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 Chhattisgarh with A Checklist from India Kailash Chandra Devanshu Gupta Received: 26 November 2012 / Revised: 5 June 2013 / Accepted: 16 July 2013 Ó 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 Á Gymnopleurini Á Checklist Á Male genitalia Á 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

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Page 1: Studies on Tribe Gymnopleurini (Coleoptera: Scarabaeidae: Scarabaeinae) from Madhya Pradesh and Chhattisgarh with A Checklist from India

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

Page 2: Studies on Tribe Gymnopleurini (Coleoptera: Scarabaeidae: Scarabaeinae) from Madhya Pradesh and Chhattisgarh with A Checklist from India

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

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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

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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

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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

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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

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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

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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

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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

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K. Chandra, D. Gupta

123

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Studies on Tribe Gymnopleurini

123

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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

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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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