studies on tribe gymnopleurini (coleoptera: scarabaeidae: scarabaeinae) from madhya pradesh and...
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RESEARCH ARTICLE
Studies on Tribe Gymnopleurini (Coleoptera: Scarabaeidae:Scarabaeinae) from Madhya Pradesh and Chhattisgarhwith A Checklist from India
Kailash Chandra • Devanshu Gupta
Received: 26 November 2012 / Revised: 5 June 2013 / Accepted: 16 July 2013
� The National Academy of Sciences, India 2013
Abstract The dung beetle tribe Gymnopleurini (Cole-
optera: Scarabaeidae: Scarabaeinae) comprise approxi-
mately 110 ball-rolling species are described within four
genera; Allogymnopleurus, Garreta, Gymnopleurus and
Paragymnopleurus. The composition of the tribe in Mad-
hya Pradesh and Chhattisgarh consisted of eight species
within three genera. Garreta smaragdifer (Walker) is
reported for the first time from both the states. The tax-
onomy, descriptions, distribution, and locality collection
data is provided for all species. Male genitalia and sexual
dimorphism of seven species except Gymnopleurus
(Metagymnopleurus) parvus MacLeay are described and
illustrated for the first time. An identification key for all
Gymnopleurini species that occur in Madhya Pradesh and
Chhattisgarh is provided along with illustrations. A
checklist pertaining 21 species distributed in all the four
recognized genera of the tribe is also prepared.
Keywords Dung beetles � Gymnopleurini � Checklist �Male genitalia � Sexual dimorphism
Introduction
The superfamily Scarabaeoidea, considered as a mono-
phyletic group [1], includes 12 families, 43 subfamilies,
118 tribes, and 94 subtribes, recognized as valid taxa [2].
This diverse scarab group has been divided in two eco-
logical groups [3]; Laparosticti (dung beetles) are charac-
terized by having most of the abdominal spiracles located
on the pleural membrane between the tergites and sternites
and the Pleurosticti (leaf-feeding beetles or leaf chafers)
characterized by having most of the spiracles located on
the upper portion of the sternites. The two groups are
commonly included in the Scarabaeidae, the biggest and
most diverse family within Scarabaeoidea. The family
Scarabaeidae includes 27,800 species worldwide, and
within Scarabaeidae, the two subfamilies; Aphodiinae and
Scarabaeinae are represented by approximately 6,850
species worldwide and the subfamilies; Orphninae, Melo-
lonthinae, Dynastinae, Rutelinae, Cetoniinae, Trichiinae
and Valginae include approximately 20,950 species [4].
The dung beetle tribe Gymnopleurini within the sub-
family Scarabaeinae of family Scarabaeidae, comprise
approximately 110 ball-rolling species, described within
four relatively species-rich genera; Allogymnopleurus,
Garreta, Gymnopleurus and Paragymnopleurus [5]. The
genus Paragymnopleurus shows a primarily Oriental dis-
tribution while the other three genera show Afro-Eurasian
(Gymnopleurus) or Afro-Oriental distributions (Allogym-
nopleurus, Garreta), and show diurnal flight activity.
Adults of the tribe are characterized by an emargination of
the lateral edge of each elytron that exposes the underlying
pleural sclerites. The middle and hind tibiae are mostly
long and thin for manipulating the dung balls whereas the
fore tibiae bear a terminal spur, four tibial teeth, and tarsi.
Separation of the genera is based mainly on metathoracic
epimera, clypeal dentition, canthi projection, abdominal
and fore femur carina and mid tibial spurs.
Depending on the geographical variation in the climate,
seasonal range and breeding, the species show variable
K. Chandra
Zoological Survey of India, New Alipore,
Kolkata 700053, West Bengal, India
e-mail: [email protected]
D. Gupta (&)
Central Zone Regional Center, Zoological Survey of India,
Vijay Nagar, Jabalpur 482002, Madhya Pradesh, India
e-mail: [email protected]
123
Proc. Natl. Acad. Sci., India, Sect. B Biol. Sci.
DOI 10.1007/s40011-013-0229-z
colouration, ranging from copper, green, blue, violet and
non-metallic shiny black [6]. Studies on nesting behavior
suggest that a dung ball intended for brood construction
is rolled away from the dung, possibly by a single beetle,
or by a pair [7] and buried at the end of a short tunnel in
the soil where a depression is excavated in the ball.
Thereafter an egg is laid at its base, and dung is pulled
over the egg to enclose it within an egg chamber, thus
forming a brood which is then coated with a mixture of
soil and dung to prevent fungal attack [5]. The shape of
the brood also varies between genera, being oval
in Garreta and Allogymnopleurus but pear-shaped in
Gymnopleurus [5, 8].
Review of literature on taxonomic studies of Gymno-
pleurini from India revealed following monographic, fau-
nal and catalogues made by Arrow [8], Janssens [9],
Balthasar, [10], Gupta [11], Biswas and Chatterjee [12],
Biswas and Ghosh [13], Chandra [14, 15], Bezdek [16],
Kabakov [17], Sewak [18, 19], Chandra and Ahirwar [20];
Chatterjee [21], Chandra and Singh [22], Mittal [23, 24],
Sabu et al. [25] and Chandra et al. [26]. The first com-
prehensive study on tribe Gymnopleurini of India was
carried out by Arrow [8], in the fauna of British India
devoted to Coprinae, wherein 20 spp. were studied and
kept in the genus Gymnopleurus. Later Janssens [9] revised
all the species of the tribe, and kept them into four cur-
rently recognized genera of the tribe viz. Allogymnopleu-
rus, Garreta, Gymnopleurus and Paragymnopleurus so for
known from the world. Further, in his monograph on the
Plaearctic and Oriental region, Balthasar [10] included
these genera as subgenera of single genus Gymnopleurus.
As far, the studies on the tribe in central India are con-
cerned, Chandra and Ahirwar [20], listed 10 spp. in the
genus Gymnopleurus from Madhya Pradesh and Chhattis-
garh. The importance of morphotaxonomy and external
genitalia has been discussed by Kirti and Gill [27].
Although from a taxonomic standpoint the tribe Gymno-
pleurini is comparatively well known but information on
the distribution, male genitalia, sexual dimorphism, and
ecology of the majority of the species from India is either
minimal or completely lacking. The study presents new
distributional and ecological data for all the species of
Gymnopleurini recorded from the states of Madhya Pra-
desh and Chhattisgarh. Compiling all the previously pub-
lished information on the group [8–26], the paper also
includes a species checklist of Gymnopleurini from India
giving geographical distribution of all the species reported
to date from the country.
Fig. 1 Map showing collection
localities
K. Chandra, D. Gupta
123
Material and Methods
Study Area
Madhya Pradesh with its capital in Bhopal and Chhattis-
garh with its capital in Raipur, occupy the very heartland of
India and together they form the unit of Central India. The
total area of these two states is 443,446 sq. km lying
between latitudes 21�–25� N and longitudes 74�–84� E,
covering about 14.5 % of the total area of India. The two
states fall under two bio-geographic zones (Zone four:
Semi Arid and Zone six: Deccan Plateau) [28] and have
18 forest types which belong to the three forest type
groups viz. tropical dry deciduous, tropical moist decidu-
ous, and tropical thorn forests [29]. Madhya Pradesh has
nine national parks, 25 wildlife sanctuaries, one biosphere
Fig. 2 Key characters: Lateral carina of a Gymnopleurus gemmatus, b Garreta mundus; Heads of c Gymnopleurus gemmatus, d G. cyaneus,
e Paragymnopleurus sinuatus ssp. assamensis, and f Garreta mundus; Pronotum of g Gymnopleurus cyaneus, h G. miliaris, i G. gemmatus,
j G. parvus, k Paragymnopleurus sinuatus ssp. assamensis, l Garreta smaragdifer, m G. mundus, and n G. dejeani
Studies on Tribe Gymnopleurini
123
reserve and five tiger reserves, while Chhattisgarh is
attributed with three national parks, ten wildlife sanctuar-
ies, one biosphere reserve and one tiger reserve.
The material for the present study was collected by the
scientific teams of Zoological Survey of India (ZSI), from
different localities and protected areas of Madhya Pradesh
and Chhattisgarh, during 1999–2011. A total of 12 districts
(Bhind, Jabalpur, Damoh, Hoshangabad, Seoni, Chhindw-
ara, Balaghat, Mandla, Umaria, Sidhi, Bilaspur and Raipur)
and ten protected areas [Pench Tiger Reserve (PTR),
Pachmarhi Biosphere Reserve (PBR), Veeragnagana Dur-
gavti Wildlife Sanctuary (VDWS), Bandhavgarh National
Park (BNP), Kanha National Park (KNP), Kanha Tiger
Reserve (KTR), Madhav National Park (MNP), Achanak-
mar-Amarkantak Biosphere Reserve (AABR), Barnawa-
para Wildlife Sanctuary (BWLS) and Bori Wildlife
Sanctuary (BWS)], were surveyed (Fig. 1).
Light trapping and handpicking methods were used for
adult beetle sampling. After collection, beetles were killed
in Benzene vapors and preserved dry pinned. The identi-
fication of the specimens was made with the help of rele-
vant literature [8–10]. For the study of male genitalia, male
specimen was dissected out and abdomen was separated
from the body and aedeagus was extracted from the
abdomen. The genitalia was then cleaned and softened in a
dish of hot water and further cleaned in a hot water solution
of 10 % KOH. If not damaged, the abdomen was again
glued to the specimen. The aedeagus was washed in 95 %
ethanol and stored in a glass vial containing 70 % alcohol.
Voucher specimens were deposited in the museum of ZSI,
Jabalpur, Madhya Pradesh (India).
For each species complete synonymies are provided.
Brief descriptions, information on sexual dimorphism,
male genitalia and geographical distributions, are inclu-
ded. An identification key for all the species from MP
and CH is presented. The keys and descriptions of each
species are accompanied by the illustrations made with
the use of digital camera attached to Leica M205A,
stereomicroscope.
Results and Discussion
Taxonomic Account
Order: Coleoptera Linnaeus, 1758.
Suborder: Polyphaga Emery, 1886.
Family: Scarabaeidae Latreille, 1802.
Subfamily: Scarabaeinae Latreille, 1802.
Tribe Gymnopleurini Lacordaire, 1856.
Identification key to genera and species of Gymnopleurini from MP and CH
1. Sides of abdomen not carinate at base (Fig. 2a)……………Gymnopleurus Illiger….....……….….…..2
Sides of abdomen carinate at base (Fig. 2b)……...…….………….……………………….………...….5
2. Clypeus quadric-dentate (Fig. 2d).………………………………….…………….…..……………….…3
Clypeus bi-dentate (Fig. 2c)...……………………………….………………...…………….……..……4
3. Upper surface without hair; pronotum convex, smooth unevenly and very strongly punctured (Fig. 2g)
…………….……………………………………………..…………..Gymnopleurus cyaneus (Fabricius)
Upper surface clothed with fine hair; pronotum granular, with a spot in middle and five spots
surrounding central spot (Fig. 2h)………….....Gymnopleurus (Metagymnopleurus) miliaris (Fabricius)
4. Pronotum with 15 shining spots (Fig. 2i)…........Gymnopleurus (Metagymnopleurus) gemmatus Harold
Pronotum with 6 spots (Fig. 2j)…………………Gymnopleurus (Metagymnopleurus) parvus MacLeay
5. Clypeus in front distinctly bi-dentate; (Fig. 2e); pronotum angulate in middle (Fig. 2k)
…........................................Paragymnopleurus Shipp….………...Paragymnopleurus sinuatus (Olivier)
Clypeus in front distinctly quadri-dentate (Fig. 2f); pronotum rounded in middle (Fig. 2 l, m, n)
…….....…………………………………………………………………………...Garreta Janssens..….6
6. Upper surface shining; pronotum very lightly punctured (Fig. 2l)…….…Garreta smaragdifer Walker
Upper surface not shining; pronotum punctured or granular……………………………………...….…7
7. Pronotum finely and distinctly punctured (Fig. 2m)…………………......Garreta mundus (Wiedemann)
Pronotum without punctures but distinctly granular throughout (Fig. 2n)…Garreta dejeani (Castelnau)
K. Chandra, D. Gupta
123
Genus Gymnopleurus Illiger, 1803
Gymnopleurus cyaneus (Fabricius, 1798) (Figs. 2d, g, 3a,
5a, b, 7a)
Copris cyaneus, Fabricius, 1798, Ent. Syst. Suppl.: 34.
Gymnopleurus cyaneus, Arrow, 1931, Faun. Brit. India,
(Lamellicornia: Coprinae), III: 49, pl. 3, f. 6.
Gymnopleurus cyaneus, Janssens, 1940, Verh. Kon. Nat.
Mus. Belg., 2 (18): 66, pl. II, Fig. 9.
Gymnopleurus (Gymnopleurus) cyaneus, Balthasar, 1963,
Mon. Scarab. Aphod. Palae. Ori. Reg., (Coleoptera:
Lamellicornia), Coprinae, I: 207.
Gymnopleurus indicus, Castelnau, 1840, Hist. Nat. Col., II:
73.
Gymnopleurus impressus, Castelnau, 1840, Hist. Nat. Col.,
II: 73.
Collection data: 104 specimens-53 male: 51 female.
Temporal distribution (all data): May (5), June (65), July (34);
Madhya Pradesh: Hoshangabad: BWS, Churna 11.vi.2009
(1#1$), 14.vi.2009 (1#1$), 10.vi.2009 (10#10#); coll.
K. Chandra. Seoni: PTR, Chhindimata, Alikatta, 29.vii.2001
(1$), 31.vii.2001 (7#3$); coll. M. L. Koshta; Chhindmata,
25.vii.2001 (1#3$); Karmajhiri, 01.vii.2001 (1#); coll.
K. Chandra. Sidhi: Bansagar canal project, Mohania,
22.vii.1999 (3#5$), coll. K. Chandra. Jabalpur: railway col-
ony, 14.vi.2004 (1#), 21.vi.2004 (2$), 22.vi.2004 (2$),
24.vi.2004 (3$1#), night collection 07.vii.2003 (2$); coll.
Subhash; ZSI colony, 17.ix.2008 (1#), coll. Rajesh.
Chhattisgarh: Bilaspur: AABR, day collection, Manjhi-
dongri, 07.vi.2004 (3#3$), 08.vi.2004 (4#4$); Atariya
Forest, 17.vi.2004 (2#), 18.vi.2004 (3#2$), 19.vi.2004
(2#2$), 27.vi.2004 (2#1$); Kotani Forest, 22.vii.2004
(1$); Lamni Forest, 07.v.2005 (1#2$); Tilaidobra, Atariya
Forest, 11.v.2005 (1#), 15.v.2005 (1#); coll. A. Singh;
Amarkantak, 05.vi.2008 (1#), 10.vi.2008 (1#); Lamni
Forest, 12.vi.2008 (2#1$); coll. Y. N. Gupta; Sarsadol
Forest, 18.vii.2005 (1#1$), coll. A. Singh. Raipur: BWLS,
Tinsa Pathar, 03.vii.2011 (2#1$), coll. K. Chandra.
Fig. 4 a Gymnopleurus miliaris
(Cooperation of # and $ beetles
in rolling dung ball),
b Paragymnopleurus sinuatus
ssp. assamensis ($ at light)
Fig. 3 Adult photographs: a Gymnopleurus cyaneus (#), b G. miliaris (#), c G. gemmatus ($), and d G. parvus ($)
Studies on Tribe Gymnopleurini
123
Diagnosis: (Fig. 3a). Bright, shining, metallic green or
blue; broad, oval and not very convex; clypeus quadri-
dentate (Fig. 2d); pronotum short, convex, strongly but
sparsely punctured (Fig. 2g); elytra with a series of large
and confluent pits.
Sexual dimorphism: Male: Spur of pro tibia flat, truncate
at end and strongly bent downwards (Fig. 5b). Female:
Spur of pro tibia slender and acute (Fig. 5a).
External male genitalia: (Fig. 7a). Phallobase longer
than parameres; parameres broad at base and irregularly
tapered distally with the tips pointed.
Remarks: Attracted towards light.
Gymnopleurus (Metagymnopleurus) miliaris (Fabricius,
1775) (Figures 2h, 3b, 4a, 5c, d, 7b)
Scarabaeus miliaris, Fabricius, 1775, Syst. Ent. App.: 817.
Gymnopleurus miliaris, Arrow, 1931, Faun. Brit. India,
(Lamellicornia: Coprinae), III: 50, Pl. II, Fig. 3.
Gymnopleurus miliaris, Janssens, 1940, Verh. Kon. Nat.
Mus. Belg., 2 (18): 66, Pl. II, Fig. 12.
Gymnopleurus (Gymnopleurus) miliaris, Balthasar, 1963,
Mon. Scarab. Aphod. Palae. Ori. Reg., (Coleoptera:
Lamellicornia), Coprinae, I: 207, pl. IX, Fig. 1.
Gymnopleurus (Metagymnopleurus) miliaris, Kabakob,
2006, Beet. subf. Scarab. Faun. Russ. adj. Coun.: 79.
Collection data: eight specimens: four male: four female.
Temporal distribution (all data): June (4), July (4);
Madhya Pradesh: Jabalpur: railway hostel, 16.vi.2004
(1$), 30.vi.2004 (3#), railway colony, 04.vii.2004 (1$),
08.vii.2004 (1$), coll. Subhash.
Chhattisgarh: Raipur: BWLS, 01.vii.2011 (1#1$), coll.
K. Chandra.
Diagnosis: (Fig. 3b). Blue black; broad not very convex;
dorsally clothed with minute grey Setae; pronotum bearing
a few shining spots; a central spot surrounded by a ring of
five spots; an outer spot on each side and lateral pit on each
side near last (Fig. 2h); elytra also bears following spots;
two spots adjoining basal margin of basal and apical parts;
one near middle of suture on each side, an outer one close
to each of last and upon apical callus.
Sexual dimorphism: Male: Pro tibia little longer and less
broad than that of female, with shorter exterior teeth and its
terminal spur very blunt and little bent (Fig 5c). Female:
Terminal spur of pro tibia strongly curved and very sharp at
end (Fig. 5d).
External male genitalia: (Fig. 7b). Phallobase longer
than parameres; parameres almost triangular in shape with
its tip having a fringe of hair.
Gymnopleurus (Metagymnopleurus) gemmatus Harold,
1871 (Figures 2a, c, i, 3c, 5e, f, 7c)
Gymnopleurus gemmatus, Harold, 1871, Col. Hefte, VIII: 117.
Gymnopleurus gemmatus, Arrow, 1931, Faun. Brit. India,
(Lamellicornia: Coprinae), III: 53, pl. III, Fig. 1. Gymno-
pleurus gemmatus, Janssens, 1940, Verh. Kon. Nat. Mus.
Belg., 2 (18): 62.
Gymnopleurus (Gymnopleurus) gemmatus, Balthasar,
1963, Mon. Scarab. Aphod. Palae. Ori. Reg., (Coleoptera:
Lamellicornia), Coprinae, I: 204, pl. VIII, Fig. 2.
Gymnopleurus (Metagymnopleurus) gemmatus, Kabakob,
2006, Beet. subf. Scarab. Faun. Russ. adj. Coun.: 79.
Scarabaeus granulatus, Fabricius (nec Olivier), 1792, Ent.
Syst.: 65.
Collection data: 12 specimens- 6 male: 6 female. Tem-
poral distribution (all data): April (2) June (9), July (1);
Madhya Pradesh: Jabalpur: railway hostel, 14.vi.2004
(1#1$), 17.vi.2004 (1#1$), 20.vi.2004 (1$), 21.vi.2004
(1$), 22.vi.2004 (1#), 30.vi.2004 (1#1$); coll. Subhash.
Bhind: 28.iv.2011 (1#1$), coll. Sandeep and Devanshu.
Fig. 5 Pro tibia of: a Gymnopleurus cyaneus ($), b G. cyaneus (#),
c G. miliaris (#), d G. miliaris ($), e G. gemmatus (#), f G. gemmatus
($), g Garreta dejeani (#), and h G. dejeani ($)
K. Chandra, D. Gupta
123
Chhattisgarh: Raipur: Barnawapara Wildlife Sanctuary,
Tinsa Pathar, 03.vii.2011 (1#), coll. K. Chandra.
Diagnosis: (Fig. 3c). Black dorsally clothed with minute
grey setae, leaving various shining spots; clypeus bi-den-
tate (Fig. 2c); pronotum with following spots; a central
spot, five adjoining front margin, four adjoining hind
margin, three and or four on each side (Fig. 2i); elytra also
bears bands; a common transverse irregular patch at middle
and a smaller common patch behind it just before apices,
three spots adjoining basal margin of each side, one near
middle of outer margin and one upon apical callus.
Sexual dimorphism: Male: Pro tibia longer and nar-
rower, a little bent before end, feebly serrate internally
and armed with three shorter teeth externally; terminal
spur bent at end and rather blunt (Fig. 5e). Female: Pro
tibia broader and armed with three longer teeth externally
while front tibial spur slender curved and very acute
(Fig. 5f).
External male genitalia: (Fig. 7c). Phallobase longer
than parameres; parameres slightly tapers distally, apical
part slightly curved inward with tips blunt and having a
fringe of short hair.
Fig. 6 Pronotum and pro legs:
a Paragymnopleurus sinuatus
ssp. assamensis (#),
b P. sinuatus ssp. assamensis ($),
c Garreta smaragdifer (#),
d G. smaragdifer ($),
e G. mundus (#), and
f G. mundus ($)
Studies on Tribe Gymnopleurini
123
Gymnopleurus (Metagymnopleurus) parvus MacLeay,
1821 (Figures 2j, 3d)
Gymnopleurus parvus, MacLeay, 1821, Horae Ent., I, 2: 517.
Gymnopleurus parvus, Arrow, 1931, Faun. Brit. India,
(Lamellicornia: Coprinae), III: 52, pl. III, Fig. 2.
Gymnopleurus parvus, Janssens, 1940, Ver. Kon. Nat. Mus.
Bel., 2 (18): 62, pl. II, Fig. 10.
Gymnopleurus (Gymnopleurus) parvus, Balthasar, 1963,
Mon. Scarab. Aphod. Palae. Ori. Reg., (Coleoptera:
Lamellicornia), Coprinae, I: 202.
Gymnopleurus (Metagymnopleurus) parvus, Kabakob,
2006, Beet. subf. Scarab. Faun. Russ. adj. Coun.: 79.
Collection data: 2$ specimens. Temporal distribution
(all data): June (1), July (1);
Fig. 7 External male genitalia of: a Gymnopleurus cyaneus, b G. miliaris, c G. gemmatus, d Paragymnopleurus sinuatus ssp. assamensis,
e Garreta smaragdifer, f G. mundus
K. Chandra, D. Gupta
123
Madhya Pradesh: Sidhi: Bansagar canal project, Moha-
nia, 27.vii.1999 (1$); Hoshangabad: PBR, Karkroi village,
11.vi.1999 (1$), coll. K. Chandra.
Diagnosis: (Fig. 3d). Dark greenish black; oval and
moderately convex; dorsally clothed with minute grayish
setae, leaving a few shining spots on pronotum and elytra;
three placed in a transverse row along middle line of
pronotum (Fig. 2j), one between median spot and base, and
one on each side between last and lateral spot, a common
spot at base of elytral suture, three along basal margin, one
on each side of suture a little behind middle, and one upon
each apical callus; posterior part of sutural margin also bare
and shining.
Genus Paragymnopleurus Shipp, 1897
Paragymnopleurus sinuatus ssp. assamensis (Waterhouse,
1890) (Figures 2e, k, 4b, 6a, b, 7d)
Gymnopleurus assamensis, Waterhouse, 1890, Ann. Mag.
Nat. Hist., (6), V: 411.
Gymnopleurus sinuatus var. assamensis, Arrow, 1931,
Faun. Brit. India, (Lamellicornia: Coprinae), III: 63.
Paragymnopleurus sinuatus, Janssens, 1940, Ver. Kon.
Nat. Mus. Bel., 2 (18): 17, 20, pl. I, Fig. 1.
Gymnopleurus (Paragymnopleurus) sinuatus, Balthasar,
1963, Mon. Scarab. Aphod. Palae. Ori. Reg., (Coleoptera:
Lamellicornia), Coprinae, I: 218, pl. X, Fig. 2.
Collection data: 36 specimens: 21 male: 15 female.
Temporal distribution (all data): June (15), July (12),
August (2), September (5), November (2);
Madhya Pradesh: Seoni, PTR: Rukhar, 12.vii.2001 (1$);
Alikatta, light trap, 07.vii.2001 (1#); Karmajhiri,
14.vii.2001 (1$); coll. R. K. Singh; Kalaphar, light trap,
11.vii.2001 (1$); Karmajhiri, 05.vi.2001 (3$), 06.vi.2001
(2$1#); 08.vi.2001 (2#1$); Turiya, 23.vi.2001 (1#); coll.
K. Chandra; Bolanala, light trap, 23.vii.2001 (1$1#);
Khamareet, 29.vii.2001 (1#); Chandrapur, 10.xi.2001
(1#); coll. M. L. Koshta; Chhindwara: Totaldoh Forest rest
house, 24.viii.2001 (1$), 26.viii.2001 (1#); 03.ix.2001
(1#), coll. Y. N. Gupta. Umaria: BNP, Tala guest house,
22.ix.2002 (2#), coll. M. L. Koshta. Mandla, KNP: Kisli
rest house, 14.ix.2002 (1#); Kanha, 19.ix.2003 (1#), coll.
M. Limje. Balaghat: Kanha Tiger Reserve: Mukki rest
house, 22.vii.2004 (1#), coll. Rajendra Gupta. Shivpuri:
Madhav National Park, landing no. 1, 23.xi.2001 (1#), coll.
Nema and Ahirwar. Damoh: VDWS, Bhaisaghat,
22.vii.2009 (1$), coll. K. Chandra.
Chhattisgarh: Bilaspur: AABR, Atariya, light trap,
17.vi.2004 (1$1#), 18.vi.2001 (1#); Achanakmar wildlife
sanctuary, Chhaparwa rest house, light trap, 03.vi.2004 (#),
14.vi.2004 (1#), 18.vii.2005 (1$); coll. A. Singh. Raipur:
BWLS, Kari Dongri forest, 04.vii.2011 (1$1#), coll.
K. Chandra.
Diagnosis: Opaque, dark red-coppery with antennal clubs
bright yellow; broad, oval not very convex; clypeus with
anterior margin bi-lobed (Fig. 2e); pronotum finely rather
closely punctured (Fig. 2k); elytra with intervals flat and
coriaceous.
Sexual dimorphism: Male: hind angles of pronotum with
slightly produced lobes; pro tibia a little elongate, with
slight internal lobe at end and terminal spur truncate and
bi-dentate (Fig. 6a). Female: Pro tibia a little broader and
its terminal spur slender and blunt (Fig. 6b).
External male genitalia: (Fig. 7d). Phallobase longer
than parameres; parameres broad at base and tapered dis-
tally with a cup shaped cavity sub apically on dorsal side
and tips blunt.
Remarks: Attracted towards light.
Genus Garreta Janssens, 1940
Garreta smaragdifer (Walker, 1858) (Figures 2l, 6c, d, 7e)
Gymnopleurus smaragdifer, Walker, 1858, Ann. Mag. Nat.
Hist., (3), II: 208.
Gymnopleurus smaragdifer, Arrow, 1931, Faun. Brit.
India, (Lamellicornia: Coprinae), III: 60, pl. III, Fig. 13.
Garreta smaragdifer, Janssens, 1940, Verh. Kon. Nat. Mus.
Belg., 2 (18): 24, 29, pl. I, Fig. 2.
Gymnopleurus (Garreta) smaragdifer, Balthasar, 1963,
Mon. Scarab. Aphod. Palae. Ori. Reg., (Coleoptera:
Lamellicornia), Coprinae, I: 226.
Collection data: Seven specimens- six male: one female.
Temporal distribution (all data): June (4), July (1), August
(1), September (1);
Fig. 8 External male genitalia of: a Garreta dejeani
Studies on Tribe Gymnopleurini
123
Ta
ble
1C
hec
kli
sto
ftr
ibe
gy
mn
op
leu
rin
ifr
om
Ind
ia
Nam
eo
fth
esp
ecie
sG
eog
rap
hic
ald
istr
ibu
tio
n
Ind
ia(S
tate
s)M
adh
ya
Pra
des
h
(dis
tric
tsan
d
pro
tect
edar
eas)
Ch
hat
tisg
arh
(dis
tric
tsan
d
pro
tect
edar
eas)
Els
ewh
ere
I.G
enu
sA
llo
gym
no
ple
uru
sJa
nss
ens,
19
40
:9
( Ty
pe
spec
ies-
Gym
no
ple
uru
sch
lori
sK
lug
,1
85
5)
1.
All
og
ymn
op
leu
rus
ma
culo
sus
(Mac
Lea
y,
18
21
)
Ass
am,
Mah
aras
htr
a,S
ou
thIn
dia
,
and
Utt
arak
han
d
––
Nep
al
2.
All
og
ymn
op
leu
rus
spil
otu
s
(Mac
Lea
y,
18
21
)
An
dh
raP
rad
esh
,B
ihar
,K
arn
atak
a,
and
Tam
ilN
adu
––
–
II.
Gen
us
Ga
rret
aJa
nss
ens,
19
40
:9
(Ty
pe
spec
ies:
Sca
rab
aeu
sa
zure
us
Fab
rici
us,
18
01
)
3.
Ga
rret
ad
ejea
ni
(Cas
teln
au,
18
40
)
Ch
hat
tisg
arh
,H
imac
hal
Pra
des
h,
Ker
ala,
Mad
hy
aP
rad
esh
,
Mah
aras
htr
aan
dT
amil
Nad
u
Dam
oh
;V
DW
SR
aip
ur;
BW
LS
–
4.
Ga
rret
ag
ille
ti(G
arre
ta,
19
14
)U
ttar
Pra
des
h–
–V
ietn
am
5.
Ga
rret
am
un
du
s(W
ied
eman
n,
18
19
)
Bih
ar,
Ch
hat
tisg
arh
and
Mad
hy
a
Pra
des
h
Ho
shan
gab
ad,
Seo
ni;
BW
S,
PT
RR
aip
ur;
BW
LS
So
uth
-Wes
tC
hin
a
6.
Ga
rret
ao
pa
cus
Red
ten
bac
her
,
18
48
Him
ach
alP
rad
esh
,Ja
mm
u&
Kas
hm
iran
dU
ttar
akh
and
.
––
Nep
al
7.
Ga
rret
aru
fico
rnis
(Mo
tsch
uls
ky
,1
85
4)
Ass
aman
dH
imac
hal
Pra
des
h–
–A
fgh
anis
tan
,C
hin
a
8.
Ga
rret
asm
ara
gd
ifer
(Wal
ker
,
18
58
)
Ch
hat
tisg
arh
,M
adh
ya
Pra
des
h
and
So
uth
Ind
ia
Seo
ni,
Ch
hin
dw
ara;
PT
RB
ilas
pu
r;A
AB
RS
riL
ank
a
9.
Ga
rret
asu
mp
tuo
sus
(Cas
teln
au,
18
40
)
Mah
aras
htr
aan
dT
amil
Nad
u–
––
10
.G
arr
eta
sylv
estr
isM
itta
l,
20
11
:2
97
Har
yan
a–
––
III.
Gen
us
Gym
no
ple
uru
sIl
lig
er,
18
03
:1
99
(Ty
pe
spec
ies-
Sca
rab
aeu
sfl
ag
ella
tes
Fab
rici
us,
17
87
)
11
.G
ymn
op
leu
rus
cya
neu
s
(Fab
rici
us,
17
98
)
Ch
hat
tisg
arh
,G
uja
rat,
Har
yan
a,
Kar
nat
aka,
Raj
asth
an,
Mah
aras
htr
a,
Mad
hy
aP
rad
esh
,O
riss
a,T
amil
Nad
u,
Utt
arak
han
d,
Utt
arP
rad
esh
and
Wes
tB
eng
al
Ho
shan
gab
ad,
Seo
ni,
Sid
hi,
and
Jab
alp
ur;
PB
R,
PT
R
Bil
asp
ur,
Rai
pu
r;
AA
BR
,B
WL
S
Ban
gla
des
h,
Nep
alan
d
Sri
Lan
ka
12
.G
ymn
op
leu
rus
bih
are
nsi
s
Arr
ow
,1
93
1
Bih
ar–
––
K. Chandra, D. Gupta
123
Ta
ble
1co
nti
nu
ed
Nam
eo
fth
esp
ecie
sG
eog
rap
hic
ald
istr
ibu
tio
n
Ind
ia(S
tate
s)M
adh
ya
Pra
des
h
(dis
tric
tsan
d
pro
tect
edar
eas)
Ch
hat
tisg
arh
(dis
tric
tsan
d
pro
tect
edar
eas)
Els
ewh
ere
13
.Gym
no
ple
uru
sb
om
ba
yen
sis
Arr
ow
,1
93
1
Mah
aras
htr
a–
–B
alu
chis
tan
14
.Gym
no
ple
uru
s
(Met
ag
ymn
op
leu
rus)
bic
all
osu
s
Fel
sch
e,1
90
9
Ass
aman
dR
ajas
than
––
Afg
han
ista
nan
dP
akis
tan
.
15
.Gym
no
ple
uru
s
(Met
ag
ymn
op
leu
rus)
gem
ma
tus
Har
old
,1
87
1
An
dh
raP
rad
esh
,B
ihar
,C
hh
atti
sgar
h,
Har
yan
a,K
arn
atak
a,K
eral
a,
Mah
aras
htr
a,M
adh
ya
Pra
des
h,
Ori
ssa,
Po
nd
ich
erry
,T
amil
Nad
u,
Utt
arak
han
dan
dU
ttar
Pra
des
h
Jab
alp
ur
and
Bh
ind
Rai
pu
r;B
WL
SA
fgh
anis
tan
,S
riL
ank
a
16
.Gym
no
ple
uru
s
(Met
ag
ymn
op
leu
rus)
hyp
ocr
ita
Bal
thas
ar,
19
60
Ori
ssa
––
–
17
.Gym
no
ple
uru
s
(Met
ag
ymn
op
leu
rus)
koen
igi
(Fab
rici
us,
17
75
)
An
dh
raP
rad
esh
,P
on
dic
her
ry,
Raj
asth
anan
dT
amil
Nad
u
––
Sri
Lan
ka
and
Ch
ina
18
.Gym
no
ple
uru
s
(Met
ag
ymn
op
leu
rus)
mil
iari
s
(Fab
rici
us,
17
75
)
Ch
hat
tisg
arh
,G
uja
rat,
Har
yan
a,
Him
ach
alP
rad
esh
,Ja
mm
u&
Kas
hm
ir,
Kar
nat
aka,
Mad
hy
a
Pra
des
h,
Mah
aras
htr
a,O
riss
a,
Raj
asth
an,
Tam
ilN
adu
,U
ttar
akh
and
and
Utt
arP
rad
esh
Jab
alp
ur
Rai
pu
r;B
WL
SA
fgh
anis
tan
,B
hu
tan
,
Pak
ista
nan
dS
riL
ank
a
19
.G
ymn
op
leu
rus
(Met
ag
ymn
op
leu
rus)
pa
rvu
s
Mac
Lea
y,
18
21
Bih
ar,
Har
yan
a,M
adh
ya
Pra
des
h,
Raj
asth
an,
Tam
ilN
adu
,U
ttar
akh
and
and
Utt
arP
rad
esh
Sid
hi,
Ho
shan
gab
ad;
PB
R–
Sri
Lan
ka
IV.
Gen
us
Pa
rag
ymn
op
leu
rus
Sh
ipp
,1
89
7:
16
6(T
yp
esp
ecie
s:S
cara
ba
eus
sin
ua
tus
Oli
vie
r,1
78
9)
20
.P
ara
gym
no
ple
uru
s
mel
an
ari
us
(Har
old
,1
86
7)
Tam
ilN
adu
and
Mah
aras
htr
a–
–S
riL
ank
a,C
hin
a,T
aiw
an,
My
anm
ar,
Vie
tnam
,L
aos,
Su
mat
raan
dJa
va
21
.P
ara
gym
no
ple
uru
ssi
nu
atu
s
ssp
.a
ssa
men
sis
(Wat
erh
ou
se,
18
90
)
Ass
am,
Aru
nac
hal
Pra
des
h,
Ch
hat
tisg
arh
,H
ary
ana,
Mad
hy
a
Pra
des
h,
Mah
aras
htr
a,M
egh
alay
a,
Nag
alan
d,
Tam
ilN
adu
,U
ttar
akh
and
and
Wes
tB
eng
al
Bal
agh
at,
Ch
hin
dw
ara,
Dam
oh
,S
eon
i,
Sh
ivp
uri
,U
mar
ia,
Man
dla
;P
TR
,
VD
WS
,M
NP
,B
NP
,K
NP
,K
TR
Bil
asp
ur,
Rai
pu
r;
AA
BR
,B
WL
S
My
anm
ar,
Nep
al,
No
rth
Ko
rea
and
Ch
ina
Studies on Tribe Gymnopleurini
123
Madhya Pradesh: Seoni: PTR, Turiya, day collection,
07.vi.2001 (3#); Khamreet, 24.vi.2001 (1#) coll.
K. Chandra. Chhindwara: PTR, Kokiwada, 28.ix.2001
(1#), coll. S. K. Mishra; Gumtara, 27.viii.2001 (1$), coll.
Y. N. Gupta.
Chhattisgarh: Bilaspur, AABR, Sarsadol Forest,
18.vii.2005 (1#), coll. A. Singh.
Diagnosis: (Fig. 2l). Dark greenish-blue, very smooth
and shining; broadly oval and rather convex; head granu-
late and opaque and clypeus quadri-dentate; pronotum and
elytra very smooth and shining; pronotum very finely but
not very closely punctured, except in front angles, where it
is minutely rugose and opaque; elytra very finely striate,
lateral margins strongly excised behind shoulders, expos-
ing meta-sternal epimera; base of abdomen sharply carinate
at sides.
Sexual dimorphism: Male: Extremity of pro tibia abruptly
incurved and truncate and terminal spur broad and truncate
at end with angles a little produced (Fig. 6c). Female:
Extremity of pro tibia comparatively not much incurved;
terminal spur acute and pointed (Fig. 6d).
External male genitalia: (Fig. 7e). Phallobase longer
than parameres; laterally parameres triangular in shape and
dorsally flat and equal in length.
Remarks: New record from Madhya Pradesh and
Chhattisgarh.
Garreta mundus (Wiedemann, 1819) (Figures 2b, f, m,
6e, f, 7f)
Gymnopleurus mundus, Wiedemann, 1819, Zool. Mag., I,
3: 162.
Gymnopleurus mundus, Arrow, 1931, Faun. Brit. India,
(Lamellicornia: Coprinae), III: 57.
Garreta mundus, Janssens, 1940, Verh. Kon. Nat. Mus.
Bel., 2 (18): 33.
Gymnopleurus (Garreta) mundus, Balthasar, 1963, Mon.
Scarab. Aphod. Palae. Ori. Reg., (Coleoptera: Lamellicor-
nia), Coprinae, I: 229.
Gymnopleurus capicola, Castelnau, 1840, Hist. Nat. Col.,
II: 70.
Collection data: 18 specimens: 7 male: 11 female.
Temporal distribution (all data): June (16), July (2);
Madhya Pradesh: Hoshangabad: BWS, Churna,
11.vi.1999 (9$5#), 10.vi.1999 (1$); Seoni: PTR, Kar-
majhiri, light trap, 13.vi.2001 (1#), coll. K. Chandra.
Chhattisgarh: Raipur: BWLS, Tinsa Pathar, 03.vii.2011
(1$1#), coll. K. Chandra (Nilgai dung).
Diagnosis: (Fig. 2m) Dull opaque and olive-green; dor-
sally without hairs; broad oval and not very convex; head
finely granular, with clypeus quadri-dentate and ocular
lobes slightly prominent (Fig. 2f); pronotum finely punc-
tured, base with a minute impression on each side of
middle; elytra very finely striate; elytral intervals finely
coriaceous and un-punctured.
Sexual dimorphism: Male: Spur of pro tibia broad and
bifid at end; clypeal dentations less acute (Fig. 6e). Female:
Spur of protibia long and acute; clypeal dentations more
acute and prominent (Fig. 6f).
External male genitalia: (Fig. 7f). Phallobase twice
longer than parameres; parameres almost triangular in
shape with its front view flat.
Remarks: Attracted towards light.
Garreta dejeani (Castelnau, 1840) (Figures 2n, 5g, h, 8a)
Gymnopleurus dejeani, Castelnau, 1840, Hist. Nat. Col., II:
70.
Gymnopleurus dejeani, Arrow, 1931, Faun. Brit. India,
(Lamellicornia: Coprinae), III: 59, pl. III, Fig. 11.
Garreta dejeani, Janssens, 1940, Ver. Kon. Nat. Mus.Bel.,
2 (18): 26, 31.
Gymnopleurus (Garreta) dejeani, Balthasar, 1963, Mon.-
Scarab. Aphod. Palae. Ori. Reg., (Coleoptera: Lamellicor-
nia), Coprinae, I: 227.
Gymnopleurus inconspicuus, Waterhouse, Ann. Mag. Nat.
Hist., (6), V: 371.
Collection data: 4 specimens- 3 male: 1 female. Tem-
poral distribution (all data): July (3), August (1);
Madhya Pradesh: Damoh: VDWS, Danital, 08.ix.2010
(1$), coll. K. Chandra.
Chhattisgarh: Raipur: BWLS, Kari Dongri Forest,
04.vii.2011 (3#), coll. K. Chandra.
Diagnosis: (Fig. 2n) Dark coppery greenish and entirely
dull above very broad and flat; head and pronotum closely
and finely granulate; clypeus rugose in front and pro-
duced into four lobes at front margin; elytra rather faintly
striate and intervals rather more finely and less closely
granular.
Sexual dimorphism: Male: Pro tibia longer than that of
female, more strongly but less regularly curved, and less
dilated from base to apex; there is a slight internal prom-
inence at extremity and terminal spur short and broad, a
little dilated at end with its outer acuminate (Fig. 5g).
Female: Pro tibial teeth more dilated than male and tibial
spur long and acute (Fig. 5h).
K. Chandra, D. Gupta
123
External male genitalia: (Fig. 8a). Phallobase twice as
longer than parameres; prameres triangular in shape; in
front view they are broad and almost circular.
Conclusion
The results of this study are based on the examination of
191 specimens that included eight species within three
genera (Garreta, Gymnopleurus and Paragymnopleurus) of
dung beetle tribe Gymnopleurini. The material examined
represented two states, 12 districts and 30 localities. All the
eight species are recorded from Madhya Pradesh. Except
Gymnopleurus (Metagymnopleurus) parvus MacLeay, the
rest of the seven species of the tribe are recorded from
Chhattisgarh. Moreover, Garreta smaragdifer (Walker) is
reported for the first time from Madhya Pradesh and
Chhattisgarh. The distribution of all species recorded to
date from Madhya Pradesh and Chhattisgarh is summarized
in Table 1. The distribution data suggests that highest
number of species (4 spp.) were collected from PTR and
BWLS followed by AABR and VDWS (2 spp. each) and
BWS, PBR, BNP, KNP, KTR, MNP, KNP (1 spp. each).
Based on the collection data, temporal distribution of all
the species is provided. Most of the specimens were
collected during June and July. All the specimens were
collected during day time only while a few specimens of
Paragymnopleurus sinuatus ssp. assamensis, Gymnopleu-
rus cyaneus and Garreta mundus were captured during
night with help of light. G. cyaneus and P. sinuatus sp.
assamensis were distributed evenly in both the states and
most abundant representing 104 and 36 specimens
respectively. G. dejeani was only recorded from, VDWS,
Damoh (MP) and BWLS, Raipur (CH). The south Indian
species, G. smaragdifer which is newly recorded from both
the states, has been only reported from PTR, Seoni (MP)
and Bilaspur (CH), extending its range towards central
India.
An identification key to the genera and species of
Gymnopleurini from MP and CH is also provided along
with illustrations (Fig. 1). The identification of species is
mainly based on the structures of clypeus, whether it is bi
or quadri-dentate and the punctation, granulation and
number of spots present on pronotum in dorsal view. As
species specificity of male genitalia is well documented
pattern in nature [30], the male genitalia of seven species
except Gymnopleurus parvus, were described and illus-
trated for the first time so as to confirm species identity
(Fig. 7). Concerning all the previously published infor-
mation, a checklist pertaining 21 species distributed in all
the 4 recognized genera of the tribe is prepared and their
geographical distribution is summarized in Table 1.
Acknowledgments The authors thank Dr. K. Venkataraman
(Director, Zoological Survey of India, Kolkata) for providing neces-
sary facilities and encouragement. Special thanks are also due to the
Chhattisgarh Forest Department, particularly the CAMPA office for
the logistic and financial support to carry out the studies on the
Protected Areas of Chhattisgarh. The authors are also thankful to Dr.
Ales Bezdek (Czech Republic), Dr. Paul Schoolmeesters (Belgium)
and Dr. James Noriega Alvarado (USA) for the literature. The authors
are also thankful to Dr. S. Sambath (Officer-in-Charge, ZSI, Jabalpur,
Madhya Pradesh) for providing necessary facilities and Mr. Sandeep
Kushwaha for his support during field visits.
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