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Stingless bees (Hymenoptera: Apidae: Meliponini) of the Indian subcontinent:

Diversity, taxonomy and current status of knowledge
Article in Zootaxa · May 2013

DOI: 10.11646/zootaxa.3647.3.1
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Zootaxa 3647 (3): 401–428 ISSN 1175-5326 (print edition)
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Copyright © 2013 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3647.3.1
http://zoobank.org/urn:lsid:zoobank.org:pub:3E2DFCFC-9D75-4245-82F5-9B9FD977160A
Stingless bees (Hymenoptera: Apidae: Meliponini) of the Indian subcontinent:

Diversity, taxonomy and current status of knowledge
CLAUS RASMUSSEN
Department of Bioscience, Aarhus University, Ny Munkegade 114, Bldg. 1540, DK-8000 Aarhus C, Denmark.
E-mail: [email protected]
Abstract
Eight named species of stingless bees are known from the Indian subcontinent: Lepidotrigona arcifera (Cockerell),
Lisotrigona cacciae (Nurse), Lisotrigona mohandasi Jobiraj & Narendran, Tetragonula aff. laeviceps (Smith), Tetragonu-
la bengalensis (Cameron), Tetragonula gressitti (Sakagami), Tetragonula iridipennis (Smith), Tetragonula praeterita
(Walker), and Tetragonula ruficornis (Smith). Lectotypes are newly designated for T. bengalensis and T. ruficornis. Keys,
comparative notes, and illustrations for species identification are provided. The distribution of stingless bees throughout
the Indian subcontinent are summarized and concluding that they are found in most parts of the Indian subcontinent, ex-
cept at higher elevation or the drier interior regions. Additional collections and studies are urgently needed to clearly de-
fine the species limits of the complex “iridipennis” species group.
Key words: Tetragonula, Lisotrigona, Lepidotrigona, Trigona, Melipona, India, Pakistan, Bangladesh, Sri Lanka, Nepal,
Bhutan, taxonomy, potential distribution
Introduction
This is an account of the specimens upon which the eight specific names of stingless bees from the Indian
subcontinent are based, as well as references to what has been published about the distribution and biology of the
stingless bees of that region. With this information, and after additional collecting of specimens and nest data, it
will be possible to prepare a complete revision of the Indian stingless bee fauna even though most of the relevant
type specimens are in museums far from India.
Bee-keeping involving several species of native honey bees (Apis spp) is a very important enterprise with a
long tradition in the Indian subcontinent (including the countries India, Pakistan, Bangladesh, Sri Lanka, Nepal and
Bhutan) (Batra 1977, Engel 1999). Less known is the fact that stingless bees have also been kept for centuries in
India, Sri Lanka and Nepal (Crane 1999, Kumar et al. 2012). As in other regions where stingless bees occur,
colonies have been kept in tree logs, wooden boxes, and clay pots for harvesting small quantities of highly prized
medicinal honey, and also for the wax and propolis, produced and gathered by the bees and used for its household
and curative properties (Crane 1999). While the medicinal properties of the honey are little explored for the Indian
subcontinent when compared to that of the Neotropical species (Rodríguez-Malaver et al. 2009, Choudhari et al.
2012, Surendra et al. 2012), the botanical origin of the honey has been explored (Phadke 1968, Joshi et al. 1998),
as have the plants visited by the bees (e.g., Ramanujam et al. 1993, Viraktamath et al. 1999, Devanesan et al. 2002,
Danaraddi 2007, Danaraddi et al. 2011). In fact, most research on stingless bees from the Indian subcontinent has
focused on their role as crop and native flora pollinators (e.g., Raju et al. 2000, Raju et al. 2009a). Earlier studies of
stingless bees have also dealt with worker communication (Lindauer 1956, Lindauer & Kerr 1958, 1960),
taxonomy and morphology (George 1934, Sakagami 1978, Jobiraj & Narendran 2004, Danaraddi et al. 2012), as
well as their nesting biology (Danaraddi et al. 2009).
Recent accounts report five named species of stingless bees from the Indian subcontinent (Sakagami et al.
1990, Jobiraj & Narendran 2004) and presumably several unnamed species that remains to be described (Sakagami
et al. 1990, Rasmussen & Cameron 2007, 2010). Additional species from the Indian subcontinent are here removed
Accepted by A. Lelej: 3 Apr. 2013; published: 10 May 2013 401

from synonymy raising the total number of named species to eight with the potential to raise further (Lepidotrigona
arcifera (Cockerell), Lisotrigona cacciae (Nurse), Lisotrigona mohandasi Jobiraj & Narendran, Tetragonula aff.
laeviceps (Smith), Tetragonula bengalensis (Cameron), Tetragonula gressitti (Sakagami), Tetragonula iridipennis
(Smith), Tetragonula praeterita (Walker) and Tetragonula ruficornis (Smith)). Stingless bees are widely known in
the Indian subcontinent as “dammer bees” or “dammar bees” (dammar is resin from in amongst dipterocarp trees),
with additional local names commonly applied, e.g., “putka” in Sikkim and Nepal (Gurung et al. 2003, Singh et al.
2011, Lepcha et al. 2012), “ngap siwor”, “ngap hamang”, and “ngap khyndew” in Khasi language (Pugh 1947),
“cherutheneecha” and “arakki” in Kerala (Nair 2003), and “kanameemessa” in Sri Lanka (Karunaratne 2005). The
scientific genus-group names for the different stingless bees from the Indian subcontinent have long been Trigona
Jurine and Lisotrigona Moure (Michener 2007), but recent data support that Trigona s.l. is not a single evolutionary
linage, as it is not a monophyletic clade. Rather, Trigona s.l. encompasses a Neotropical clade and a distantly
related Indo-Malayan/Australasian clade (Rasmussen & Cameron 2007, 2010). The solution has been to restrict the
usage of Trigona to the Neotropical lineage while treating the names proposed by Moure (1961) for the Indo-
Malayan/Australasian taxa at the genus level (Rasmussen 2008, Michener 2013), including Lepidotrigona Schwarz
and Tetragonula Moure. The latter was proposed with Trigona iridipennis Smith, 1854 from Sri Lanka as the type
species.
The few genera of stingless bees from the Indian region can easily be separated using published keys (e.g.,
Sakagami et al. 1990, Michener 2007). However, the species-level identification is not trivial. A notoriously
complex genus of stingless bees is Tetragonula, with more than 30 described species and no global identification
key (Rasmussen 2008). Specimens of Tetragonula from the Indian subcontinent were treated as a single species in
the key by Sakagami (1978). Older keys, such as those of Bingham (1897) and Schwarz (1939), cannot reliably be
used for species identification; in particular, the first reference contains several misinterpretations of species and
both references include many species from outside the region.
No comprehensive summary of stingless bee distribution from the region has been published, with the
exception of Sri Lanka (Karunaratne et al. 2005b). Within India, many scattered distribution records exist for states
in north and south, east and west, as well as from the distant Andaman Islands (Sakagami 1978, National
Horticulture Mission 2012). Stingless bees are also reported from Nepal (Partap 1999, Gurung et al. 2003, Sharma
2004) and Bangladesh (Islam 1997, Hannan 2007), although the exact localities and species identity in those
countries needs to be established. In the Neotropical region, stingless bees are known from 34.90°S (Montevideo)
in Uruguay (Camargo & Pedro 2007) up to 27.03°N (Álamos, Sonora) in Mexico (Búrques 1997). In Africa they
are distributed between 28.54°S (Eshowe) in South Africa up to 18.00°N (Njala) in Sierra Leone (information
extracted from gazetteer in Eardley 2004), while in the Indo-Malayan/Australasian region they are known from
36.41°S in Australia (Dollin et al. 1997) up to 24.23 °N in Taiwan (Sakagami & Yamane 1984, 1987, Sung et al.
2006). However, the northernmost global record of extant stingless bees is Dehra Dun, Uttar Pradesh (30.32°N) in
India (Sakagami 1978) with several other Indian records above 28°N (Nogueira-Neto 1951, Goel & Kumar 1990,
Gurung et al. 2003, Chaudhary & Singh 2007, Gupta et al. 2011). It thus appears that stingless bees are known
from most parts of the Indian subcontinent, at least up to 1000 m a.s.l. in India and Nepal (Sinu & Shivanna 2007).
In South America and Asia they become rare above 2500 m a.s.l., although exceptionally they have been recorded
up to 4000 m a.s.l. in the Andes of Peru and Bolivia (Camargo & Moure 1996, Sung et al. 2006).
Given the steady interest in pollination by stingless bees in India and a renewed global interest in cultivation
and propagation of stingless bee colonies for conservation, the purpose of the present paper is to summarize
information about the described species of stingless bees and provide morphometric data on the type specimens, as
well as photographic documentation of their habitus. This will facilitate species identification in the region. In
addition, I compile and map all published records of stingless bees from the Indian subcontinent. As such data are
fragmentary with many states unrepresented or with few presence points, I further apply species distribution
modeling to identify the potential range where these bees may naturally occur. This should inspire more and much
needed studies of stingless bees from the region. Hopefully, new collections can be made of these bees for
taxonomic purposes, particularly capturing male specimens with associated workers (collected from the same
nests) and photographic documentation of the nest entrances. The hidden male genitalia have proven to be valuable
for diagnosing species, even in cases where almost no external morphological differences can be recognized
(Sakagami 1978). Longer series of workers are necessary to describe and account for the variation in coloration and
pilosity within a single species. Lastly, sometimes external nest features such as nest entrances are the most
402 · Zootaxa 3647 (3) © 2013 Magnolia Press RASMUSSEN

diagnostic field traits observed, distinguishing closely related species (Camargo & Pedro 2003, 2004, Rasmussen
2004, Rasmussen & Camargo 2008). It should also be established as a standard procedure to collect additional
workers from the sampled nests into cold-kept ethanol for subsequent DNA studies and molecular confirmation of
identification. Only through such careful and rigorous collection and documentation across the subcontinent, can a
comprehensive revision of the true diversity of stingless bees from the region be made. Likely the diversity in any
parts of the Old World will not reach more than 50 species per site as in highly diverse parts of South America
(Roubik pers. comm., Rasmussen & Gonzalez 2009), but there is certainly a potential for encountering several
species of stingless bees at any site in India.
MAP 1. All localities from the Indian subcontinent where stingless bees have been studied according to the references listed in
appendix 1. Notice occurrences near the border of Pakistan and within Bangladesh and Nepal. Possibly stingless bees can be
found in parts of Pakistan and Bhutan as well.
Material and methods
Morphological terminology follows that of Michener (2007) and Camargo & Pedro (2009), including the traits
measured in the latter. Photographs were prepared using a Canon 7D digital camera with a MP-E 65 mm lens and
stacked with the Zerene Stacker v1.04 software package. Measurements in mm were made with an ocular
micrometer on a Leica stereomicroscope. Abbreviations used are HW = head width, WL = wing length, WD =
STINGLESS BEES OF THE INDIAN SUBCONTINENT Zootaxa 3647 (3) © 2013 Magnolia Press · 403
wing diagonal which is the distance between M-Cu bifurcation and basal tip of marginal cell, HTL = hind tibia
length. The primary types of all species described from India and Sri Lanka, except Lisotrigona mohandasi, were
examined; the loaning institutions are BMNH = Natural History Museum, London, England; OUMNH = Oxford
University Museum of Natural History, Oxford, England; USNM = National Museum of Natural History,
Washington DC, USA.
To produce distribution map 1, all references listed in Appendix 1 were searched for occurrence records of
stingless bees (only records with detailed locality indication or complete coordinates were included). Due to
uncertainty of the identification of species of Tetragonula, I have not distinguished between the different species
reported and all records have simply been entered as a distribution record at a higher taxonomic rank than genus. A
total of 155 unique records were entered as coordinates in DIVA-GIS v7.5 (Hijmans et al. 2001). To estimate the
potential full distribution of stingless bees from the Indian subcontinent, I used Maxent v3.3.3k software with
default parameters for modeling of species’ geographic distributions (Phillips et al. 2006). Modeling was based on
19 bioclimatic variables from the WorldClim database (Hijmans et al. 2005) at a spatial resolution of 2.5 arc-min
(bioclimatic variables included: annual mean temperature, mean diurnal range, isothermality (temperature mean/
range), temperature seasonality (coefficient of variation), max temperature of warmest period, min temperature of
coldest period, temperature annual range, mean temperature of wettest and driest quarter, mean temperature of
warmest and coldest quarter, annual precipitation, precipitation of wettest and driest period, precipitation
seasonality, precipitation of wettest and driest quarter, precipitation of warmest and coldest quarter). Model results
were processed and visualized using the DIVA-GIS v7.5 (Hijmans et al. 2001).
Systematics
Key to the identification of the known stingless bees from the Indian subcontinent (workers only)
1. Inner surface of hind basitarsus uniformly setose, without a differentiated basal sericeous area (Fig. 2f) . . . . . . . . . . . . . . . . . 2
-. Inner surface of hind basitarsus with a differentiated basal sericeous area of short, dense, hairs (Figs 1a–b). (Tetragonula
Moure 1961) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2. Head and thorax with dense tessellation and without shiny interspaces; densely plumose (“scale-like”) hairs present on the
margin of mesoscutum (Fig. 7c); body and wing each more than 4 mm in length (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . Lepidotrigona arcifera (Cockerell 1929) [or other species of the genus Lepidotrigona Schwarz].
-. Head and thorax with brilliant and polished integument (Fig. 3i); mesoscutum without densely plumose hairs; body and wing
each less than 3 mm in length. (Lisotrigona Moure 1961) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Head and thorax dark brown to black with metasoma brown (Fig. 3); clypeus with minute hairs; metanotum finely imbricate
(see also text for a discussion of the two Indian Lisotrigona species) . . . . . . . . . . . . . . . . . . Lisotrigona cacciae (Nurse 1907)
-. Head and thorax all black with metasoma brown; clypeus with plumose hairs; metanotum reticulate anteriorly. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lisotrigona mohandasi Jobiraj & Narendran 2004
4. Larger species with forewing length, including tegulae, between 4.2 and 4.5 mm; head width more than 1.7 mm. Mesoscutum
with bands of pubescence weakly defined, not as clearly defined with glabrous interspaces as on Fig. 1d. . . . . . . . . . . . . . . . 5
-. Smaller species with forewing length, including tegulae, between 3.5 and 4.2 mm; head width 1.5–1.8 mm. Mesoscutum with
distinct bands of pubescence separated by broad glabrous interspaces (Fig. 1d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . “iridipennis” species group (See tables 1 and 2).
5. Antenna ventrally mostly testaceous to ferruginous . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragonula aff. laeviceps (Smith 1857)
-. Antenna ventrally totally blackish brown to black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragonula gressitti (Sakagami 1978)
Lepidotrigona arcifera (Cockerell 1929)

(Figs 2a–l, map 2)
Trigona arcifera Cockerell 1929: 591–592: Holotype (BMNH 17b.1081, worker): examined, “Type” (red border), “B.M.
TYPE / HYM. / 17B.1081”, “Trigona / arcifera Ckll / TYPE”. “Trigona / ventralis Sm / ♀ from Sladen”, “Teesta / Bridge /
Himalayas / India / 10.1.97”, “T.D.A. Cockerell / B.M.1936-415”. Type locality: INDIA, Sikkim, Teesta bridge (on
January 10, 1897) [ca. 27.11°N, 88.47°E].
Provenance: Theodore Dru Alison Cockerell (1866–1948) received a single worker specimen probably directly
through the collector, Frederick William Lambert Sladen (1876–1921) (see Sladen 1915). Sladen, who was an
apiculturist and bumble bee worker, had collected numerous bees in the Himalayan parts of India (Anon 1921),

which were described by Cockerell and other authors (e.g., Blüthgen 1926). Cockerell misread the handwritten
specimen label (Fig. 2a) as it is not “Testa” Bridge, but rather the old “Teesta” Bridge across the river Teesta or
Tista, flowing for most of the length of the Sikkim state bordering West Bengal. The bridge joins the road between
Kalimpong and Darjeeling. The locality is actually near the border of Nepal and thus suggests the hitherto formally
unrecorded presence of the genus Lepidotrigona in Nepal.
FIGURE 1. 1A. Scanning electron microscope image demonstrating the inner surface of hind tibia and basitarsus, the latter
with the differentiated basal sericeous area of short, dense, hairs characteristic of the genus Tetragonula of the Indian
subcontinent (specimen is Heterotrigona itama Cockerell from Malaysia). 1B. Close-up of the hind basitarsus with sericeous
area of short, dense, hairs. 1C. Lepidotrigona with diagnostic dense tessellation on head and thorax and densely plumose hairs
on the margin of mesoscutum (Lepidotrigona cf. arcifera from India, Nagaland, collection 21). 1D. The “iridipennis” species
group is characterized by distinct bands of pubescence separated by broad glabrous interspaces on the mesoscutum (specimen
from India, Tamil Nadu, collection 17). Notice that the latter two specimens are laterally glued directly to the entomological
pin, thus, avoiding to pin through the mesoscutum and obscure characters from this area.
Notes on the type: The holotype is unfortunately slightly damaged, with the left hind leg (Figs 2f–g), the left
antenna (Fig. 2d), and abdomen (Figs 2i–j) glued to a point-mount. The right hind leg and the right antenna are
missing. Due to the glue on the abdomen, the extent of the dark band on the pale metasomal tergum 1 cannot be
properly determined (Fig. 2i). It appears to form a complete band, with a projection in the center (condition B3 in
Fig. 11 of Sakagami 1975); the hairs on vertex, scutellum and mid tibia are very light to pale brown.
Comments: This species was tentatively recognized as a valid species within the “ventralis” species group
(Rasmussen 2008), although Schwarz (1939) suggested it should be synonymized with L. flavibasis (Cockerell
1929) and Moure (1961) suggested synonymization with L. hoozana (Strand 1913b). Within Lepidotrigona, the
“ventralis” species group is characterized by smaller body size (body and forewing length each less than 5 mm),
FIGURE 2. Lepidotrigona arcifera (Cockerell): holotype of Trigona arcifera. 2A. Labels. 2B. Dorsal view of head and
mesosoma (metasoma glued separately to the point mount). 2C. Frontal view of head. 2D. Antenna glued to point mount. 2E.
View of propodeum. 2F. Inner (posterior) view of left hind leg, glued to point mount. 2G. Outer (anterior) view of left hind leg.
2H. Left lateral view of head and mesosoma. 2I. Dorsal view of metasoma, notice adherent; hiding part of the basal tergum. 2J.
Ventral view of metasoma. 2K. Right ventral view of head and mesosoma. 2L. Right forewing.

and the diagnostic densely plumose mesoscutal hairs usually whitish (not yellow) and not extending to the
scutellum (Figs 2b, 7c). The scutellum is bordered with rather long, erect hairs. The hind tibia in the “ventralis”
group is not as apically expanded (spoon-like) as it is in some of the other species of the genus (i.e., “nitidiventris”
species group). No taxonomic revision exists, but species are traditionally distinguished by a combination of body
size, integument coloration on the first and apical metasomal terga and coloration of hairs on the vertex, scutellum,
and posterior margins of the mid and hind tibiae (Schwarz 1939, Sakagami 1975). Two species of the same species
group that occur the nearest and could be expected in the subregion are L. ventralis (Smith) and L. flavibasis. The
first species can be recognized by the two lateral and separate dark spots on the otherwise pale metasomal tergum 1
(forms a complete band in L. arcifera) while the second species can be recognized by the semi-circle of black
integument, partly enclosing the basal depression on the otherwise pale tergum 1, as in L. arcifera; L. flavibasis can
be recognized by the apical metasomal terga brownish to blackish (yellowish on L. arcifera, Fig. 2i) and the fore
and middle tibiae with blackish hairs on the external surface (Schwarz 1939). Such characters do show intraspecific
variation over a large geographical range, which makes it difficult to separate and delimit the species convincingly.
Likely, the separation of the different species has to be done while examining longer series of specimens from more
localities, and including associated males, information on nest architecture, and molecular data.
Lisotrigona cacciae (Nurse 1907)

(Figs 3a–i, map 2)
Melipona cacciæ Nurse 1907: 619: Lectotype (BMNH 17b.1103, worker): examined, “LECTOTYPE” (blue border), “Type”
(red border), “B.M. TYPE / HYM. / 17B.1103”, “Melipona / cacciæ / (Nurse)”, “Col. C.G. Nurse / Collection. / 1920-72”,
“Hoshung- / -abad / Type”, “LISOTRIGONA / cacciae (Nurse) / det. M. S. Engel, 1999”, “LECTOTYPE / Melipona /
cacciae Nurse / design. J. S. Moure, 1961 / (ref: Engel, Oriental Insects, 2000)”. Type locality: INDIA, Madhya Pradesh,
Hoshangabad [ca. 23.25°N, 78.2°E].
Provenance: Charles George Nurse (1862–1933) based his description on specimens collected by A.M.F. Caccia,
of the Indian Forest Department, and presented to Nurse though the courtesy of Isaac Henry Burkill (1870–1965),
reporter on Economic Products to the Government of India since 1902.
Comments: Engel (2000) synonymized Trigona scintillans Cockerell, 1920 from Sandakan, Borneo, under L.
cacciae (from India). Engel (2000) found no structural differences between the specimens and although coloration
varied both within and between specimens from different locations, he pointed out that even the original type series
of L. cacciae from a single locality, as described by Nurse (1907), included specimens with variable coloration
(“abdomen dark red … [to] almost black in some specimens”). Jobiraj and Narendran (2004) in their account of
Lisotrigona from India did not have access to the type specimen of L. cacciae and made a few misinterpretations in
their comparisons to this species: in L. cacciae the head is near black in some specimens (not brown to dark
brown), the gena is finely punctate (not impunctate) (Fig. 3g), pubescence on clypeus is too short to clearly
distinguish whether or not it is plumose (stated as being simple) (Fig. 3d), but the hypoepimeron (immediately
above the scrobal groove) has very fine hairs (not without hairs). In addition, the metanotum is imbricate but
approaching reticulate sensu Harris (1979). The diagnostic characters for L. mohandasi thus approach L. cacciae,
and a direct comparison of the two primary type specimens is necessary to encounter reliable diagnostic characters
for their separation.
Lisotrigona mohandasi Jobiraj & Narendran 2004

(map 2)
Lisotrigona mohandasi Jobiraj & Narendran 2004: 39, 40–43, figs 1–4: Holotype (Calicut University Zoology Department,
worker); paratypes (Jobiraj collection (2)): Not examined. Type locality: INDIA, Kerala, Thrissur, Peechi, Kerala Forest
Research Institute (on April 3rd, 2000) [ca. 10.53°N, 76.35°E].
Provenance: T. Jobiraj and Thekke Curuppathe Narendran described this species based on three specimens
collected by Dr. K. Mohandas.
FIGURE 3. Lisotrigona cacciae (Nurse): lectotype of Melipona cacciae. 3A. Habitus, dorsal view. 3B. Frontal view of head.
3C. Habitus, left lateral view. 3D. Lower frontal view of head, including partially hidden mandibles. 3E. Labels. 3F. Habitus,
right lateral view. 3G. Lateral view of head including genal area. 3H. Right forewing. 3I. View towards propodeum.
Comments: The holotype of L. mohandasi was not examined. Some of the characters indicated by Jobiraj &
Narendran (2004) for separating the present species from L. cacciae are incorrect as discussed under L. cacciae.

Tetragonula species of the “iridipennis” species group
Diagnosis: Amongst Tetragonula species groups members of the “iridipennis” group are characterized by having a
dark mesoscutum with four distinct hair bands separated by broad glabrous interspaces (Fig. 1d), and by their
smaller body size, with the forewing length, including tegula, measuring 3.0 to 4.3 mm. In addition, Sakagami
(1978) reported that the male gonostylus arises dorsad (not laterad) to the gonocoxite in all species of this group.
Comments: Besides the Indian species of Tetragonula (i.e., T. bengalensis (Cameron), T. iridipennis (Smith),
T. praeterita (Walker) and T. ruficornis (Smith)), the group also includes the smallest member of the genus, the
apparently widespread T. fuscobalteata (Cameron 1908) and T. clypearis (Friese 1909) from Australia and New
Guinea (Dollin et al. 1997). A last species in the group is T. pagdeni (Schwarz 1939) described based on the
holotype (in USNM) male collected from Nakhon Si Thammarat in southern Thailand (Sakagami 1978), but
widespread in all of southeast Asia (Sakagami et al. 1990). Sakagami (1978) reported that T. pagdeni workers were
only statistically different from T. iridipennis s.l. from India (the former with paler coloration of the anterior
corbicular fringe and more plumose frontal hairs), although males of T. pagdeni did differ in genitalic
characteristics from the Indian specimens of T. iridipennis s.l. Preliminary molecular results (Rasmussen &
Cameron 2007, 2010) placed four genetically distinct but morphologically “iridipennis”-like specimens from India
in a highly supported clade together with specimens identified as T. pagdeni (from Malaysia), T. clypearis (from
Australia) and, surprisingly, also T. minor (Sakagami 1978) (from Thailand). Tetragonula minor is a very small
member of the genus and is tentatively placed in this species group, as the male is unknown and it cannot be
assessed whether the male gonostylus arise dorsad from the gonocoxite. Workers of Tetragonula minor are
distinguished from the other members by the darker hairs of the mesoscutum and corbicular fringe. The principal
measurements of the species of this species group are given in table 1 (on average HW=1.6–1.8, WL=3.7–4.3,
WD=0.9–1.2, HTL=1.4–1.7) and as follows for species not encountered in the Indian subcontinent: T.
fuscobalteata: HW=1.35–1.5, WL=3.1–3.4, WD=0.9–1.0, HTL=1.2–1.4; T. clypearis: HW=1.57, WL=3.55,
WD=0.97, HTL=1.46; T. minor: HW=1.6, WL=3.6–3.8, WD=1.0, HTL=1.5; T. pagdeni: HW=1.6–1.8, WL=3.9–
4.1, WD=1.0–1.2, HTL=1.4–1.7. The amber fossil Meliponorytes devictus Cockerell 1921 from the Hukawng
Valley, Myanmar, has been placed under junior synonymy of T. iridipennis (Zeuner & Manning 1976), but could
represent an additional member of this species group. Although Hukawng amber generally is recognized as old
(Cruickshank & Ko 2003), this particular piece is copal of much more recent but unknown age (Grimaldi et al.
1995).
TABLE 1. Measurements (mm) of the examined primary type specimens of species described from the Indian
subcontinent and listed above.
Lepidotrigona Lisotrigona Tetragonula T. T. T.
arcifera cacciae iridipennis praeterita ruficornis bengalensis
Total length of body 4.70 2.95 3.55 3.33 3.45 3.55
Width of head 1.89 1.19 1.60 1.52 1.66 1.70
Length of head (clypeal apex-vertex) 1.51 1.01 1.30 1.32 1.32 1.34
Length of compound eye 1.13 0.83 1.10 1.04 1.13 1.13
Width of compound eye 0.47 0.33 0.40 0.40 0.42 0.43
Upper interorbital distance 1.23 0.75 1.00 1.04 1.02 1.02
Maximum interorbital distance 1.34 0.85 1.13 1.12 1.14 1.13
Lower interorbital distance 1.13 0.65 0.82 N/A 0.79 0.87
Diameter of median ocellus 0.08 0.11 0.15 0.14 0.14 0.15
Interocellar distance 0.36 0.27 0.33 0.35 0.38 0.38
Ocellorbital distance 0.32 0.18 0.23 0.24 0.24 0.23
Interalveolar distance 0.19 0.13 0.15 0.15 0.17 0.17
Alveolorbital distance 0.32 0.19 0.30 0.27 0.31 0.26
Alveolocellar distance 0.75 0.54 0.71 N/A 0.67 0.67
......continued on the next page
TABLE 1. (Continued)
Lepidotrigona Lisotrigona Tetragonula T. T. T.
arcifera cacciae iridipennis praeterita ruficornis bengalensis
Alveolar diameter 0.10 0.06 0.14 0.15 0.12 0.12
Length of clypeus 0.36 0.24 0.33 N/A 0.39 0.33
Maximum width of clypeus 0.71 0.42 0.70 N/A 0.71 0.65
Intertentorial distance, width of clypeus 0.56 0.36 0.48 N/A 0.48 0.51
Clypeocellar distance 0.24 0.06 0.96 N/A 0.06 0.14
Length of malar space 0.13 0.02 0.03 N/A 0.05 0.06
Length of scape 0.60 0.37 0.57 0.59 0.57 0.57
Diameter of scape 0.13 0.07 0.10 0.08 0.10 0.10
Diameter of third flagellomere 0.14 0.11 0.12 0.13 0.13 0.13
Length of pedicel + flagellomeres 1.35 N/A 1.24 N/A 1.25 1.30
Length of first flagellomere 0.10 0.06 0.09 0.05 0.07 0.11
Length of second flagellomere 0.10 0.06 0.07 0.12 0.11 0.11
Length of third flagellomere 0.12 0.07 0.10 0.10 0.11 0.11
Length of mandible 0.65 0.45 0.62 N/A 0.62 0.60
Width of mandible 0.21 0.12 0.19 N/A 0.21 0.18
Length of forewing (excluding tegula) 4.25 2.37 3.44 3.44 3.20 3.50
Length of forewing (including tegula) 4.60 2.65 3.80 3.70 3.50 3.80
Width of forewing 1.65 0.95 1.32 N/A 1.28 1.25
Length of pterostigma 0.71 0.36 0.48 0.48 0.51 0.48
Width of pterostigma 0.19 0.12 0.11 0.11 0.13 0.13
Length of marginal cell 1.40 0.89 1.21 1.21 1.23 1.21
Width of marginal cell 0.32 0.18 0.30 N/A 0.30 0.29
Length of first abscissa of M 0.77 0.38 0.58 0.60 0.52 0.55
Length of first abscissa of Cu 0.89 N/A 0.71 0.70 0.70 0.71
Length of wing diagonal 1.33 0.73 1.01 0.95 0.95 0.95
Hamuli 6 6 5 5 5 5
Length of mesoscutum 1.08 0.71 0.87 0.85 0.83 1.00
Width of mesoscutum 1.26 0.93 1.01 1.00 1.05 1.13
Width of scutellum 0.94 0.48 0.57 0.61 0.69 0.79
Length of scutellum 0.28 0.23 0.33 0.24 0.33 0.28
Length of tibia III 1.43 0.86 1.55 1.47 1.43 1.51
Width of tibia III 0.57 0.31 0.54 0.54 0.52 0.54
Length of basitarsus III 0.62 0.35 0.50 0.48 0.48 0.50
Width of basitarsus III 0.26 0.18 0.29 0.27 0.27 0.29
Width of tergum III 1.58 1.05 1.36 1.28 1.32 1.28
Length of hairs on clypeus <0.01 <0.01 <0.01 <0.01 <0.01 <0.01
Length of hairs on frons 0.02 0.02 <0.01 <0.01 <0.01 <0.01
Length of hairs on vertex 0.19 0.04 0.13 0.11 0.07 0.05
Length of hairs on scutellum apex 0.21 0.13 0.12 0.12 0.10 0.15
Problems: More than one hundred years ago four names were proposed for four very similar Tetragonula
specimens from India and Sri Lanka. Those four species were poorly characterized in their time and they have
never since been directly compared. This creates an uncertainty as to the true identity of these four validly proposed

species. They might simply be synonymous, or they may be the correct names for some of the several species of
Tetragonula that inhabit the Indian subcontinent. I have had the opportunity to examine and document with
photographs the primary types of all four species. The external morphology of the primary types might provide
sufficient and reliable characters for separating these species, but intraspecific variation (size, pubescence, and
coloration) is at the same time obscuring the limits of each of the biological species. Ideally, additional characters
from the male genitalia and molecular data will provide enough data to convincingly separate and define the
species, but such data is not available at this point. It is beyond the scope of the present work to locate male
specimens from the type localities or sample fresh specimens for use in molecular studies, to provide an assessment
of the validity of each of these four species. Generalities about each of the species are provided below, in addition,
detailed measurements of the type specimens are found in table 1, photographs of the type specimens are provided
in figures 4–7, and lastly, table 2 provides direct comparison of each of the four primary types.
TABLE 2. Notable differences found by direct examination of the four primary types of the “iridipennis” species group
in the Indian subcontinent, other than Tetragonula ruficornis being the generally lightest and T. praeterita the darkest of
the four primary types.
T. iridipennis T. praeterita T. ruficornis T. bengalensis
Erect setae on vertex dark brown dark brown light brown light or slightly darker
brown
Plumose hairs on abundant sparse, maybe worn abundant abundant
clypeus and frons
Mandible chestnut-brown without (not visible) amber-yellow with near amber-yellow with near
black apical area black apical area black apical area
Erect setae on margin a few dark brown very light very light very light
of scutelum
Pubescence on almost none, except on almost none, except on sparsely covered almost none, except on
metasomal T3-T4 apical third apical third apical third
Stout setae of outer dark brown dark brown white brown
surface of hind tibia
Inner surface of hind basal sericeous area basal sericeous area basal sericeous area basal sericeous area
basitarsus more than half the more than half the less than half the length more than half the
length of basitarsus length of basitarsus of basitarsus length of basitarsus
Marginal cell of lightly infuscate, as rest lightly infuscate, as rest less infuscate, as rest of less infuscated, as rest
forewing of wing (Fig. 4i) of wing (Fig. 4i) wing (Fig. 6g) of wing (Fig. 6g)
Tetragonula iridipennis (Smith 1854)

(Figs 4a–j, map 2)
Trigona iridipennis Smith 1854: 413–414: Lectotype (BMNH 17b.1114, worker): examined, “Type” (orange border),
“iridipennis / Type Sm.”, “B.M. TYPE / HYM. / 17B.1114”, “TRIGONA / iridipennis / TYPE. Smith.”, “Ceylon” (reverse
side “53 / 23”). In addition “LECTOTYPE Trigona iridipennis Smith Design. JS Moure 1961 / C Rasmussen 2013”; Type
locality: SRI LANKA, Central Province, Kandy [ca. 7.27°N, 80.60°E, ca. 467m a.s.l.].
Provenance: Frederick Smith (1805–1879) described this species merely as from Sri Lanka (then Ceylon).
However, the label code of the lectotype indicates that the specimen was purchased by the Natural History Museum
in London as part of lot 23 in 1853. According to the register at the museum, this lot was sold by Hugh Cuming
(1791–1865), an insect dealer from London, and originated from Sri Lanka where it had been collected by George
Henry Kendrick Thwaites (1811–1882). Thwaites went to Sri Lanka in 1849 as director or superintendent of the
historical and important Royal Botanical Gardens of Peradeniya, near the city of Kandy in the Central Province
(Anon 1882a, 1882b). Presumably the type specimen must have been collected around this area, although Thwaites
also traveled to other parts of Sri Lanka during his time in Sri Lanka.
FIGURE 4. Tetragonula iridipennis (Smith): lectotype of Trigona iridipennis. 4A. Habitus, dorsal view. 4B. Frontal view of
head. 4C. View towards propodeum and metasoma. 4D. Lower frontal view of head, including partially hidden mandibles. 4E.
Mesepisternum 4F. Lateral view of head, including genal area. 4G. Outer view of right hind tibia. 4H. Inner view of right hind
tibia. 4I. Left fore- and hindwing. 4J. Labels.

Lectotype: Smith (1854) did not state how many specimens he saw of T. iridipennis. In the case of additional
type specimens (or syntypes) it is therefore necessary to designate a lectotype to stabilize the future use of the
name. However, as Moure (1961) already referred to the above specimen as the holotype, the ICZN Article 74.6
(ICZN 1999) allows the inference by Moure (1961) as a valid lectotype designation, as the original description did
not imply syntypes.
FIGURE 5. Tetragonula praeterita (Walker): lectotype of Trigona praeterita. 5A. Habitus, dorsal view. 5B. Upper frontal view
of head, notice from previous photo that the specimen is glued directly to the pin mount. 5C. Habitus, left lateral view. 5D.
Genal area and lower face, including mandibles. 5E. Habitus, right lateral view. 5F. View towards propodeum. 5G. Outer view
of left hind leg. 5H. Inner view of left hind leg. 5I. Labels (“prateria W” is written on front and “d) Specimen” is written on rear
of same label).
Tetragonula praeterita (Walker 1860)
(Figs 5a–i)
Trigona præterita Walker 1860: 305–306: Lectotype (BMNH 17b.1185, worker): examined, “63 / 52”, “Type” (green border),
“præterita / W.”, “Trigona”, “B.M. TYPE / HYM. / 17b.1185” (taxonomy). In addition “LECTOTYPE Trigona praeterita
Walker Design. JS Moure 1961 / C Rasmussen 2013”; Type locality: SRI LANKA.
Provenance: Francis Walker (1809–1874) only stated that the specimen came from Sri Lanka (then Ceylon). The
label code indicates that the type specimen was purchased by the Natural History Museum in London as lot 52 in
1863. Unfortunately the register only indicates that this lot, containing 116 Hymenoptera from Sri Lanka, was
presented to the museum by the Entomological Society and included the types described by Walker in the Annals
and Magazine of Natural History during 1858, 1859 and 1860. The original collector or the type locality can
therefore not currently be further traced.
Lectotype: Walker (1860) did not state how many specimens he saw of T. praeterita. In the case of additional
type specimens, it is therefore necessary to designate a lectotype to stabilize the future use of the name. However,
as Moure (1961) already referred to the above specimen as the holotype, the ICZN Article 74.6 (ICZN 1999)
allows the inference by Moure (1961) as a valid lectotype designation, as the original description did not imply
syntypes.
Notes on the type: The type is glued with the entire ventral side adhered to the point mount. The lower part of
the face is not clearly visible. In addition, it appears that the specimen was conserved in alcohol first, as the
compound eyes are deflated rendering the head width measurements smaller.
Comments: Tetragonula praeterita was first synonymized under T. iridipennis by Schwarz (1937), although
Schwarz noticed that the description indicated a larger species (2.5 English lines in body length and 4 lines in wing
length equals 5.3 mm and 8.4 mm, respectively). As the specimen otherwise agrees with the description and the
label information, and the register is consistent with the interpretation that this specimen is a type specimen seen
and described by Walker, the mistake in measurements must be attributed to the general standard and brevity of
Walker’s scientific work (Evenhuis 2011). The synonymy of T. praeterita under T. iridipennis was later followed
by Moure (1961) upon examination of the type and by Sakagami (1978) without examining the type.
Tetragonula ruficornis (Smith in 1870)

Trigona ruficornis Smith in Horne & Smith 1870: 185, 194: Lectotype (BMNH, worker): examined, “India” (typed, with hand-
written reverse “69 / 86”), “SYNTYPE” (blue border), “SYNTYPE [worker symbol] / Trigona / ruficornis / F. Smith,
1870: 194 / det. D. Notton, 2012”). In addition “LECTOTYPE Trigona ruficornis Smith Design. C. Rasmussen 2013”;
Type locality: INDIA, Uttar Pradesh, Varanasi (formerly Benares) (on April 4th, 1863) [25.28°N, 82.96°E].
Melipona smithii Bingham 1897: 560, 563: Unnecessary replacement name for Trigona ruficornis Smith, nec “Lamarck”.
Lamarck 1817 (and repeated in 1835) only listed these five stingless bee species in the genus Melipona: M. favosa, M.
amalthea, M. ruficrus, M. postica, and M. pallida. The replacement name therefore remains enigmatic as there was no M.
ruficornis, unless Bingham had it mistaken for M. ruficrus Latreille (not Lamarck, and -crus, Latin, in reference to leg, not
-cornis, horned, in reference to the antennae).
Provenance: Frederick Smith (1805–1879) described this species based on several specimens and the report of a
single nest collected by Charles Horne (1824–1872). Horne had located the nest in Varanasi (formerly Benares),
Uttar Pradesh, but Smith cited confusingly and probably incorrectly the type locality as Mainpuri, Uttar Pradesh,
almost 500 km NW of Varanasi (many of the other insect species from this publication were in fact collected in
Mainpuri and maybe hence the confusion?). The register of the Natural History Museum in London indicates that
lot 69 from 1886 included, among others, three specimens of T. ruficornis from India presented by Horne as part of
type specimens described earlier. David Notton of the BMNH also located the remains of a nest (BMNH specimen
number 650787) labeled as Trigona from India 84/38. Referring to the register, this was part of a lot of
Hymenoptera from India, collected by the late Charles Horne, and donated by his widow but also relating to Horne
& Smith (1870) probably representing the nest described in that paper. The nest is apparently mostly batumen and
cerumen from the internal parts of the nest, thus, not including diagnostic features such as the nest entrance or
brood combs.

FIGURE 6. Tetragonula ruficornis (Smith): lectotype of Trigona ruficornis. 6A. Habitus, dorsal view. 6B. Frontal view of
head. 6C. Habitus, right lateral view. 6D. Lower frontal view of head, including mandibles. 6E. Outer view of left hind leg. 6F.
Inner view of left hind leg. 6G. Right forewing. 6H. View towards propodeum. 6I. Labels.
Lectotype: Although a single specimen was separated earlier and labeled as the “holotype” (specimen BMNH
17b.1187, labeled “India” (reverse “69 / 86”), “Type” (red border), “Trigona / ruficornis / (Type) Smith”, “B.M.
TYPE / HYM. / 17b.1187”), the account by Smith clearly indicates that the species was described from a nest, and
multiple specimens must have been acquired by the museum. Two additional specimens with label data “69 / 86”
have been located by D. Notton in the general collection under the name T. smithii, and are now considered
syntypes. As already pointed out by Moure (1961), the “holotype” is headless and it would therefore not be
desirable to select this particular specimen as the lectotype. Fortunately, Moure (1961) treating this headless
specimen as holotype does not qualify as lectotype by inference (ICZN 1999, Article 74.6), as the biological
account accompanying the original description clearly indicate that there was a syntype series (ICZN 1999).
Instead, I here designate a complete syntype specimen as the lectotype in order to stabilize the future use of the
name.
Comments: Moure (1961) suggested the headless specimen should be a callow, but the pigmentation appears
to be complete, and all of the lecto- and paralectotype specimens are rather light colored compared to other Indian
species. Apparently Schwarz (1939) was the first to suggest the synonymy of T. ruficornis under T. iridipennis, but
he later changed his mind and identified bees from India as T. ruficornis (see Nogueira-Neto 1949, 1951).
Tetragonula bengalensis (Cameron 1897)

Trigona bengalensis Cameron 1897: 143–144: Lectotype (OUMNH, worker): examined, “Trigona / bengalensis / Cam.”. In
addition “LECTOTYPE Trigona bengalensis Cameron Design. C Rasmussen 2013”; Type locality: INDIA, West Bengal,
26 km N of Kolkata (formerly Calcutta) on the east bank of the Hooghly river (also known as Hugli) (sometime during
1872–1886 or 1893) [ca. 22.68°N, 88.38°E].
Provenance: Peter Cameron (1847–1912) described this species based on at least five specimens collected by
George Alexander James Rothney (1849–1922). These five specimens are all in agreement with the original
description of T. bengalensis (only one labeled as such), but one of these individuals, standing over the name
“bengalensis” in the Rothney drawer in OUMNH, is labeled in different handwriting and on a more recent paper
label as T. iridipennis. As Cameron did not refer to T. iridipennis in his account Hymenoptera Orientalia, he must
have been unaware of the species proposed by Smith (from Sri Lanka) and the label could have been added
subsequently by an unknown as a sign of synonymy. However, Rothney (1903) himself later provided an account
of the type locality stating that T. bengalensis was rare compared to T. iridipennis.
Lectotype: I here designate as the lectotype the only specimen from the original Rothney drawer labeled as
“bengalensis” in order to stabilize the future use of the name. The remaining four specimens standing over the
drawer label “bengalensis Cam.” are considered conspecific paralectotypes. An additional OUMNH drawer label is
printed, in red, “BARRACKPORE: / Rothney” in support of these being authentic type specimens.
Comments: Tetragonula bengalensis was first synonymized under T. iridipennis by Bingham (1897) without
further discussion. Most later authors followed this, although Sakagami (1978), in an addendum (p. 247), pointed
out that males from Sri Lanka (type locality of T. iridipennis) and India (type locality of T. bengalensis) differed in
the genitalia, but were otherwise identical. Therefore Sakagami proposed to use T. iridipennis for the Sri Lankan
population and T. bengalensis for the Indian population (ignoring the fact that T. ruficornis was an earlier name),
although he cautioned that T. pagdeni (Schwarz 1939) from Thailand might not differ from the latter.
Tetragonula species of the “laeviceps” species group
Members of this species group usually have a mesoscutum almost as evenly banded with hair as the “iridipennis”
group, but these species are larger, with forewing length, including tegula, measuring between 4.2 and 4.8 mm.
Sakagami (1978) referred to this species group as the taxonomically most difficult group within Tetragonula. A
single species has been reported in India.
Tetragonula aff. laeviceps (Smith 1857)
Specimens similar to Trigona laeviceps Smith 1857: 51: Neotype (BMNH, worker): examined; Type locality: SINGAPORE,
Upper Peirce Reservoir Park.

FIGURE 7. Tetragonula bengalensis (Cameron): lectotype of Trigona bengalensis. 7A. Habitus, dorsal view. 7B. Frontal view
of head. 7C. Habitus, right lateral view. 7D. Lower frontal view of head, including partially hidden mandibles. 7E. Habitus, left
lateral view. 7F. Labels. 7G. View towards propodeum. 7H. Mesepisternum. 7I. Outer view of right hind leg.
Comments: Recognizable type material of T. laeviceps could not be found and a neotype from the type locality,
Singapore, was recently designated to stabilize the usage of the name (Rasmussen & Michener 2010). Specimens
similar to the highly variable T. laeviceps have been reported from Bangalore in India by Sakagami (1978) and
Sakagami et al. (1990). In addition to reporting this species from mainland India, Sakagami (1978) examined a
worker of presumably T. aff. laeviceps from the Andaman Islands. Sakagami et al. (1990) also reported a “forma
B” of Tetragonula for India, but it is uncertain which species that would represent.
MAP 2. Type localities for the species of stingless bees from the Indian subcontinent. a. Tetragonula iridipennis, b. Lisotrigona
mohandasi, c. L. cacciae, d. T. ruficornis, e. T. bengalensis, f. Lepidotrigona arcifera. Tetragonula praeterita was described
from “Ceylon”, or Sri Lanka, with no further indication of the locality.
Tetragonula species of the “gressitti” species group
The only species in this group is T. gressitti, which Sakagami (1978) defined by having the male mandible receded
and apex of gonocoxite clavate.
Tetragonula gressitti (Sakagami 1978)
Trigona (Tetragonula) gressitti Sakagami 1978: 214–216: Holotype (not located, see Rasmussen (2008), worker): not
examined; Type locality: VIETNAM, Lâm Đồng province in the Central highlands.

This species is characterized by distinctly melanic coloration including ventral side of entire antenna, and relatively
long malar space and scape. This species was recently reported in from Hunli and Pashighat, Lower Dibang Valley
district of Arunachal Pradesh in the extreme northeastern Himalayan region of India, close to China (Rathor et al.
2013).
Distribution
Stingless bees are most abundant in the southern parts of India and along the coast in the Bay of Bengal but can
also be locally common elsewhere (map 3). The interior plains of India are the least favorable areas for stingless
bees. Apparently stingless bees are also to be expected in the southern parts of Pakistan, Nepal, Bhutan, and most
of Bangladesh. Stingless bees of the Indian subcontinent represent the northernmost distributed stingless bees,
globally, although the precise distribution in the region is only fragmentarily known. Inhabiting such northern
latitudes for a social insect with limited adaption to cold conditions for extended periods, even below freezing for
multiple days, should motivate new behavioral and physiological studies to be conducted in the northern parts of
India.
MAP 3. Stingless bees can be expected from most parts of the Indian subcontinent. The darkest green represents areas with the
highest probability of habitat suitability and lighter shows lower probabilities of habitat suitability. The potential distribution of
stingless bees is based on localities from map 1 and Maxent (see text for details).
Discussion
The full diversity of stingless bees in the Indian subcontinent remains unknown. As a first step towards such
knowledge all known information about the name-bearing type specimens from the region is here summarized.
However, species of the “iridipennis” species group are extremely similar in external morphology of the workers,
and a taxonomic revision of the species of India should include morphological characters of the male genitalia and
molecular data. It is clear from Rasmussen & Cameron (2007, 2010), that at least four species from the
“iridipennis” group are found in India, but until their males are discovered, further collections are made, and
already described species from southeast Asia are compared with Indian specimens, it remains premature to
describe and propose additional species of Tetragonula from India.
Based on the distribution in Sri Lanka, I tend to favor the synonymy of T. praeterita with T. iridipennis, but
because of the shrunken eyes (and reduced head width) of the type specimen of T. praeterita, in addition to the glue
and worn or reduced pubescence, I tentatively leave the name as a separate species in recognition of several minor
differences (table 1 and 2). Even in Sri Lanka at least three species of stingless bees are found, including possibly
two species of Tetragonula (W.A.I.P. Karunaratne, pers. comm.) and the rarely observed Lisotrigona cacciae (as
first reported by Sakagami et al. 1990). Tetragonula ruficornis appears to be the best defined species based on the
inner surface of hind basitarsus (table 2) and shorter wing (table 1) and T. bengalensis is probably also distinct
(table 1 and 2), as suggested by Sakagami (1978).
In addition to the eight species here listed, more species are expected to occur in India. Some of those will be
species new to science, whereas others will represent the rich stingless bee fauna already known from Myanmar,
China, Thailand, Laos, Vietnam, Cambodia and Malaysia. Undoubtedly, new country records should be sought in
the NE areas of the Indian subcontinent, where closer contact with more diverse faunas of other countries could
lead to the discovery of extended species ranges. The present paper highlights some gaps in our current knowledge
of the diversity of stingless bees from the Indian subcontinent and it is hoped it will inspire new approaches,
collaborators and efforts to understand these bees.
Acknowledgment
Charles Michener, Victor Gonzalez, Deborah Smith, David Roubik, and Arkadiy Lelej provided many very helpful
comments that improved the manuscript and Gonzalez further assisted in locating many of the references with
additional citations supplied by Aluri Raju, Inoka Karunaratne, Pavithra Nayak, Raju Vyas, S. Basavarajappa, and
Shashidhar Viraktamath. Vikram S. Rathor, M. Muthuraman, and S. Viraktamath discussed the identity of Indian
stingless bees. David Notton was extremely helpful in locating all type specimens from BMNH, including
additional information on the provenance from records kept at the museum, while James Hogan kindly located the
type specimens of T. bengalensis in OXUM and Brian Harris the type specimen of T. pagdeni from USNM.
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APPENDIX 1.
List of all references searched for occurrence records of stingless bees from the Indian subcontinent. References were located
by database searches in Zoological Record, Scopus and Web of Science. In addition Rasmussen (2008), scholar.google.com,
and books.google.com were searched. In all searches the taxon name as “stingless bees”, Trigona, Tetragonula, Lisotrigona,
Lepidotrigona or Melipona was used in combination with each of the countries (India, Pakistan, Bangladesh, Sri Lanka (also as
Ceylon), Nepal, and Bhutan) or with each of the different species epithets. The website www.biodiversitylibrary.org was also
searched by taxon name.
Smith 1854, Tennent 1859, 1861, Motschoulsky 1863, Castets 1893, Bingham 1896, Dalla Torre 1896, Bingham 1897,
Cameron 1897, Nurse 1907, Strand 1913a, Dover 1925, George 1933, 1934, Pugh 1947, Nogueira-Neto 1949, 1951, Lindauer
1956, Lindauer & Kerr 1958, 1960, Percy et al. 1968, Phadke 1968, Jain & Kushwaha 1972, Kareem et al. 1976, Batra 1977,
Kshirsagar & Chauhan 1977, Sakagami 1978, Koeniger & Vorvohl 1979, Divan 1981, Pande & Sumita 1987a,b, Bichee &
Sharma 1988, Mohana Rao 1988, Kumar & Kumar 1989, Percy 1989, Raju & Reddi 1989a,b, Mohana Rao & Suryanarayana
1989, Goel & Kumar 1990, Raju 1990a, 1990b, Prasad & Hemanth 1992, Mohana Rao & Suryanarayana 1992, Prakash et al.
1993, Raju & Reddi 1993, Sadakathulla 1993, Ramanujam et al. 1993, Taori & Chakravarty 1994, Reddi et al. 1995a,b, Raju &
Reddi 1996a,b, Reddi et al. 1996, Islam 1997, Rama Das et al. 1997, Reddi & Raju 1997, Reddi et al. 1997, Joshi et al. 1998,
Raju et al. 1998, Partap 1999, Viraktamath et al. 1999, Lakshmi & Suryanarayana 1999, Rao et al. 1999, Atluri et al. 2000,
Chaudhary & Kumar 2000, Mohan & Devanesan 2000, Raju et al. 2000, Koshy et al. 2001, Nair & Nair 2001, Raju et al. 2001,
Viraktamath et al. 2001, Wijesekara 2001, Devanesan et al. 2002, Jagadish et al. 2002, Karunaratne & Edirisinghe 2002,
Kuberappa et al. 2002, Kusumawathie & Edirisinghe 2002, Raju & Ezradanam 2002, Thomas et al. 2002, Devanesan et al.
2003, Gurung et al. 2003, Nair 2003, Jobiraj & Narendran 2004, Liyanage & Edirisinghe 2004, Muthuraman & Saravanan
2004, Rangaiah et al. 2004, Sharma 2004, Eswarappa et al. 2005a,b,c, Gajanan et al. 2005, Karunaratne 2005, Karunaratne et
al. 2005a,b, Kuberappa et al. 2005, Raju et al. 2005, Rao et al. 2005, Raju & Rao 2006a,b, Raju et al. 2006, Chaudhary &
Singh 2007, Danaraddi 2007, Hannan 2007, Rasmussen & Cameron 2007, Sinu & Shivanna 2007, Sivaram 2007, Atluri et al.
2008, Karunaratne & Edirisinghe 2008a,b, Raju et al. 2008, Danaraddi & Viraktamath 2009, Danaraddi et al. 2009, Kulloli &
Sreekala 2009, Kulloli et al. 2009, Kuriakose et al. 2009, Raju et al. 2009a,b, Sheetal & Basavarajappa 2009, Stanley et al.
2009, Thomas et al. 2009, Basavarajappa 2010, Mupade & Kulkarni 2010, Punekar & Kumaran 2010, Rashmi et al. 2010,
Sharma et al. 2010, Danaraddi et al. 2011, Gupta et al. 2011, Jadhav et al. 2011, Kulloli et al. 2011, Raju et al. 2011a,b,
Ramasubbu et al. 2011, Regupathy & Ayyasamy 2011, Singh et al. 2011, Vyas 2011, Aswani & Sabu 2012, Choudhari et al.
2012, Danaraddi et al. 2012, Gorain et al. 2012, Kumar et al. 2012, Lepcha et al. 2012, Mohapatra & Sontakke 2012, National
Horticulture Mission 2012, Prasad & Sunoj 2012, Raju et al. 2012a,b, Surendra et al. 2012, Vyas 2012, Wanigasekara &
Karunaratne 2012, Nayak et al. 2013.
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Stingless bees (Hymenoptera: Apidae: Meliponini) of the Indian subcontinent:

Article in Zootaxa · May 2013

Stingless bees in Argentina View project

Native Danish bees - also in the future View project

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Stingless bees (Hymenoptera: Apidae: Meliponini) of the Indian subcontinent:

Accepted by A. Lelej: 3 Apr. 2013; published: 10 May 2013 401

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Material and methods

Lepidotrigona arcifera (Cockerell 1929)

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Lisotrigona cacciae (Nurse 1907)

Lisotrigona mohandasi Jobiraj & Narendran 2004

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Tetragonula iridipennis (Smith 1854)

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Tetragonula ruficornis (Smith in 1870)

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Tetragonula bengalensis (Cameron 1897)

Tetragonula species of the “laeviceps” species group

Tetragonula aff. laeviceps (Smith 1857)

416 · Zootaxa 3647 (3) © 2013 Magnolia Press RASMUSSEN

Tetragonula species of the “gressitti” species group

Tetragonula gressitti (Sakagami 1978)

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428 · Zootaxa 3647 (3) © 2013 Magnolia Press RASMUSSEN

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