antennae 11 segmented, scape extending to posterior margin of head about half its length, antennal insertions touching the posterior margin of clypeus; a pair of erect setae present little behind antennal insertions, and a pair at about mid-length of the head; clypeus dorsally convex, anterior clypeal margin convex with two pair of erect setae with a downwardly directed median seta, posterior margin with a pair of erect setae; compound eyes broadly oval, convex and placed at about mid-length of the head; three ocelli present with a pair of erect setae between lateral ocelli.

Mesosoma
Pronotum and about half of the mesonotum forming an arch in profile view (Fig. 7A); pronotum with two standing setae; mesometanotum strongly constricted (Fig. 7B) and lower than rest of mesosoma; propodeum armed with a pair of short spines; propodeal declivity slanting.

Metasoma
Petiole upright, dorsally with teeth-like apical corners (Fig. 7A) and emarginated, sides angular; gastral segment with erect setae on their distal margin and basal part of first gastral sternite with 2 erect setae; acidopore welldeveloped with the fringe of hairs. Promesonotum convex, higher than remaining mesosoma in profile view (Fig. 7D); propodeum with broad, thick, posteriorly diverging spines and propodeal declivity steep; fore tibiae having pectinate spur at the distal end, basitarsus having a hairy notch at the proximal end.

Metasoma
Petiole dorsally with angular sides, without distinct teeth or spines, shallowly emarginated, and sides slightly convex; acidopore well-developed and fringed with hairs. Head, mesosoma, and metasoma covered with abundant whitish long erect setae (Fig. 7D-F); legs with sub-erect to erect comparatively short whitish setae; antennal scape with sub-erect to erect setae while funicular segments with sub-erect to decumbent pubescence; mandibular surfaces with fine setae.

Sculpture and colour
Head, mesosoma, and metasoma are smooth and shining except for rugose mesometapleuron. Body uniformly black; antennal scape (except the distal end), tarsi of legs yellowish-brown; mandible, the distal end of antennal scape, funicular segments of antennae brown.

Comments
Lepisiota layla seems to be a rare species. It shows similarities with the South African species, Lepisiota capensis junodi (Forel, 1916), in having numerous whitish erect setae all over the body but it differs by lack of bispinose petiole. Distribution in India: Himachal Pradesh, Madhya Pradesh*.

Forel (1892) (Description of worker)
Head subquadrate with somewhat convex sides (Fig. 8C). Mandibles narrow. Clypeus convex at middle and subcarinate. Antennal scape barely protruding 1/3 rd of their length to the posterior margin of the head.

Description (Workers): Head
Head slightly longer than broad, subquadrate, sides of head weakly convex, posterior margin straight with posterolateral corners rounded (Fig. 8F) and head covered with appressed pubescence; posterior margin of head with 4-5 erect setae (including a pair of erect setae between lateral ocelli), three pairs of setae at middle of head; palp formula 6,4 and third maxillary segment from base longest of all segments, sixth maxillary segment distinctly longer than fifth segment; mandible with five teeth on masticatory margin, third tooth from apex smaller than fourth tooth; antennae 11 segmented with scape extending to posterior margin of head not more than 1/3 rd of its length; antennal insertions touching posterior clypeal margin; antennae covered with appressed to decumbent pubescence; clypeus dorsally convex, subcarinate at middle, having decumbent pubescence, posterior margin having a pair of long yellowish erect setae while anterior margin with two pairs of long yellowish erect setae with a downwardly directed long median seta; anterior clypeal margin convex; compound eyes broadly oval and positioned at about mid-length of head; three ocelli present.

Mesosoma
Promesonotum convex in profile view, metanotum little lower than promesonotum (Fig. 8D); propodeal spine broad at the base and blunt at apex, propodeal declivity steep; fore tibiae with a pectinate spur at the distal end and basitarsus with a notch at the proximal end; dorsum of pronotum with more than 10 erect yellowish setae, mesonotum with two pairs of erect yellowish setae, metanotum with a pair of erect yellowish setae and propodeum with three pairs of erect yellowish setae; mesosoma with very sparse appressed to decumbent pubescence.

Metasoma
Petiole upright, dorsally bispinose, and deeply emarginate (Fig. 8D), lateral sides of petiole weakly convex in frontal view with a pair of long setae little above petiolar spiracles; gastral segments sparsely covered with yellowish erect setae mostly on posterior half except first gastral segment which is almost completely covered with erect setae; gaster with sparse decumbent pubescence; acidopore well-developed and fringed with hairs.

Sculpture and colour
Dorsum of head and mesosoma reticulate-punctate ( Fig. 8E-F); mesopleuron rugose; mandible, clypeus smooth and shiny; gaster microreticulate and subopaque. Body bicoloured; head brown to dark brown; antennal scape yellowish-brown while antennal funicles brown; mesosoma, and petiole golden brown to dark brown, first gastral segment golden brown to brown while remaining gaster black in most studied specimens. The first gastral segment was golden brown in the Pachmarhi population with the remaining segments being black while from the Jabalpur population four specimens had complete black gaster.

Comments
Lepisiota pulchella differs from L. opaca in the sculpture of gaster, which is microreticulate and subopaque in the former while smooth and shiny in the latter. Distribution in India: Karnataka, Kerala, Madhya Pradesh*, Maharashtra.