REPRODUCTIVE BIOLOGY AND NESTING BEHAVIOUR 
OF THE PROMINENT DUNG BEETLES 
(SCARABAEINAE: COLEOPTERA) IN THE 
AGRIBELTS OF MALABAR COAST 
 
 
 
Thesis submitted to the 
 
UNIVERSITY OF CALICUT 
For the award of the Degree of 
DOCTOR OF PHILOSOPHY IN ZOOLOGY 
(Under the Faculty of Science) 
 
 
 
BY 
 
PRAMEELA K. 
 
 
Under the Guidance of 
Dr. SABU K. THOMAS 
 
 
 
 
P. G. & RESEARCH DEPARTMENT OF ZOOLOGY, 
ST. JOSEPH’S COLLEGE (AUTONOMOUS), DEVAGIRI, 
KOZHIKODE - 673 008, KERALA. 
MARCH 2022 






CONTENTS 
  Page No. 
Chapter  1:  INTRODUCTION                                                  
1.1 Distribution of dung beetles 2 
1.2 Ecological importance of dung beetles 3 
1.3 Prominent dung beetles in  different regions 4 
1.4 Prominent dung beetles in India 5 
1.5 Nesting behaviour and biology of dung beetles  5 
1.6 Nesting behaviour and biology of dung beetles in India 8 
Chapter 2: REVIEW OF LITERATURE   
2.1 Taxonomy of dung beetles of the world 13 
2.2 Distribution of dung beetles in India 15 
2.3 Prominent dung beetle species in the world 17 
2.4 Prominent dung beetles species in India   18 
2.5 Reproductive biology of dung beetles in the world 19 
2.6 Reproductive biology of dung beetles in India  22 
Chapter 3: MATERIALS AND METHODS   
3.1 Rearing of dung beetles 24 
3.2 Study of nesting pattern 27 
Chapter 4: RESULTS   
4.1 Life biology of Onthophagus  cervus (Fabricius,1798) 29 
4.2 Life biology of Onthophagus  fasciatus (Boucomont,1914) 31 
4.3 Life biology of Tiniocellus  spinipes (Roth,1851) 33 
4.4 Life biology of Sisyphus longipes (Olivier,1789)  35 
   
4.5 Life biology of Tibiodrepanus setosus (Wiedemann, 1823) 37 
Chapter 5: DISCUSSION   
5.1 Onthophagus cervus 48 
5.2 Onthophagus fasciatus 52 
5.3 Tiniocellus spinipes 55 
5.4 Sisyphus longipes 57 
5.5 Tibiodrepanus setosus 59 
   
Chapter 6: CONCLUSION  61 
Chapter 7 : REFERENCES  63 
 LIST OF FIGURES  85 
 LIST OF TABLES  87 
 
 
 
 
 
 
 
 
 
 
 
 
 

                                                   INTRODUCTION 
Dung beetles belonging to the subfamily Scarabaeinae is a highly diverse and 
broadly distributed group, characterized by dung and organic debris at both the adult 
and larval stages (Hanski and Cambefort, 1991).The dung beetles include three 
subfamilies of Scarabaeidae (Insecta: Coleoptera), Aphodiinae, Geotrupinae and 
Scarabaeinae. Within the subfamilies, Scarabaeinae is the only group that is mainly 
coprophagous (faeces eating). The majority of Aphodiinae and Geotrupinae are 
saprophagous (eaters of decaying organic matter), not true dung beetles (Halffter and 
Matthews, 1966; Scheffler, 2002). Dung beetles are categorised into 12 tribes which 
included Coprini, Dichotomini, Phanaeini, Oniticellini, Onitini, Onthophagini, 
Eucraniini, Eurysternini, Canthonini, Gymnopleurini, Scarabaeini and Sisyphini 
(Lawrence and Newton, 1995). 
Based on their nesting strategies dung beetles are divided into three functional 
groups, namely, rollers (telecoprid nesters), tunnelers (paracoprid nesters) and dwellers 
(endocoprid nesters) (Cambefort and Hanski, 1991). In Scarabaeinae, dwelling is 
associated with tribe Oniticellini and tunneling with tribes Coprini, Onitini and 
Onthophagini, dung rolling is associated with tribes Scarabaeini, Gymnopleurini, 
Sisyphini and Canthonini (Halffter and Edmonds, 1982). Regarding the morphological 
changes between the functional groups, the tunneling dung beetles have comparatively 
smaller hind legs and the front legs are well suited for digging. The presence of horns is 
common in tunneling dung beetles. The rollers commonly have long hind legs. The 
rollers roll the dung ball using their back legs (Hanski and Cambefort, 1991; Scholtz et 
al., 2009).  
Most dung beetles use one of the three wide nesting strategies, tunnellers, rollers 
and dwellers. Tunneller (Paracoprid) species bury brood balls in vertical chambers in 
1 
 
the locality of the original deposition site. Roller (telecoprid) species carry balls to some 
horizontal distance away, before burial below the soil surface. Dweller (endocoprid) 
species brood their young inside the dung mass itself (Halffter and Edmonds, 1982). 
This functional stratification permits dung beetles to decrease the intense struggle for 
limited food and space and protect the food from adverse environmental conditions such 
as heat and extreme dryness (Halffter and Edmonds, 1982; Cambefort and Hanski, 
1991). 
 1.1. Distribution of dung beetles. 
   Provincial lists of dung beetles are obtainable from South Africa (Péringuey, 
1900; Chown et al., 1995), African Tropical region (Gillet, 1908, 1911), Sumatra 
(Gillet, 1924), China (Gillet, 1935; Nakane and Shirahata, 1957; Bin -Hong Ho, 2018), 
Central America, the West Indies South America (Blackwelder, 1944) Afghanistan 
(Balthasar, 1955), Japan (Nakane and Tsukamoto, 1956), Florida (Woodruff, 1973), 
Panama and Costa Rica (Howden and  Young, 1981; Howden and  Gill, 1987; 
González-Maya and Mata-Lorenzen, 2008), Nebraska (Ratcliffe, 1991; Wagner et al., 
2020), Europe (Baraud, 1992), Colombia (Lopera, 1996), Nearctic Realm (Smith, 
2003) and Palaearctic region (Löbl and  Smetana, 2006). World checklists of dung 
beetles were prepared by Krajcik (2006) and Schoolmeesters (2019). Checklists of 
dung beetles of Pakistan (Siddiqui et al., 2014), Northern Brazil (Pacheco and Vaz-
de-Mello, 2019), the Mediterranean region (Löbl and Löbl, 2016), Ecuador (Espinosza 
and Noreiga, 2018), Mexico (Sanchez - Hernandez et al., 2020), Southwest Arabia 
(Paulian, 1938; Ziani, 2021), Neotropical region (Gonzalez and Vaz-de-Mello, 2021) 
and Cambodia (Han et al., 2021) are available. 
Arrow (1931) and Balthasar (1963a, b) provided comprehensive information 
about the dung beetles in the Indian subcontinent.  Recent works on the taxonomy of 
2 
 
dung beetles from different parts of India are Biswas and Chatterjee,1985; 
Veenakumari and Veeresh,1996; Mittal,1999; Chatterjee and Biswas, 2000; Chandra, 
2005; Sewak, 2006; Schoolmeesters and Sabu, 2006; Chandra and Ahirwar, 2007; 
Vinod and Sabu, 2007; Vinod, 2009; Sarkar et al., 2010; Latha et al., 2011; Sabu et 
al., 2011; Venugopal et al., 2012; Sarkar et al., 2015; Sathiandran et al., 2015; 
Karimbumkara and Priyadarsanan, 2013, 2016  ; Gajendra and Prasad, 2016; Subha 
and Sabu, 2017; Kalawate, 2018; Patole, 2019 and Kharel et al., 2020). 
1.2. Ecological importance of dung beetles.  
Dung beetles are an economically important group of beetles that play a central 
role in forest soil conditioning, as they are the chief agents in soil aeration, improving 
soil structure and water circulation and modify organic debris, making it usable for 
other organisms (Halffter and Mathews, 1966; Bornemissa and Williams, 1970; 
Nealis, 1977; Mittal, 1993). Faeces decompose four times faster in the presence of 
dung beetles (Gillard, 1967), thus the rate of nutrient cycling is increased (Miranda et 
al., 1998). Dung beetles have very low efficiency of assimilation of energy (Holter, 
1975) and much of what they ingest is quickly egested and is thus rapidly available to 
other organisms (Scheffler, 2002). Dung beetles also accelerate bacterial growth and, 
unlike earthworms, incorporate manure and carrion and plant material in the soil (Lutz, 
1931). Dung beetles are among the most important invertebrates to dung 
decomposition in temperate and tropical agricultural grasslands (Gittings et al., 1994; 
Davis, 1996; Horgan, 2001; Lee and Wall, 2006; Slade et al., 2011; Kaartinen et al., 
2013). Dung removal, seed dispersal, nutrient cycling and reduction of greenhouse gas 
emissions are the major ecosystem services provided by dung beetles (Slade et al., 
2011; Lugon et al., 2017; Menendez et al., 2016; Nervo et al., 2017 and Piccini et al., 
2017).  
3 
 
1.3 Prominent dung beetles in different regions.  
Dominant dung beetle species varies among different regions. Oniticellus 
pseudoplanatus Balthasar, 1964 in moist forests of Ivory Coast (Cambefort and Walter, 
1991); Onthophagus vulpus Harold, 1877 and Sisyphus thoracicus Sharp, 1875in the 
rainforest in Malaysia (Davis, 2000); Dichotomius amplicollis Harold, 1869, 
Deltochilum gibbosum (Fabricius, 1775) and Onthophagus landolti Harold, 1880 in 
Mexican dry forest (Andresen 2005, 2008); O. wallacei Harold, 1871, and 
O.fuscostriatus Boucomont,1914 in Indonesian forest (Shahabuddin,2010); 
Dichotomius nisus (Olivier, 1789), Trichillum externepunctatum (Preudhomme de 
Borre, 1880), Canthon podagricus (Harold,1868), Onthophagus hirculus 
(Mannerheim, 1829), Pseudocanthon perplexus (LeConte, 1847), Ontherus sulcator 
(Fabricius, 1775)  and  Ataenius platensis (Blanchard, 1846) in the Argentine cattle 
ranches (Damborsky et al., 2015), Canthon histrio Serville, 1828, Onthophagus 
hirculus (Mannerheim, 1829) and Deltochilum verruciferum Felshe, 1911 in Brazilian 
dry forest (Novais et al., 2016), Canthonquinque maculatus Castelnau,1840 Canthon 
conformis Harold, 1868, Dichotomius serices (Harold, 1867) in Southern Atlantic 
forest of Argentina (Andrés Gómez, 2017), Ontherus pubens Genier, 1996 in Ecuador 
(Espinosza and Noreiga, 2018), Deltochilum mexicanum (Burmeister, 1848) and 
Dichotomius satanas (Harold, 1867) in Mexico (Barretto et al., 2019),  Onthophagus 
hecate (Panzer, 1794), O. pennsyl vanicus Harold, 1871 and Diapterna pinguella 
(Brown, 1929) in the Nebraska Sandhills Ecosystem (Wagner et al., 2020), Eurysternus 
caribaeus (Herbst,1789), E. nigrovirens Génier, 2009, Dichotomius carbonarius 
(Mannerheim, 1829), Onthophagus hirculus (Mannerheim, 1829), O. buculus 
Mannerheim, 1829, Canthon simulans (Martinez, 1950), C. fortemarginatus 
4 
 
Balthasar,1939, and Canthidium barbacenicum (Preudhomme de Borre,1886) in  south 
Western Brazil Cerrado (Silva et al., 2021).  
1.4. Prominent dung beetles in India.  
 Dominant dung beetles in India are Digitonthophagus gazella (Fabricius, 
1787), Onthophagus rectecornutus Lansberge, 1883, Copris repertus Walker,1858, C. 
fricator (Fabricius, 1787) in Deccan region in south India (Veenakumari and Veeresh, 
1996, 1997); Caccobius ultor  Sharp, 1875 in the forests of Haryana in North-Western 
India (Mittal, 2005; Kakkar and Gupta, 2009, Kakkar, 2010); Caccobius vulcanus 
(Fabricius, 1801), C. ultor Sharp,1875, Onthophagus centricornis Fabricius, 1798, 
O.cervus (Fabricius, 1798), O. fasciatus Boucomont, 1924, O. dama (Fabricius, 1798), 
Tiniocellus spinipes (Roth, 1851), Sisyphus longipes (Olivier, 1789) and 
Tibiodrepanus setosus (Wiedemann, 1823) in the moist belts of south  India (Vinod, 
2009; Thomas et al., 2011; Venugopal et al., 2012; Nithya, 2012; Nithya et al.,2015; 
Shobhana, 2016; Sabu, 2012 and Subha, 2018). Patole (2019) reported Catharsius 
pithecius (Fabricius, 1775) and Gymnopleurus cyaneus (Fabricius, 1798) from the 
agriculture belts in Maharashtra. Two Tiniocellus species, Tiniocellus imbellis (Bates, 
1891) and T. spinipes (Roth, 1851) were reported in the Tropical Forest of the 
Himalayan foothills, West Bengal, India. (Sarkar and Kharel, 2020). 
1.5. Nesting behaviour and biology of dung beetles. 
Like all insects, scarabs undergoes four stages in their life cycle: egg, larvae, 
pupae and adult. (Halffter and Mathews,1966; Hanski and Cambefort, 1991; Scheffler, 
2002). All known Scarabaeinae exhibit some degree of nesting behaviour, Juvenile 
development takes between 30⎼50 days from egg to adult and in adverse conditions 
over a year. After emergence from the nest adults undergoes a prolonged feeding 
period for three to four months to develop gonads and eggs. The average life span of 
5 
 
60 days to three years (Scheffler, 2002). Nest preparation and brood mass construction 
are the pioneers of the next step in the breeding process (Halffter and Edmonds, 1982). 
Reproductive biology and nesting behaviour of dung beetles have been broadly studied 
and critically analysed in different parts of the world. Klemperer (1983) described the 
subsocial behaviour of  Oniticellus cinctus (Fabricius,1775), from Birmingham. 
Biology and nesting behaviour of Onitis viridulus Boheman, 1857, O. fulgidus 
Castelnau, 1840, O. obscures Lansberge,1886 O. alexis Klug, 1835, O. perpunctatus 
Balthasar, 1963a, O. caffer Boheman,1857. O. aygulus (Fabricius, 1781), O. tortuosus 
Houston, 1983 O. receptor, O. uncinatus Klug, 1855, O. picticollis Boheman, 1857  
and O. pecuarius Lansberge,1875 from South Africa were studied by Edwards and 
Aschenborn(1987). Life history of Onthophagus medorensis Brown, 1929 Hunter et 
al., (1991). Studies on reproductive biology and nesting of Onthophagus stylocerus 
Graells, 1851 by Romero and Piera (1995). Studies on the brood care behaviour and 
nest structure of the dung beetle Onthophagus vacca (Linnaeus, 1767) by  Sowig 
(1996). Studies on the life history of O. depressus Harold, 1871 by Hunter et al., 
(1996). The reproductive biology of the O.incensus (Say, 1835) was studied in Mexico 
by Martínez et al., (1998). Biology of the dung beetle O. hirculus Mannerheim, 1829 
done by Gonzalez-Vainer and Morelli (1999). Reproductive biology of Onitis belial 
(Fabricius, 1789) from Morocco, O. anthracinus Felsche, 1907 and O.vanderkelleni 
Van Lansberge,1886 from Kenya, by Palestrini et al., (2002). The cost of reproduction 
of dung beetle Onthophagus binodis (Thunberg, 1818) was studied in Australia by 
Kotiaho and Simmons (2003). Comparative analysis of reproductive and nesting 
behaviour in several species of  Eurysternus Dalman, 1824 by Huerta et al.,(2003). 
Studies on fecundity and offspring survival of Copris tripartitus Waterhouse, 1875 
from Mexico was done by Huerta and Bang (2004). Studies on the life cycle, 
6 
 
preimaginal development and phenology of Onthophagus landolti Harold, 1880 in 
Mexico was done by Pérez-Cogollo et al., (2015) and the cost of reproduction of 
Callosobruchus maculatus (Fabricius,1775) was done by Paukku and Kotiaho(2005). 
The rolling and tunneling behaviour of large-sized subsocial African dung rolling 
beetle Scarabaeus catenatus (Gerstaecker,1871) were studied by Sato (2007). 
Reproductive development and seasonal activity of two Korean native Coprini species 
Copris ochus (Motschulsky, 1860) and C. tripartitus Waterhouse, 1875 studied by 
Bang et al., (2008). Reproductive activity of Onthophagus granulatus Boheman, 1858 
was studied in New Zealand by Forgie (2009). Studies on pre-imaginal stages of O. 
incensus (Say, 1835) by Huerta et al., (2010). Nesting behaviour of O. incensus (Say, 
1835) was studied in Mexico by Huerta and Hernández (2013). Pérez - Cogollo et 
al.,(2015) studied the life history of O. landolti Harold, 1880, in Mexico. The feeding, 
reproductive and nesting behaviour of Canthon bispinus (Germar, 1824) from 
Uruguay (González-Vainer, 2015). Comparative studies on the nesting and food 
relocation behaviour of Eucranium Brulle, 1834 with that of the morphologically 
similar South African subgenus Scarabaeus (Pachysoma) Macleay,1821 was done by 
Ocampo and Philip (2017). The life history of the dung beetle Onthophagus lecontei 
Harold, 1871 was studied in Mexico by Arellano et al., (2017). The reproductive 
biology of Euoniticellus intermedius (Reiche, 1848) was studied in Mexico by 
Martinez et al., (2019). Medina et al., (2020) conducted a study on the feeding and 
reproductive behaviour of the dung beetle Canthon rutilanscyanescens Harold, 1868 
in Brazil.  
 
 
 
7 
 
1.6. Nesting behaviour and biology of dung beetles in India.   
Only scant information is available on the life-cycle of Indian dung beetle 
species and the details are as follows; Joseph (1994) carried out a study on sexual 
dimorphism and intra sex variations of the giant dung beetle Heliocopris dominus 
Bates (1868). Veenakumari and Veeresh (1996b) studied the feeding and breeding 
behaviour of Gymnopleurus gemmatus (Harold, 1871) and G.miliaris (Fabricius 
1775). Studies on the reproductive biology of the two commonly occurring south 
Indian species Onthophagus gazella (Fabricius, 1787) and O.rectecornutus Lansberge, 
1883 was done by Veenakumari and Veeresh (1996c). Subsociality in Copris repertus 
Walker, 1858 and C. indicus Gillet, 1910 was studied by Veenakumari and Veeresh 
(1997). Joseph (1998) conducted studies on the life biology and breeding behaviour of 
Heliocopris dominus Bates (1868). Studies on the life cycle, ecological role and 
biology of immature stages of H. dominus Bates, 1868 was done by Joseph (2003). 
Nidification behaviour of three dung beetle species, Onthophagus catta (Fabricius 
1787), Onitis philemon (Fabricius, 1801) and Liatongus rhadamistus (Fabricius, 1775) 
were studied in Maharashtra by Gaikwad and Bhawane (2015). Study of nesting and 
biology of the dung beetle Scaptodera rhadamistus (Fabricius, 1775) from 
Maharashtra was done by Khadakkar et al., (2018). Analysis of the nesting 
architecture, life cycle, and brood ball morphometry of the dung beetle Oniticellus 
cinctus (Fabricius, 1775) was studied in Dehradun by Singh et al., (2019).  
It is impossible to interpret the exact mechanism behind the seasonality and 
abundance of individual species and genera due to lack of knowledge on the biology 
and ecology of prominent dung beetles species (Sabu, 2012; Vinod, 2009; Nithya, 
2012; Latha, 2011; Sobhana, 2016; and Subha, 2018). The present study has been 
undertaken to understand the life-history traits of the five prominent dung beetle 
8 
 
species, Onthophagus cervus (Fabricius, 1798), Onthophagus fasciatus Boucomont, 
1924, Tiniocellus spinipes (Roth, 1851), Sisyphus longipes (Olivier, 1789), and 
Tibiodrepanus setosus (Wiedemann, 1823) in the agribelts of Malabar Coast region in 
south India.  
 
The genus Onthophagus Latreille, 1802 is a hyperdiverse and largest genus, 
within the tribe Onthophagini of the subfamily Scarabaeinae. The peculiarity of the 
Onthophagus genus is characterized by their excavation habit of construction 
galleries beneath the dung, and females are produced a large number of eggs 
(Fecundity is high) (Halffter and Edmonds, 1982; Delgado,1997; Pulido and 
Zunino,2007). Onthophagus is a well-adapted species capable of surviving in a 
variety of habitats including disturbed habitats like crop fields and may produce 
several broods per year as common in small tunnelers (Cambefort and Hanski, 1991). 
The tribe Onthophagini belong to two species, Onthophagus cervus (Fabricius, 1798) 
and Onthophagus fasciatus Boucomont, 1924 are the prominent tunneller species in 
the moist belts of south India (Vinod, 2009; Thomas et al., 2011; Venugopal et al., 
2012; Nithya, 2012; Nithya et al; 2015; Shobhana, 2016; Sabu, 2012 and Subha, 
2018). 
 The tribe Oniticellini Kolbe, 1905 include 4 subtribes namely Drepanocerina 
van Lansberge, 1875, Oniticellina Kolbe, 1905, Helictopleurina Janssens, 1946 and 
Eurysternina Volcano, Martinez and Pereira, 1960 (Branco, 2010 and Philips, 2016). 
Tiniocellus spinipes (Roth, 1851) comes under the soil tunneling genus Tiniocellus of 
the subtribe Oniticellina (Cambefort and Lumaret, 1983). Oniticellina sub-tribe differ 
from other sub-tribes due to these characters are as follows, the pygidium lacks the 
basal transverse carina, the dorsal face is either glabrous or with simple pilosity, and 
9 
 
the scutellum is small but always distinct (Janssens, 1949). So far only the data 
available on the biology of subtribe Oniticellina species namely, Liatongus 
rhadamistus (Fabricius, 1775) in West India (Gaikwad and Bhawane, 2015) and 
Euoniticellus intermedius (Reiche, 1848) in Mexico (Martinez et al., 2019) have been 
reported. Tiniocellus spinipes is a small-sized beetle having, Length of 6 mm, a 
breadth of 2.5mm; and inhabited in Mammalian dung. Tiniocellus Péringuey, 1901 is 
a species-poor genus and has only seven species throughout the world.  Tiniocellus. 
spinipes (Roth,1851), The Asiatic T. imbellis (Bates, 1891); T. asmarensis Balthasar, 
1968 the African T. setifer (Kraatz,1895), T. praetermissus (Branco, 2010), T. dolosus 
(Branco,2010) and T. eurypygus (Branco, 2010).Two species, T. imbellis (Bates, 
1891) and T. spinipes (Roth, 1851) are known from India, Schoolmeesters (2019). 
        Sisyphini is regarded as the true dung ball rollers (Daniel et al., 2020) and 
Sisyphus longipes (Olivier, 1789) belongs to the tribe Sisyphini. As a member of the 
roller guild, it has the following features that assist the species in dung rolling. It 
contains eight antennal articles, comparatively short bodies that are laterally 
compressed and flattened, especially at the sides of the pronotum, elytra that are broad 
proximally but attenuate posteriorly and the exceptionally long middle and hind legs. 
The tribe contains species of minor to moderate body size averaging 7.0⎼10.0 mm. 
The Sisyphus species in India are Sisyphus longipes (Olivier, 1789) S. neglectus Gory, 
1833, S. araneolus Arrow 1927, S. hirtus Wiedemann, 1823 (Arrow 1927), and S. 
indicus Hope, 1831. Sisyphus longipes as the dominant roller species in the moist belts 
of South India (Vinod, 2009; Sabu, 2011; Mathew, 2011; Simi, 2012; Nithya, 2012; 
Nithya et al., 2015 Subha, 2018) and showed significant seasonal variation with high 
abundance in the Monsoon season (Sabu, 2011).  
10 
 
     The genus Tibiodrepanus was introduced by Krikken (2009) Tibiodrepanus 
setosus (Wiedemann, 1823) comes under the dwelling subtribe Drepanocerina. The 
subtribe Drepanocerina is composed of 11 genera and 46 species (Schoolmeesters, 
2019). The prominent dweller species in the moist belts of South India is 
Tibiodrepanus setosus (Vinod, 2009; Sabu, 2011; Mathew, 2011; Simi, 2012; Nithya, 
2012; Nithya et al., 2015 Subha, 2018). 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
11 
 
 
                                   OBJECTIVES               
   Study of following aspects of prominent dung beetle species in the agricultural 
fields in the Malabar Coast region. 
1. Life biology and  
2. Fecundity, egg mortality, larval and pupal duration, adult life span, nesting 
strategies and voltinism of prominent dung beetles species.  
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
12 
 


REVIEW OF LITERATURE 
 
 2.1. Taxonomy of dung beetles of the world 
         The dung beetles were categorized under subfamily Scarabaeinae and members 
of the suborder Lamellicornia were included by Linnaéus (1758) under a single genus, 
the Scarabaeus. The dung beetles derived from the Linnean Scarabaeus and constituted 
a new genus Copris by Fourcroy (1785). Latreille (1796) placed the species with 11-
jointed antennae under the name Geotrupes. Two new genera Oryctes and Aphodius 
were introduced by Illiger (1798). Fabricius (1798) separated genus Onitis from genus 
Copris. Creutzer (1799) suggested the name Actinophorus for the ball rolling beetles 
now comprised in the genera Scarabaeus and Gymnopleurus. Weber (1801) introduced 
the name Ateuchus sacer for Scarabaeus sacer. The largest dung beetle genus, 
Onthophagus was introduced by Latreille (1802). The genus Gymnopleurus was 
introduced by Illiger (1803). Latreille (1807) introduced the genus Sisyphus. Oniticellus 
genus was introduced by Serville (1825). Genus Drepanocerus was introduced by 
Kirby (1828). The two new genera, Catharsius and Heliocopris comprising large dung 
beetles introduced by Hope (1837). The genus Caccobius was introduced by Thomson 
(1863). The genus Liatongus was introduced by Reitter (1892) and the genus 
Tiniocellus by Péringuey (1900). Boucomont (1914) introduced the genus Phacosoma. 
Due to homonymy, Vaz-de-Mello (2003) renamed the genus Phacosoma as 
Ochicanthon. The genus Tibiodrepanus was introduced by Krikken (2009) which is 
formerly supposed to be the genus Drepanocerus Kirby, 1828.  
Arrow (1931) discarded the classification system suggested by Lacordaire 
(1856) and classified dung beetles in four divisions (= tribes) viz. Scarabaeini, 
13 
 
Sisyphini, Coprini, and Phanelini under subfamily Coprinae with which he considered 
the Scarabaeinae synonymous. Scarabaeinae subdivided into six tribes: Coprini, 
Eurysternini, Oniticellini, Onitini, Onthophagini and Scarabaeini by Janssens (1949). 
Later, Balthasar (1963a,b) positioned the group as a family including two 
behaviourally diverse subfamilies: Coprinae and Scarabaeinae. The previous 
subfamily involved the tribes Coprini, Dichotomini, Phanaeini, Oniticellini, Onitini 
and Onthophagini while the latter subfamily included the tribes Eucraniini, 
Eurysternini, Canthonini, Gymnopleurini, Scarabaeini, and Sisyphini. A study by 
Zunino (1984), which focused on the systematics of the subfamily Scarabaeinae based 
on the comparative analysis of the male and female genitalia disputed the monophyly 
of the tribes Onitini, Coprini, and Dichotomini.   
Lawrence and Newton (1995) placed all 12 tribes in the subfamily Scarabaeinae 
which they consider the Coprinae synonymous. Krikken (2009) revised and discussed 
the taxonomic and biogeographic status of the genus Drepanocerus Kirby, 1828 and 
split the genus into five new subgenera namely, Afrodrepanus, Clypeodrepanus, 
Latodrepanus , Sulcodrepanus and Tibiodrepanus. 
Dung beetles were recorded from South Africa (Péringuey ,1900; Chown et al., 
1995; Davis ,2002), African Tropical region (Gillet ,1908, 1911), Sumatra (Gillet 
1924), China (Gillet 1935; Nakane and Shirahata 1957 and Bin -Hong Ho 2018), 
Southwest Arabia (Paulian , 1938; Ziani, 2021), Central America, the West Indies and 
South America (Blackwelder 1944; Afghanistan (Balthasar, 1955), Japan (Nakane and 
Tsukamoto 1956), Florida (Woodruff ,1973), Panama and Costa Rica (Howden and  
Young 1981; Howden and  Gill 1987; González-Maya and Mata-Lorenzen 2008), 
Nebraska (Ratcliffe 1991; Wagner et al., 2021), Europe (Baraud, 1992), Colombia 
14 
 
(Lopera ,1996), Nearctic Realm (Smith, 2003) and Palaearctic region (Löbl and  
Smetana, 2006). Krajcik (2006) and Schoolmeesters (2011) prepared world checklists 
of dung beetles. Siddiqui et al., (2014) gave an annotated list of scarabs collected from 
vicinities of Pakistan with the faunal composition. The distribution and species 
diversity of dung beetles from the Mediterranean region were reported by Löbl and 
Löbl (2016). Philips (2016) described the tribe, Oniticellini Kolbe, 1905 and it has four 
subtribes, viz. Drepanocerina van Lansberge, 1875, Oniticellina Kolbe, 1905, 
Helictopleurina Janssens, 1946 and Eurysternina Volcano, Martinez and Pereira, 1960 
and provided the worldwide data on the tribe Oniticellini Kolbe, 1905 and it belongs 
to 26 genera and 252 species Branco (2010).  Silva (2017) presented an annotated list 
of the dung beetle species from, Southern Brazil. Espinosza and Noreiga (2018) 
provided distributional data and recorded 14 genera and 54 species from Ecuador. 
Salomao et al., (2019) analyzed a total of 945 species from the mosaic habitat at the 
ecotone of Savanna ecosystems in North-Eastern Brazil. Schoolmeesters (2019) 
provided worldwide records of dung beetles.  Sanchez-Hernandez et al., (2020) 
reported 112 species and 7 subspecies belonging to 23 genera, 7 tribes, and 4 sub-
tribes of the subfamily Scarabaeinae from Natural Protected Areas in Mexico. 
González-Alvarado and Vaz-de-Mello (2021) provide a complete taxonomic revision 
of the Neotropical dung beetle of sub genus Deltochilum (Deltohyboma) Lane 1946. 
2.2. Distribution of dung beetles in India. 
Data on the occurrence of dung beetles are mainly based on the publications of 
the Zoological Survey of India and the works carried out regionally from different 
parts of India. Arrow (1931) recorded 48 species of dung beetles from the western 
slopes of the south Western Ghats. Biswas and Chatterjee (1985) reported 7 new 
15 
 
species namely, Oniticellus namdaphensis, Oniticellus subhendui, Oniticellus gayeni, 
Onthophagus tirapensis, Onthophagus arunachalensis, Onthophagus songsokensis 
and Onthophagus royi from Namdapha wildlife sanctuary. Biswas and Chatterjee 
(1986) reported 3 new species namely Onthophagus keralicus, Onthophagus sahai, 
and Onthophagus taruni, and recorded 16 species from the Silent Valley National 
Park.  Veenakumari and Veeresh (1996a) described 61 species of Scarabaeinae 
belonging to three tribes from Bangalore in the Deccan region. Chatterjee and Biswas 
(2000) reported 27 species from Tripura State. Chandra (2000) published an inventory 
of 96 species of scarab beetles and their dispersal from the protected areas of Madhya 
Pradesh. Biswas and Mulay (2001) noted 71 species from Nilgiri Biosphere Reserve. 
As part of the biodiversity documentation program by the Kerala Forest Research 
Institute, 37 species of dung beetles from Kerala was reported by Mathew (2004). An 
account of the scarabaeid beetles of Himachal Pradesh was published by Chandra 
(2005). A new species, Onthophagus devagiriensis from a moist deciduous forest in 
the Wayanad region of Kerala State was reported by Schoolmeesters and Sabu (2006). 
Chandra and Ahirwar (2007) provided a comprehensive account of the scarab beetles 
of Chhattisgarh and Madhya Pradesh. Vinod (2009) prepared a checklist of 58 species, 
including 13 genera and 7 tribes of the Wayanad region. Latha et al., (2011) gave a 
revision of the taxonomic status of the Scarabaeinae genus Ochicanthon Vaz-de-Mello 
2003 and 15 species were reported with 8 new species from the Western Ghats. Sabu 
et al., (2011) prepared a checklist of 142 species from the moist South Western Ghats 
including five new species. Karimbumkara and Priyadarsanan, (2013) published a 
comprehensive list of 145 species of dung beetles belonging to 9 tribes and 23 genera 
reported from Karnataka. Sarkar et al., (2015) described the systematics of 19 
Scarabaeinae species under 6 genera reported from Buxa Tiger Reserve, West Bengal. 
16 
 
Sathiandran et al., (2015) published an illustrated checklist of 36 species of dung 
beetles from the Periyar Tiger Reserve in the southern Western Ghats. Mittal and Jain 
(2015) studied the taxonomy of the Indian dung beetle and recorded 420 species in 38 
genera. Karimbumkara and Priyadharsanan (2016) reported three new species 
Onthophagus jwalae (Kerala), O. pithankithae (Karnataka), and O. tharalithae 
(Assam) from India. Subha and Sabu (2017) reported bioindicator dung beetles from 
a shaded coffee plantation in the Nilgiri Biosphere Reserve of the south Western Ghats.  
Kalawate (2018) gave a preliminary study on the dung beetles of the Northern Western 
Ghats, Maharashtra, Khadakkar et al., (2019) collected and identified a total of 97 
scarab beetles species of 39 genera belonging to 7 subfamilies, where 10 species were 
newly recorded from different habitats of the Vidarbha region of Central India.  Patole 
(2019) analyzed the diversity and relative abundance of dung beetles from Sakri tahsil, 
Dist Dhulia Maharashtra. Latha and Sabu (2018) collected 34 species, belonging to 11 
genera and 7 tribes from Nelliampathy.  Sarkar and Kharel (2020) provided the first 
faunistic account on the Onthophagus Latreille, 1802 of the Nadia district, West 
Bengal. Chauhan and Uniyal (2020) prepared a checklist of dung beetles of 
Uttarakhand. Sarkar and  Kharel (2020) published the first faunistic study on the 
tribe Oniticellini Kolbe, 1905 (Coleoptera: Scarabaeidae) of Baikunthapur Tropical 
Forest in the Himalayan foothills, West Bengal. 
2.3. Prominent dung beetle species in the world  
Dominant dung beetle species vary among different global regions. Cambefort 
and Walter (1991) reported Oniticellus pseudoplanatus Balthasar, 1964 as the 
prominent species in moist forests of the Ivory Coast. Othophagus vulpus Harold, 1877 
and Sisyphus thoracicus Sharp, 1875 were reported as the dominant dung beetle 
17 
 
species in the tropical rainforests of Malaysia Davis (2000). Dichotomius amplicollis 
(Harold,1869), Deltochilum gibbosum (Fabricius,1775) and Onthophagus landolti 
Harold,1880 were recorded as the prominent beetles from Mexican dry forest 
(Andresen 2005, 2008). Shahabuddin (2010) recorded O. wallacei Harold, 1871 and 
O. fuscostriatus Boucomont, 1914 as the dominant species from an Indonesian forest. 
Onitis crassus Sharp, 1875 was reported as the major species from Pakistan by Ali et 
al., (2015). Canthon histrio Serville, 1828, Onthophagus hirculus Mannerheim, 1829, 
and Deltochilum verruciferum Felshe, 1911 were reported as the prominent species in 
Brazilian dry forest by Novais et al., (2016). Canthon quinque maculatus Castelnau, 
1840, Canthon conformis Harold,1868 and Dichotomius sericeus Harold,1867  were 
reported as the  dominant species in the southern Atlantic forest of Argentina (Gomez 
Cifuentes et al., 2017). Espinosza and Noreiga (2018) reported Ontherus pubens 
Genier, 1996 as the abundant species from Ecuador. Wagner et al., (2021) reported 
Onthophagus hecate Panzer, 1794, O. pennsylvanicus Harold, 1871 and Diapterna 
pinguella Brown, 1929 as the abundant species from the Nebraska Sandhills 
Ecosystem in the USA.  
2.4. Prominent dung beetles species in India   
Onthophagus gazelle Fabricius, 1787, O. rectecornutus Lansberge, 1883, 
Copris repertus Walker, 1858, and C. fricator Fabricius, 1787 were reported as the 
prominent species from the Deccan region in south India (Veenakumari and Veeresh 
1996a,1997). Three prominent species, Tiniocellus spinipes (Roth, 1851), 
Tibiodrepanus sinicus Harold, 1868 and Caccobius ultor Sharp, 1875 were reported 
from the forests of Haryana (Mittal 2005; Kakkar and Gupta, 2010). Vinod, (2009); 
Sabu, (2011); Simi et al., (2012); Nithya, (2012); Latha,(2013); Shobhana, (2016); and 
18 
 
Subha, (2018) reported Caccobius vulcanus Fabricius, 1801, C. ultor Sharp, 1875, 
Onthophagus centricornis Fabricius, 1798, O. cervus (Fabricius,1798), O. dama 
(Fabricius, 1798), Tiniocellus spinipes (Roth, 1851), Tibiodrepanus setosus 
Weidemann, 1823 and Sisyphus longipes (Olivier, 1789) as the prominent species from 
the ‘moist belts of South India. Catharsius pithecius (Fabricius, 1775) and 
Gymnopleurus cyaneus (Fabricius, 1798) were reported as dominant dung beetle in 
the agriculture belts of Maharashtra Patole (2019). Sarkar and Kharel (2020) reported 
Tiniocellus imbellis (Bates,1891) and T.spinipes (Roth,1851) as the dominant species 
in the Tropical Forest of the Himalayan foothills, West Bengal. 
2.5. Reproductive biology of dung beetles in the world  
Several features of the biology of dung beetles had been broadly studied and 
critically analyzed in different regions of the world (France, South American countries, 
South Africa, Germany, etc.). Halffter and Edmonds (1982) compiled information on 
the nesting behaviour of subfamily Scarabaeinae and analyzed the relationship 
between ecological conditions, morphology and behaviour of dung beetles. Subsocial 
behaviour of Oniticellus cinctus (Fabricius, 1775) was described by Klemperer (1983) 
from Birmingham and found that the nest chambers of Oniticellus cinctus contained 
twenty brood balls, and the whole period of development took one month. Edwards 
and Aschenborn (1987) observed the nesting biology of Onitis viridulus 
Boheman,1857, O. fulgidus Klug,1855, O. obscurus Lanseberge, 1886,  O. alexis 
Klug, 1835,  O. perpunctatus Balthasar, 1963, O. Caffer Boheman, 1857, O. aygulus 
(Fabricius, 1781), O. tortuosus Houston, 1983, O. deceptor Peringuey, 1901, O. 
uncinatus Klug, 1855,  O. picticollis Boheman 1857 and O. pecuarius Lanseberg, 1875 
from South Africa. Hunter et al., (1991) worked on the life history studies of 
19 
 
Onthophagus medorensis Brown, 1929   and described each stage of its life cycle and 
the whole developmental period took 46 days. A study on the reproductive biology, 
nesting and ontogenetic development of O. stylocerus Graells, 1851 revealed that the 
species was univoltine (Romero and Piera, 1995). Studies on the brood care behaviour 
and nest structure of the dung beetle O. vacca (Linnaeus, 1767) was done by Sowig 
(1996). Hunter et al., (1996) studied the life history of O. depressus Harold, 1871. Sato 
(1997) detailed the nesting, rolling and tunnelling behaviour of large-sized subsocial 
African dung rolling beetle Scarabaeus catenatus (Gerstaecker, 1871). The 
reproductive biology of the Onthophagus incensus (Say, 1835) in Mexico has been 
studied by Martínez et al., (1998). Gonzalez-Vainer and Morelli (1999) studied the 
biology of the dung beetle Onthophagus hirculus Mannerheim, 1829 from Uruguay. 
Palestrini et al., (2002) worked on the reproductive biology of   Onitis belial Fabricius, 
1789 from Morocco, O. anthracinus Felsche, 1907 and O. vanderkelleni Van 
Lansberge, 1886 from Kenya and it was revealed that O. vanderkelleni Lansberge, 
1886 constructed a poorly enlarged nest and laid a greater mean number of eggs, and 
O. belial Fabricius 1798, O. anthracinus Felsche, 1907 spent a more amount of energy 
for the construction of the nest. The longevity cost of reproduction for males and 
females in the dung beetle Onthophagus binodis Thunberg, 1818 was studied in 
Australia by Kotiaho and Simmons (2003). Huerta et al., (2003) analyzed the 
reproductive biology and nesting behaviour of several species of Eurysternus Dalman, 
1824 and observed that Eurysternus, a morphologically quite homogeneous genus, 
showed two distinct types of nesting behaviour. Analysis of the fecundity and 
offspring survival of Copris tripartitus Waterhouse 1875 from Mexico was done by 
Huerta and Bang (2004). The cost of reproduction in Callosobruchus maculatus 
(Fabricius 1775) was studied by Paukku and Kotiaho (2005).The study of reproductive 
20 
 
development and seasonal activity of   Copris ochus Motschulsky, 1860 and C. 
tripartitus Waterhouse, 1875 from Korea was done by Bang et al., (2008) and their 
study revealed that C. ochus and C. tripartitus appeared as univoltine species. A study 
of reproductive activity of Onthophagus granulatus Boheman, 1858 from New 
Zealand by Forgie (2009), showed that O. granulatus was univoltine species. Huerta 
et al., (2010) worked on pre-imaginal stages of the O. incensus (Say ,1835) from 
Mexico. Simmons and Ridsdill-Smith (2011) studied the sexual dimorphism, 
reproductive success and the parental investment of two Onthophagus species O. 
taurus (Schreber, 1759), and O. vacca (Linnaeus, 1767). Nesting behaviour of O. 
incensus (Say ,1835) from Mexico was studied by Huerta and García-Hernández 
(2013). Pérez - Cogollo et al., (2015) studied the life history of O. landolti Harold 
(1880) from Mexico and described its pre-imaginal stages of development. The 
feeding, reproductive, and nesting behaviour of Canthon bispinus (Germar, 1824) was 
studied in Uruguay by González-Vainer (2015). Ocampo and Philip (2017) Studied 
the biology and food relocation behaviour of Eucranium species and compared it with 
South African subgenus Scarabaeus (Pachysoma) Macleay,1821 .Study on nesting 
biology and life history of the dung beetle Onthophagus lecontei Harold, 1871 from 
Mexico by Arellano et al., (2017), revealed that a type 1 pattern of nesting behaviour 
was observed and pairs built one to seven brood masses. This study showed the pre-
nesting period (Feeding) lasted for 16 days, the egg stage for 2 days, the larval period 
for 22 days and the pupal period for 11 days. Studies on the reproductive biology of 
Euoniticellus intermedius (Reiche, 1848) by Martinez et al., (2019) observed that 
development from egg to imago ranged from 25 to 28 days and the lifespan of this 
species was recorded as 30 to 60 days. Hernandez et al., (2020) provided details of the 
21 
 
Feeding and reproductive behaviour of the dung beetle Canthon rutilans cyanescens 
Harold, 1868 from Brazil.  
2.6. Reproductive biology of dung beetles in India  
Very few reports are available on the life history and behaviour of the dung 
beetle species in India. Joseph (1994). Studied the Sexual dimorphism and intra sex 
variations of the giant dung beetle Heliocopris dominus Bates, 1868.  Feeding and 
breeding behaviour of Gymnopleurus gemmatus (Harold, 1871) and Gymnopleurus 
miliaris (Fabricius, 1775) done by Veenakumari and Veeresh (1996b) detailed that 
feeding, ball making and rolling, mating, competition, and predation of two species. 
Life history of two commonly occurring south Indian species, Onthophagus gazella 
(Fabricius, 1787) and O. rectecornutus Lansberge, 1883 were done by Veenakumari 
and Veeresh (1996c). Subsocial behaviour in Copris repertus Walker, 1858 and Copris 
indicus Gillet,1910 was studied by Veenakumari and Veeresh (1997).  Studies on the 
life cycle, ecological role and biology of immature stages of Heliocopris dominus 
Bates, 1868 have been done by (Joseph, 1998, 2003). Gaikwad and Bhawane (2015) 
studied the nidification behaviour of three dung beetle species, Onthophagus catta 
(Fabricius, 1787), Onitis philemon (Fabricius, 1801) and, Liatongus rhadamistus 
(Fabricius, 1775) from Maharashtra and analyzed that Onthophagus catta constructed 
a simple nest composed of a single unbranched vertical gallery, Onitis philemon made 
a simple unbranched numerous and extensive vertical galleries and Liatongus 
rhadamistus have constructed a tunnel just beneath the dung pads up to 9 cm deep. 
Studies on the life cycle and nesting behaviour of dung beetle Onthophagus catta 
(Fabricius, 1787) from Maharashtra by Gaikwad and Bhawane (2016) showed that the 
adult longevity ranged between 42-85 days. Nesting and biology of dung beetle 
22 
 
Scaptodera rhadamistus (Fabricius, 1775) was studied in Maharashtra by Khadakkar 
(2018). Singh et al., (2019) studied the nesting architecture, life cycle, and brood ball 
morphometry of the dung beetle Oniticellus cinctus (Fabricius, 1775) in Dehradun the 
study revealed that the total period for the development of the beetles took one month.  
 
Analysis of the literature revealed that no data is available on the prominent 
dung beetle species, reproductive biology and nesting behaviour in the Agribelts of the 
Malabar Coast region. 
23 
 

METHODOLOGY 
 
 
3.1. Rearing of Dung Beetles  
Adult beetles were collected using dung baited pitfall traps and handpicking 
from the agricultural fields in different regions of the Malabar Coast, namely, an open 
agricultural field consisting of mainly coconut plantation with intervening grasslands close 
to Devagiri College campus (11015’N, 75048’E), Kozhikode district, an open 
agricultural field at Naduvattam (10052'55.92"N, 7600'29.59"E) and paddy field at 
Thavanoor (10.8412°N,75.9938°E) Malappuram district, Kerala (India) and paddy 
field at Kumbidi (10.8337°N,76.0489°E) Palakkad district, Kerala (India), were 
collected during June 2016 to December 2017 period. To collect live dung beetles, 
pitfall traps made of plastic basins, 10 cm in diameter and 15 cm deep with the 
minimum quantity of water to prevent the drowning of the fallen beetles, were placed 
in the field from 8:00 am to 12:00 pm. Preliminary verification, separation, and sexing 
of the collected beetles were done by comparing with verified specimens present in the 
insect collections of St. Joseph’s College, Devagiri, Kozhikode and taxonomic keys in 
Arrow (1931). Based on morphological characters such as small body size and colour, 
beetles of uniform age were selected and grouped. Adult Onthophagus cervus were 
sexed with the male having a pair of horns behind the eyes, slanting backward, wide at 
the base but not united, each bent at a right angle inside just beyond the base, curved 
outward and rapidly narrowed. The clypeus is slightly produced, but truncate and not 
pointed, very shining, lightly punctured with intermixed large and small punctures and 
separated from the forehead by not very strong carina. The clypeus is powerfully and 
closely punctured and not shining, separated from the forehead by a strong carina, and 
24 
 
there is a similar carina between the eyes are present in females Onthophagus fasciatus 
.In male, the clypeus is feebly punctured in the middle and more intensely and closely 
at the sides produced to a point, gently reflexed and divided by a slightly curved carina 
from the sparsely punctured forehead. The posterior margin of the head is produced 
backward and gently curved upward, the median part developing a curved tongue-like 
process at the sides ,a pair of closely parallel horns, which is  wide at the base and 
tapering to the tips. The pronotum is almost vertical and finely and sparsely punctured 
in front. Females, having features such as the clypeus is transversely rugose, the sides 
are convergent, and the front margin is strongly reflexed and nearly straight in the 
middle, the forehead is relatively strongly punctured and separated from the clypeus by 
a strong nearly straight carina  and there is a second strong carina upon the vertex. The 
pronotum bears a well-marked transverse carina in the middle just behind the front 
margin), and Tiniocellus spinipes were sexed based on the pronotum shape (the 
clypeus is short  with its margin rounded and extremely feebly excised in the middle. 
The front tibia is broad with four short sharp external teeth almost at right angles in 
males and the female. The clypeus is slightly produced and distinctly excised at the 
front margin. The front tibia is broad  with very strong external teeth, the terminal one 
.very oblique). Sisyphus longipes males showed remarkable peculiarities in the legs. In 
Tibiodrepanus setosus the head is rather narrow,unevenly and unequally punctured 
with the sides nearly straight and parallel behind. The pronotum has a small anterior 
lateral depression on each side and a big posterior depression, from the middle of the 
latter springs, a slender dorsal horn directed obliquely forward, its extremity with a 
little bifurcate, but the tips scarcely diverging. The males having a horn present.  
Female having the pronotum with a rather large median posterior depression  a smaller 
one in front of it, and an anterior lateral depression on each side (Arrow, 1931). 
25 
 
 Ten mating pairs were selected. Each pair was placed in an individual wide-
mouthed earthen pot with diameter of 51.5 cm, thickness 0.9 cm and length 14 cm 
and filled with finely sieved clay soil collected from the collection site and 
moistened with water by a depth of 13.5 cm and fresh cow dung on top for food and 
the construction of brood balls and each pair were provided with fresh cow dung 
twice a week. The top of the earthen pots was covered with a mesh net (mesh size 
0.053 µm) to prevent the escape of the beetles and the pots were kept at controlled 
room conditions (Temperature 230C-250C; humidity 75%) and in plastic troughs 
containing moist sand. Water was sprayed with a mist sprayer on alternate days to 
prevent desiccation (Fig.2 A-F). Daily observations for all life events, such as brood 
ball formation, egg-laying, egg hatching, duration of the larval and pupal phase and 
adult emergence were noted, and parallel laboratory culture was maintained for 
observing each life cycle stage of the development and also for studying the nest 
architecture. To monitor the life cycle and development of the egg, different stages 
of larval development, pupa and until adult emergence were recorded by making a 
small opening on each brood ball, which was closed by pasting with a layer of dung 
and soil after each observation and the brood masses/balls were retained in individual 
earthen pots arranged with moist soil. Observations were made twice a week until the 
emergence of new adults. The number, length and width of the brood masses, 
number of larvae, pupae and adults, and the size of the adults were recorded. Newly 
emerged beetles were collected, paired and counted, and transferred to new 
individual earthen pots topped with fresh cow dung and were kept until their natural 
death. Adult longevity (after emergence from their brood ball) is known only in 
laboratory-reared specimens and the survival period was noted for each beetle. The 
experiment setup was kept moist by sprinkling water to prevent desiccation. After 
26 
 
two weeks, the earthen pot was opened with care and notes were being made on the 
nest architecture, ball making, and the length of the tunnel was taken. Photographs 
were taken using Nikon digital camera D90 and LEICAS8APO (Trinocular stereo 
zoom microscope).  
3.2. Study of Nesting Pattern  
Adult beetles got from the collection site were placed in a wide-mouthed 
earthen pot. The earthen pot was filled with moist soil and topped with fresh cow 
dung droppings. The top of the earthen pot was covered with a mesh net, after 
introducing the beetles to prevent their escape. These wide-mouthed circular earthen 
pots (16×40cm) were found to be the most suitable for the rearing of roller and 
dweller species. 
Preliminary analysis was done in the field beneath the dung pats to get an 
idea about the tunneling behaviour of tunneling species, brood ball construction, 
nesting preparation and also done by open the tunnels by digging in the agricultural 
field from where the beetle collections were made. For the study of the nest 
architecture of tunneling species, adult beetles got from the collection site, were 
placed in plastic pots (15×1×16 cm) which were cut into half lengthwise and rejoined 
with masking tape to retain their original shape. The rejoined plastic pot was filled 
with moist soil up to a depth of 12 cm and topped with fresh cow dung droppings. 
Beetles were shifted to the pre-arranged plastic pot containing soil and cow dung 
topping. The top of the plastic pot was covered with a mesh net, after introducing the 
beetles to prevent their escape. The experimental setup was kept moistened by 
sprinkling water to prevent desiccation. After two weeks, the rejoined plastic pot was 
opened with care and the notes were being made on the nest architecture, ball 
making and the length of the tunnel was taken (Fig.3 A-E). For data analysis, 
27 
 
(Mean±SD) values are calculated from raw data values with the use of Microsoft 
excel 2010. 
 
 
 
28 
 



RESULTS 
 
      4.1. Life biology of Onthophagus cervus (Fabricius, 1798) 
Life biology involved four stages namely egg, larva, pupa and adult. The egg stage 
lasted for 3.60±0.51 days, the larval stage for 16.70 ±1.87 days, the pupal stage for 
10.20±1.03 days and the adult stage for 60.17± 2.08 days.  
Brood mass and eggs (Fig 4. A-N): Adult beetles constructed brood balls after 
12.4±0.69 days. A single mating pair produced 14.10±5.69 brood balls during its 
period of the life cycle. Oval- shaped brood balls have a length of 20.4±0.97 mm, width 
32.8±1.62 mm and were coated by a layer of soil and dung (Fig 4. A). The brood 
masses were formed of dung mass with an egg chamber with the egg glued to the wall 
of the egg chamber (Fig 4. B). Brood masses were attached to the wall and end of the 
tunnels. Eggs were elongated oval in appearance and creamy white, during the first two 
days. Before hatching (3rd day), the egg became yellowish and the eggshell became 
transparent (3rd and 4th day) and the larva was visible through the chorion. The egg 
stage lasted for 3.6±0.51 days. A single mating pair produced 21.7±6.69 surviving eggs 
during its lifetime. Low egg mortality (14.57%) was recorded (Table.1). 
Larva: Three larval instars (Fig 4. C, D, E) were recorded. Newly emerged larvae were 
transparent with the tips of the mandible being dark brown. Larvae were found in a 
cavity inside a brood ball and they consumed the dung ball from inside. Newly hatched 
larvae were creamy white fleshy “grubs”. All larvae have the characteristic “coprine 
hump” and the flattened, fleshy-lobed anal segment. The larval period lasted 16.7±1.87 
29 
 
days. Low larval mortality (16.12%) was recorded. A single mating pair produced 
18.2±6.58 surviving larvae. 
Pupa: Pupae were present inside the thin-walled pupal cell or cocoon constructed by 
larva inside the brood ball. The inner surface of the pupal cell was smooth and was 
coated with soft dried dung and soil (Fig 4. F). Newly formed pupae were creamy 
white, shiny with four pairs of finger-like processes on the dorsolateral region of the 
abdomen and a large  blunt pronotal projection extending over a posterior portion of 
the head. Later on the pupae turned golden brown (Fig 4. G, H). The pupal period 
lasted for 10.2  ± 1.03 days. Pupal mortality (27.48%) was recorded. A single mating 
pair produced 13.2±4.88 pupae. 
Adult: Teneral period lasted 2.40 ±0.51 days. The teneral adult was light orange-red 
(Fig. 4. I). Adult emerged by cutting a hole in the brood ball (Fig 4. J). 67.42% of 
adults emerged (30 females and 10 males) and the sex ratio of 3:1 was observed. 
Newly formed adults took 1.40±0.52 days for the complete melanization. On exit from 
the brood ball, newly emerged beetles constructed the tunnels. Sexual maturity was 
attained by 11±1.05 days of emergence. Adult male (Fig 4. K) duration of  35.2±8.65 
days and female (Fig 4.L) duration of  60.17± 2.08 days were observed. Egg to teneral 
adults took 28.2 ±1.03 days. A single mating pair produced 4±2.21 surviving adults 
during its lifetime.  
Nesting behaviour:  Adult beetles (males and females), upon releasing, made 
vertical (Fig 4. M) and horizontal tunnels (Fig.4.N). Both males and females were 
involved in tunnel construction and handling of dung. Both vertical and horizontal 
tunnels were made and were interconnected. Vertical tunnels with a depth of 
6.96±1.30 cm and horizontal tunnels  with a length of 2.25±0.59 cm, were observed. 
30 
 
Brood masses were present at the bottom of the tunnels. Brood balls were seen in 
single or in mass.  
4.2. Life biology of Onthophagus fasciatus Boucomont, 1914  
The lifecycle comprises four phases namely egg, larva, pupa, and adult. The egg stage 
took 3.1±0.57 days, the larval stage for 18.1±0.31 days, the pupal stage for 11.1±0.57 
days and the (egg to teneral adult) for 38.5 ±0.52 days. Adult period for 64.1± 2.42 
days.  
  Egg and Brood ball (Fig 5.  A-I ) : 
Egg: Eggs were elongated ovoid in appearance and creamy white, during the first 2 
days (Fig 5. A). Earlier to hatching (3rd day), the egg developed yellowish and the 
eggshell became transparent (3rd and 4th day) and the larva was visible through the 
chorion. The egg period took 3.1±0.57. A single mating pair formed 12.5 ±10.60 eggs 
during its lifetime.  Low egg mortality (13.29 %) was recorded (Table 2).  
Brood ball:  Adult beetles constructed brood balls subsequently 12.2±0.42 days. A 
single mating pair made 13.2±3.68 brood balls throughout its period of the lifecycle. 
Oval-shaped brood balls have a length of 14.4±1.42 mm, width 22.9±2.23 mm, and 
were layered by a film of soil and dung (Fig 5. B) . Brood masses are attached to the 
wall and end of the tunnels. The brood masses are formed of dung mass with an egg 
chamber with the egg attached to the wall of the egg chamber 
Larva: Three larval instars were observed (Fig 5. C, D, E). Newly developed larvae 
were clear with the tips of the mandible being dark brown. Larvae were seen in a cavity 
inside a brood ball and they used up the dung ball from inside. Newly produced larvae 
were creamy white fleshy “grubs”. “Coprine hump” and flattened, fleshy-lobed anal 
segment are the characteristics feature of all larvae. The larval period lasted for 18.1 
31 
 
±0.31 days. Low larval mortality (28.03 %) was recorded.The third segment of the 
abdomen with a setose present.  
Pupa: The newly formed pupae were creamy white, shiny, with large, blunted pronotal 
projection lengthening over the posterior portion of the head and small mesonotal and 
metanotal projections present. Large finger-like lateral tergal projections on segments 
3-6 and  caudal projections callous-like. (Fig 5. F). Pupae were present inside the thin-
walled pupal cell or cocoon made by larva inside the brood ball. The inner surface of 
the pupal cell was smooth and was covered with soft dried dung and soil (Fig 5. G). 
The pupal period lasted for 11.1±0.57 days.  Pupal mortality was recorded at 61.68 %. 
Adult: Teneral adults continued in the pupal cell for 3 days. The teneral adult is light 
orange-red. Adult emerged by cutting a hole in the brood ball 36.84 % of adults 
emerged (25 females and 10 males) and the sex ratio of 5:2 was observed. Newly 
formed adults took 1.4±0.52 days for the complete melanization.  On leaving the 
brood ball, newly emerged beetles constructed the tunnels. Sexual maturity was 
attained by 11±1.05 days of emergence. Adult male (Fig 5. H) duration of 51.4±5.19 
days and female (Fig 5. I) duration of   64.1± 2.42 days were observed. Egg to teneral 
adults took 38.5 ±0.52 days.  
Nesting behaviour: Paired males and females  upon releasing, made vertical and 
horizontal tunnels.  Construction of tunnels and handling of dung is done by both 
males and females, vertical and horizontal tunnels were made. Vertical tunnels with a 
depth of 7.3±0.63 cm and horizontal tunnels with a length of 4.3±0.49 cm, were 
observed. Brood masses were present at the bottom of the tunnels. Brood balls are 
seen in single or in mass.  
 
32 
 
4.3  . Life   biology of Tiniocellus spinipes (Roth, 1851) 
Life biology contains four stages namely egg, larva, pupa, and adult. The egg stage 
lasted for 4.2±0.42 days, the larval stage for 20.6 ±1.26   days and the pupal stage for 
12.3±0.82 days and the developmental period (egg to teneral adult) for 39.2 ± 0.63 
days.  
  Egg   and  brood mass (Fig 6. A-H): 
A single mating pair produced 14.3±5.57 brood balls during its period of one life 
cycle. Oval-shaped brood balls have a length of 29.5±0.70 mm, width 50.5±2.83 mm 
and are covered by a layer of soil and dung. The brood masses are made of dung 
mass with an egg chamber with the egg attached to the wall of the egg chamber. 
Brood masses are produced after 6.6±0.69 days and are attached to the end of the 
tunnels.  
Egg: The egg is usually found adhering to the wall of the brood ball (Fig 6. A). As 
development continues, there is an increase in width, so that the eggs just before 
hatching the chorion is transparent (3rd and 4th day), the dark mandibles and the 
segmentation of the body can be distinguished through it. Eggs were lengthened oval 
in appearance and creamy white, during the first 2 days. The egg stage lasted 4.2 ± 
0.42 days. 24.4 ± 6.91 eggs are produced during their lifetime. Egg mortality 28.27 % 
and egg hatchability 71.72% were recorded. (Table 3). 
 Larva: Three larval instars  (Fig 6. B, C, D)   were recorded. Newly developed larvae 
were transparent with the tips of the mandible being dark brown. Larvae were found in 
a cavity inside a brood ball and they consumed the dung ball from inside. Newly 
hatched larvae were creamy white fleshy “grubs”, “Coprine hump” and the flattened, 
fleshy-lobed anal segment. The larval period lasted for 20.6 ±1.26 days. Larval 
33 
 
mortality (18.85 %) was recorded. A single mating pair produced 14.2±6.52 surviving 
larvae. Larval survivability 81.14% was recorded. 
Pupa: Pupae were present inside the thin-walled pupal cell (Fig 6. E) or cocoon 
constructed by larva inside the brood ball. The internal surface of the pupal cell was 
smooth and was coated with soft dried dung and soil. The newly formed pupae were 
creamy white, shiny, with four pairs of finger-like processes on the dorsolateral area of 
the abdomen and a large, blunt pronotal projection prolonging over a  posterior portion 
of the head. The pupal period lasted 12.3±0.82 days. Pupal mortality was recorded at 
48.59 %. A single mating pair produced 7.3 ±3.27 pupae. Pupal survivability of 
51.40% were observed. 
 Adult: Teneral adults remained in the pupal cell for 2.4 ±0.51 days and 41.09% of 
adults are emerged by making a hole in the brood ball (Fig .6, F). The teneral adult is 
light orange-red in colour. Complete melanization of teneral adults required 1.4±0.52 
days. Sexual maturity was attained by 12.5±0.98 days after emergence. Adult female  
(Fig 6. G) duration of  70.7± 6.42 and male (Fig 6. H)  duration of  72±3.65 days and 
days were recorded. Egg to teneral adults, took 39.2 ± 0.63 days. A single mating pair 
produced 3.±2.05 surviving adults during its lifetime, 41.09% of adult survivability 
were observed (30 females and 20 males ) and the sex ratio of 3:2 was observed. Adult 
mortality (58.90 %) was recorded. 
Nesting behaviour: Adult beetles made shallow vertical tunnels. Both males and 
females were involved in tunnel construction and handling of dung. Vertical tunnels 
with a depth of  9.65±0.74 cm (tunnel length of  2.7±0.49 cm inside the dung pat; 
6.95±0.25 in the soil).  
 
34 
 
4.4.   The life biology of Sisyphus longipes (Olivier, 1789)  
Life biology contains four stages namely egg, larva, pupa, and adult. The egg stage 
lasted for 6.8±1.22 days, the larval stage for 25.3± 0.68 days and the pupal stage for 
15.5± 0.70 days and the developmental period (egg to teneral adult) for 47.3 ±1.63 
days. 
Brood ball construction: (Fig 7. A-J):  Paired beetles built brood balls after 
13.56±2.05 days, A single mating pair created 20.1±5.13 brood balls and food balls 
9.6 ±4.40 during its period of the life cycle. The adult beetles (males and females) 
upon releasing, make a brood ball from the dung pad and it is rolled away from the 
dung. The dung ball is buried with a depth of 3.6±0.69 cm or the sides of the earthern 
pot and some are not buried in the soil. When the brood ball is completely formed 
from the dung and it is rolled away by the pair or by the single adult alone, using the 
long hind limbs with pushing using the back legs and pulling using the front legs and 
fashioned into spherical balls. Food balls are smaller than brood balls or they may be 
of similar size. Spherical-shaped brood balls have a diameter of 12.1±1.51 mm and 
the feeding ball has a width of 6.37 mm. The adult-making brood balls in which eggs 
are laid may be coated with soil. 
Egg: The adult beetle constructed a spherical brood ball (Fig 7. A) and is buried at 
the end of a shallow tunnel underneath the moist soil in the earthern pot and lays an 
egg in the brood ball and eggs are present inside the brood ball. Eggs were 
lengthened oval in appearance and creamy white, soft and yolky. The egg stage lasted 
for 6.8±1.22 days (Fig 7. B). 
 Larva: The larvae present inside the brood ball, (Fig 7. C, D, E) they eat the dung 
from inside the brood ball. The newly formed larvae were translucent, fleshy “grubs”, 
35 
 
C-shaped body, the adjacent area of the mandible with two setae, pronotum with 
separate shields bearing anterior angels; legs with prominent lengthen terminal 
papillae, third abdominal segment absent dorso middle prominence; maxillary 
stridulatory teeth absent. Legs two- or three-segmented. The hindgut enlarged portion 
of larvae stores larval excrement that has grey-brownish paste. The larval period 
lasted for 25.3±0.68 days. 
Pupa: The newly formed pupae are shiny, creamy white, with numerous projections 
present on its dorsal surface, finger-like processes are detected on the dorsolateral 
region of the abdomen and pronotal projections absent, mesonotal and metanotal 
projections present, adjacent tergal projections finger-like, present on the abdomen 
(Fig 7. F). The pupal period lasted for 15.5± 0.70 days in the laboratory. Pupae are 
present inside the pupal cell (Fig 7. G) formed by the larva. Pupal survivability 44% 
were recorded (Table.4). 
Adult: Adults are formed in the larval chamber prepared by larvae, they construct a 
hole in the chamber and the adults emerged  (Fig 7. H) from the chamber after 15.5± 
0.70 days of pupation , 44 % of adults arisen from the brood ball in which 
survivability of adults is 40.90%  were observed  (9 females and  9 males) and the sex 
ratio 1:1 were observed. The newly formed immature adults stay in the dung and the 
sexual maturity was attained after 14.5±0.52 days. Adult male (Fig 7. I) duration of  
33±7.52 days and female (Fig 7. J) duration of 42.5± 2.63 days were observed in the 
laboratory.The developmental period from egg to teneral adults took 47.3 ±1.63 days 
were observed. 
Nesting behaviour:  Nest provision and brood mass creation are the symbols of the 
following stage in the breeding process. The adult beetles (males and females), built 
36 
 
a brood ball from the dung pad  rolled away from the dung mass  and the eggs were 
positioned in the brood ball  buried in the pit with the depth of 3.6±0.69 cm     
4.5.     Life biology of Tibiodrepanus setosus (Wiedemann, 1823)  
Biology of Tibiodrepanus setosus: (Fig 8. A-I):  Life biology involves four stages 
namely egg,   larva, pupa, and adult. The egg stage lasted for 3.9 ±0.31 days, the larval 
stage for 15.4±1.07 days and, the pupal stage for 8.6 ±0.51 days and egg to teneral 
adult 30 ±0.44 adult stage for 50.9±6.79 days   
Brood mass and Egg (Fig 8. A):  Adult beetles laid eggs  using much drier dung after 
7.5±0.52 days. A single mating pair produced 10.2 ±1.54 eggs during its period of the 
life cycle. The eggs are laid directly in the food source  with the eggs are present in the 
dung. 
Egg: Eggs were elongated oval in appearance and creamy white, during the first 2 
days (Fig 8. A). Before hatching (3rd day), the egg became yellowish and the eggshell 
became transparent (3rd and 4th day) and the larva was visible through the chorion. 
The egg stage lasted 3.9 ±0.31. A single mating pair produced 10.2 ±1.54 eggs during 
its lifetime. Low egg mortality of 13.72% was recorded (Table.5).  
Larva: Three larval instars, (Fig 8 .B,C,D) were recorded. Larvae were found in a 
cavity inside a brood ball. Newly hatched larvae were creamy white fleshy “grubs”. All 
larvae have the characteristic “Coprine hump”.  Most are c-shaped, white fleshy 
“grubs” with a well sclerotised head capsule and well-developed legs. The larval period 
lasted for 15.4 ±1.07 days. Low larval mortality of   22.72 % was recorded.  
Pupa: Pupae were present inside the thin-walled pupal cell constructed by larva inside 
the brood ball. (Fig 8 .E). The inner surface of the pupal cell was smooth and was 
37 
 
coated with soft dried dung and soil. The newly formed pupae were creamy white, 
shiny, with four pairs of finger-like processes on the dorsolateral region of the 
abdomen and a large, blunt pronotal projection lengthening over a posterior portion of 
the head (Fig 8. F). Later on, the pupae turned golden brown. The pupal period lasted 
for 8.6 ±0.51 days. Pupal mortality was recorded at 47.05 %. 
Adult: Teneral adults (Fig 8. G) remained in the pupal cell for 3.1±0.31 days. Adult 
emerged by cutting a hole in the pupal cell.  Adults emerged 55.55 % (12 females and 
8 males) and the sex ratio of   3:2  was observed. Newly formed adults took 1.3 ±0.49 
days for the complete melanization. Sexual maturity was attained by 12.4±0.51days of 
emergence. Adult male (Fig 8. H) duration of 50.9±6.79 days and female (Fig 8. I) 
duration of  50 ± 8.01 days were observed. Egg to teneral adults took 30 ±0.44 days  
Nesting behaviour 
Beetles use plenty of drier dung for building brood masses and each brood mass has 
one egg. Type 4 nests were observed, the beetles do not form nesting chambers or 
burrows, and the whole development within the dung pat itself.  
 
 
 
38 
 
Table 1. Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Onthophagus cervus in the agribelts of 
the Malabar Coast region. 
 
  Hatchability/ 
Parameters Mean ±SD Mortality/ Survivability (%) 
Fecundity 25.4±6.67 - 
Egg hatchability 21.7±6.69 85.43 
Egg mortality 3.7±2.31 14.57 
Larval survivability 18.2±6.58 83.88 
Larval mortality 3.5±1.50 16.12 
Pupal survivability 13.2±4.88 72.52 
Pupal mortality 5±2.62 27.48 
Adult survivability 4±2.21 30.30 
Adult mortality 9.2±2.57 69.69 
 
 
 
 
39 
 
Table 2.  Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Onthophagus fasciatus in the  
agribelts of the Malabar Coast region.  
 
  Hatchability/ 
Mortality/ Survivability (%) 
Parameters Mean ±SD 
Fecundity 12.5± 10.60 - 
Egg hatchability 13.7±4.28 86.7 
Egg  mortality 2.1±1.29 13.29 
Larval  survivability 10.7±3.49. 71.97 
Larval  mortality 3.0±0.94 28.03 
Pupal  survivability 4.1±0.88 38.31 
Pupal  mortality 6.6±3.40 61.68 
Adult survivability 2.4±0.84 58.53 
Adult mortality 1.5±0.70 41.47 
 
 
 
 
 
 
 
 
 
40 
 
Table  3. Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Tiniocellus spinipes in the  
agribelts of the Malabar Coast region. 
 
  Hatchability/ 
Mortality/ Survivability(%) 
Parameters  Mean±SD 
Fecundity  24.4 ± 6.91 - 
Egg hatchability 17.5±6.68 71.72 
Egg  mortality 6.9±3.38 28.27 
Larval  survivability 14.2±6.52 81.14 
Larval  mortality 3.3±0.82 18.85 
Pupal survivability  7.3±3.27 51.40 
Pupal mortality 6.9±3.51 48.59 
Adult survivability  3±2.05 41.09 
Adult mortality  4.3±1.77 58.90 
 
 
 
 
 
 
 
41 
 
Table  4. Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Sisyphus longipes in the agribelts of 
the Malabar Coast region. 
 
  Hatchability/ 
Parameters Mean ±SD Mortality/ Survivability 
(%) 
Fecundity 15±5.66 - 
Egg hatchability 12.8±4.75 85.33 
Egg mortality 2.2±1.61 14.67 
Larval survivability 10±4.88 78.12 
Larval mortality 5.6±2.50 21.88 
Pupal survivability 4.4±2.83 44 
Pupal mortality 5.6±2.50 56 
Adult survivability 1.8±0.79 40.90 
Adult mortality 2.6±2.17 59.09 
 
 
 
42 
 
Table 5.   Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Tibiodrepanus setosus in the  
agribelts of the Malabar Coast region. 
 
  Hatchability/ 
Mortality/Survivabilit
Parameters Mean ±SD y (%) 
Fecundity  10.2 ±1.54 - 
Egg hatchability 8.8±1.48 86.27 
Egg  mortality 1.4±0.51 13.72 
Larval  survivability 6.8±1.39 77.28 
Larval  mortality 2±1.69 22.72 
Pupal  survivability 3.6±0.97 52.94 
Pupal  mortality 3.2±1.69 47.05 
Adult survivability 2±0.81 55.55 
Adult mortality 1.6±0.51 44.44 
 
43 
 
Table  6. Number and size of brood balls and duration (days) of different life cycle stages of various Onthophagus     
                               species (Author details are provided in parenthesis; ND: no data available) 
 Number of Brood ball Egg Larval Pupal Teneral Total duration Adult 
Species brood balls (Incubatio duration duration adult days (Egg to longevity 
 Length in Width in n period) teneral adult) 
mm mm days 
O.  cervus 14.1±5.69 20.4±0.97 32.8±1.62 3.6±0.51 16.7 ±1.87 10.2±1.03 2.4±0.51 28.2 ±1.03 60.17± 2.08 
(present study) 
      
O. catta          
(Gaikwad and Bhawane, 22.5±17.67 27.7±3.79 5.4±1.49 2.38±0.8 31.5 ±6.37 13.46±0.8 3.5±0.70 48.33±4.49 66.7±11.98 
2016) 
O. gazella          
(Veenakumari and Veeresh, 6 40.6±0.03 16.0±0.08 5.4±0.54 26.2±1.22 11.16±0.98 3.5±0.70 41.4±2.60 ND 
1996c ) 
O. rectecornutus          
Veenakumari and    12±2.5 34.7±0.33 11.3±0.11 4.0±0.47 19.0±2.00 10.88±1.05 4 31.8±1.93 ND 
Veeresh,1996c) 
O. lecontei          
(Arellano et al., 2017) 3.50±1.74 23.47±1.52 23.14±0.91 2 22±1.14 11±0.87 4±0.95 39 60±2.3 
O. incensus          
(Huerta et al., 2010) 5± 5.65 25±7.07 12.5±3.53 4 22 10±2.82 ND 36 ±2.82 93 
O. landolti          
(Perez cogollo et al., 2015) 14.5±13.43 ND ND 2.2 ±0.70 21±1.41 7±1.41 ND 30 60 
O. medorensis          
(Hunter et al., 1991) ND 10.27±4.63 ND 4 28 11.5±0.70 4±1.41 49.5 ±4.94 53±26.88 
O. depressus          
(Hunter et al., 1996) ND 22±4.24 16±1.41 3.4±1.28 27 12 3 46.5±14.84 50 
O. stylocerus 
(Romero and Piera,1995) 19.75±2.16 31.5±9.19 15.5±3.53 7.5±3.53 22.33±3.05 14±4.24 15 60.5±13.43 ND 
44 
 
 
Table 7:  Number and size of brood balls and duration (days) of different life cycle stages of various Onthophagus 
species. 
(Author details are provided in parenthesis; ND: no data available) 
 Number of Brood ball Egg Larval Pupal Teneral Total Adult 
Species brood balls (Incubatio duration duration adult duration longevity 
 Length in Width in n period) days (Egg to 
mm mm teneral 
adult) days 
O.  fasciatus 13.2±3.68 14.4±1.42 22.9±2.23 3.1±0.57 18.1 ±0.31 11.1±0.57 3 38.5 ±0.52 64.1± 2.42 
(present study) 
       
O. catta          
(Gaikwad and Bhawane, 2016) 22.5±17.67 27.7±3.79 5.4±1.49 2.38±0.8 31.5 ±6.37 13.46±0.8 3.5±0.70 48.33±4.49 66.7±11.98 
O. gazella          
(Veenakumari and 6 40.6±0.03 16.0±0.08 5.4±0.54 26.2±1.22 11.16±0.98 3.5±0.70 41.4±2.60 ND 
Veeresh, 1996c ) 
O. rectecornutus          
(Veenakumari and 12±2.5 34.7±0.33 11.3±0.11 4.0±0.47 19.0±2.00 10.88±1.05 4 31.8±1.93 ND 
Veeresh,1996c) 
O. lecontei          
(Arellano et al., 2017) 3.50±1.74 23.47±1.52 23.14±0.91 2 22±1.14 11±0.87 4±0.95 39 60±2.3 
O. incensus          
(Huerta et al., 2010) 5± 5.65 25±7.07 12.5±3.53 4 22 10±2.82 ND 36 ±2.82 93 
O. landolti          
(Perez cogollo et al., 2015) 14.5±13.43 ND ND 2.2 ±0.70 21±1.41 7±1.41 ND 30 60 
O. medorensis          
(Hunter et al., 1991) ND 10.27±4.63 ND 4 28 11.5±0.70 4±1.41 49.5 ±4.94 53±26.88 
O. depressus          
(Hunter et al., 1996) ND 22±4.24 16±1.41 3.4±1.28 27 12 3 46.5±14.84 50 
O. stylocerus 
(Romero and Piera,1995) 19.75±2.16 31.5±9.19 15.5±3.53 7.5±3.53 22.33±3.05 14±4.24 15 60.5±13.43 ND 
 
 
45 
 
 
Table  8. Number and size of brood balls and duration (days) of different life cycle stages of various sub-tribe 
Oniticellina species 
  Brood ball Egg Larval Pupal Teneral Total duration Adult longevity 
 Number of incubation duration duration adult days (egg to teneral 
Species brood balls period adult) days 
Length in Width in  
mm mm 
T. spinipes 96.33±1.3 29.5±0.70 50.5±2.83 4.2±0-42 20.6 ±1.26 12.3±0.82 2.4 ±0.51 39.2 ± 0.63 72±3.65 
(Roth,1851) 
(Present study) 
E. intermedius 111±15 - - 1.5±0.70 31 28 - 56 45± 21.21 
(Reiche,1848) 
L. rhadamistus - - 20.5±1.28 3.94±0.7 30±32.52 17±1.41 5.5±0.70 69.14±6.17 80.45±14.95 
(Fabricius,1775) 
 
 
 
46 
 
Table 9.  Number, shape and size of brood balls and duration (days) of different life stages of Sisyphus species. 
 Brood ball Duration of different life stages Adult number Adult longevity 
Species No Shape Diameter Egg Larva Pupa Egg adult Male Female Summer Winter 
Number Duration 
Summer Winter 
S. longipes 20.1±5.1 Sphere 12.1±1.5 15±5.66 NA 6.8±1.22 25.3±0.68 15.5± 0.70 47.3±1.63 9 9 42.5±2.63  
(Present study ) 3 1 
S. sordidus - Dome on - 15±7.07 - 8 - - 66.2 - - - 300-700 
 sphere 
S. seminulum - Dome on - 26.0 17.5 6.5±2.12 - - 47.1 - - 125.2 186 
 sphere 
S. mirabilis - Sphere with 16.2 46.8 46.2 9±1.41 - - 77.3 175 175 144.1 214.6 
very short tip 
S. fortutus - Sphere with tip 17.1 54.5 57.5 12±2.82 - - 73.2 54 68 153.5 201.0 
S. spinipes - Sphere with 17.6 43.7 34.5 8.5±0.70 - - 51.8 108 129 104.2 153.2 
 long tip 
S. infusticatus - Sphere with tip 13.6 56.4 36.2 9±1.41 - - 53.7 112 120 114.1 127.3 
S. rubrus 40 Sphere with 14.7 36.4 24.8 - - - 64 131 136 116.8 148.6 
flat dome 
S. calcaratus 48 Orange 10.6 41.2 25.5 7.5±0.70 - - 59.7 160 178 133.6 221.0 
S. muricatus - Orange - 14  - - - 58 4 4 395 - 
S. fasciculatus - Orange - 18 - - - - 60 18 21 395 - 
 
S. barbarossa - Orange - - - - - - -  9 9 - 
 
S. tibialis - Sphere - - - - - - - - - - - 
 
47 
 






DISCUSSION 
 
   The present study provides data on the reproductive biology and nesting 
behaviuor of the five prominent dung beetle species Onthophagus cervus, O. fasciatus 
, Tiniocellus spinipes, Sisyphus longipes, and Tibiodrepanus setosus in the agribelts of 
the Malabar Coast region and the basic reasons for their abundance and dominance in 
the region. 
5.1. Onthophagus cervus (Fabricius, 1798). 
Comparison of data of brood mass production, fecundity, duration of egg, 
larval, pupal, adult stages, adult mortality, and life span of Onthophagus cervus with 
other Onthophagus species (listed out in Table.6) revealed that a broad categorization 
of Onthophagus species based on the life cycle characteristics is possible. Data on the 
brood mass production of different Onthophagus species showed that Onthophagus 
species can be categorized as high and low brood mass producers. Onthophagus 
stylocerus Graells,1851 (Romero and Piera, 1995); O. rectecornutus Lansberge,1883 
(Veenakumari and Veeresh, 1996c); O. landolti Harold,1880 (Pérez-Cogolloet al., 
2015); O.catta (Fabricius,1787) (Gaikwad and Bhawane, 2016), and O. cervus 
(Fabricius,1798) comes under the category of high brood mass producers with a brood 
mass range of 1– 40 and; O. hirculus Mannerheim,1829 (Gonzalez- Vainer and 
Morelli, 1999); O. incensus Say,1835 (Huerta and Hernandez, 2013); O. lecontei 
(Harold,1871) (Arellano et al., 2017) falls under the category of low brood mass 
producers with a brood mass range of 1–10. Similarly based on the size of brood ball,  
two categories of Onthophagus species are recognizable with a large-sized brood ball 
category consisting of, O. stylocerus (Romero and Piera, 1995); O. rectecornutus 
(Veenkumari and Veeresh, 1996c); O. catta (Gaikwad and Bhawane, 2016) and small 
48 
 
brood ball category of O. medorensis Brown,1929 (Hunter et al., 1991); O. depressus 
Harold,1871 (Hunter et al., 1996); O. hirculus Mannerheim,1829 (Gonzalez –Vainer 
and Morelli, 1999); O. lecontei (Arellano et al., 2017) and O. cervus. 
Duration of egg incubation revealed a pattern of longer egg incubation period 
in O. medorensis (Hunter III et al., 1991); O. stylocerus (Romero and Piera, 1995); O. 
depressus (Hunter et al., 1996); O. rectecornutus (Veenakumari and Veeresh, 1996c); 
O. hirculus (González-Vainer and Morelli, 1999); O. incensus (Huerta et al., 2010), O. 
cervus  and short egg incubation period in O. landolti (Pérez-Cogolloet al., 2015); O. 
catta (Gaikwad and Bhawane, 2016) and in O. lecontei (Arellano et al., 2017). 
Comparison of larval duration showed that O. cervus and O. rectecornutus 
(Veenakumari and Veeresh, 1996) belong to the shorter larval duration category 
compared to O. medorensis (Hunter et al.,1991); O. stylocerus (Romero and Piera, 
1995); O. depressus (Hunter et al., 1996); O. incensus (Huerta and Hernandez, 2013); 
O. landolti (Pérez-Cogolloet al., 2015); O. catta (Gaikwad and Bhawane, 2016); and 
O. lecontei (Arellano et al., 2017) with long larval duration period. 
Comparison of pupal duration among the various Onthophagus species show 
that O. landolti (Pérez-Cogolloet al., 2015) has a short pupal period compared to 
longer pupal duration in O. medorensis (Hunter et al., 1991); O. stylocerus (Romero 
and Piera,1995); O. depressus (Hunter et al., 1996); O. rectecornutus (Veenakumari 
and Veeresh, 1996c); O. catta (Gaikwad and Bhawane, 2016); O. cervus and O. 
lecontei (Arellano et al., 2017). Higher variability in egg hatchability, larval and pupal 
survivability under uniform conditions in many samples indicate that wider variation 
exists in the population and the exact reasons are not understood and could be genetic. 
The developmental period of O. cervus (egg to a teneral adult) and Mexican 
species O. landolti (Pérez-Cogolloet al., 2015) was the shortest among the various 
49 
 
Onthophagus species. The teneral adult period was shorter in O. cervus compared to 
other Onthophagus species. Comparison of adult duration showed that O. cervus and 
O. medorensis (Hunter et al., 1991); O. depressus (Hunter et al., 1996); O. landolti 
(Pérez-Cogollo et al., 2015); O. lecontei (Arellano et al., 2017); were species with 
short adult longevity whereas,O. stylocerus (Romero and Piera,1995); O. rectecornutus 
(Veenakumari and Veeresh, 1996c); O. incensus ((Huerta and Hernandez 2013); and 
O. catta (Gaikwad and Bhawane, 2016); were with longer adult duration. Low pupal 
survivability compared to the high egg hatchability, larval survivability, and adult 
survivability of O. cervus indicated that the pupal phase is the crucial phase in the life 
cycle of O. cervus. 
Type 1 pattern of nesting was present in Onthophagus cervus with a simple, 
shallow tunnel with a bottom containing brood masses and vertical and horizontal 
tunnels (Halffter and Edmonds, 1982). Similar type 1 pattern was reported in O. taurus 
(Fabre, 1918); O. fucatus (Main, 1922); O. coenobita (Burmeister, 1930); O. catta 
(Fabricius,1787) (Gaikwad and Bhawane, 2016); and O. lecontei Harold,1871 (Arellano 
et al.,2017). Some Onthophagus species constructed compound nest (Type 2) with 
galleries that may have one or more branches, which ended into brood cells in 
O.nuchicornis Linnaeus,1758,  O. fracticornis (Burmeister, 1930), O. medorensis 
(Hunter et al., 1991), O. stylocerus (Romero and Piera, 1995), O. rectecornutus 
(Veenakumari and Veeresh, 1996c)  and O. incensus (Huerta and Hernandez 2013). 
Among the tunneling species, large species tend to bury their brood balls at a 
deeper depth and small species at a shallower depth, which is thought to help reduce 
overall competition for nesting space (Hanski, 1991a; Rougon and Rougon, 1991; 
Hernández et al., 2011). Tunnels were dug roughly perpendicular to the interface 
between soil and dung, resulting in interference competition for nesting space 
50 
 
underneath dung pads, especially in areas where tunnels branch out into nesting 
chambers (Halffter and Edmonds, 1982; Hanski, 1991b; Macagno et al., 2016). Higher 
longevity of females and female-biased sex ratio were seen in O. cervus. Why females 
live longer than male is generally unknown, either metabolic differences or differences 
in patterns of resource allocation between males and females probably account for the 
gender difference in lifespan (Fox et al., 2003). Alternatively, males may allocate a 
more significant proportion of their biomass to reproduction, or allocate those resources 
sooner, such that they become resource-stressed at a younger age. Gender-difference in 
energy expenditure explains at least some of the gender-difference in lifespan. Some of 
the difference in lifespan and mortality rates between genders is due to faster energy-
water loss in males than in females (Fox et al., 2003).  
Observed sex ratio bias in O. taurus females, is caused by the higher mortality of 
males and suggested that this might be linked to higher demand for nutritional resources 
during offspring development (House et al., 2011). Differential mortality is common in 
species like dung beetles with both the sexes having distinct nutritional requirements and 
energy expenditures due to differential mobility and investment in parental care (Veran 
and Beissinger, 2009). Evaluation of the cost of male production studies with other 
groups (Jokela et al., 1997; Wolinska and Lively, 2008; Macagno et al., 2019), have 
suggested a cost of producing males. Also, as per LMC (local mate competition), a 
female-biased sex ratio is favored if the mating competition takes place between male 
offspring (Hamilton, 1967), whereas an equal sex ratio is expected under random mating 
(Fisher, 1930). Hence the female-biased sex ratio noticed in O. cervus indicates that 
mating competition takes place between male offspring and the high cost of producing 
males might have led to the reduction in the ratio of males to females in O. cervus. 
51 
 
  With traits that are common in an r-selective species such as high fecundity, 
multivoltine, small body size, low egg mortality, shorter larval duration, early maturity 
onset, and shorter developmental period (short generation time enables attaining 
maturity earlier together with female-biased sex ratio), longer duration of females 
(favoring high egg production) and shallow tunnels (which enable easy and fast 
tunneling process and development in thin soil topsoil layer) all of these morphological 
and physiological traits contribute to the higher abundance of O. cervus and make it the 
prominent dung beetle species in the agribelts of  Malabar Coast in south India. 
5.2   Onthophagus fasciatus Boucomont, 1914 . 
    The current study showed that the nesting pattern presented by Onthophagus 
fasciatus was Type 1 with a simple, shallow tunnel with a bottom containing brood 
masses and with the tunnels not going deeper into the soil and creating vertical and 
horizontal tunnels (Halffter and Edmonds, 1982).  Similar type 1 pattern was reported 
in O. catta (Gaikwad and Bhawane, 2016); O. lecontei (Arellano et al., 2017); O. 
coenobite (Burmeister1930); O. tauruss (Fabre, 1918); and O. fucatus (Main, 1922). 
Some Onthophagus species constructs compound nest (Type 2) with galleries that 
may have one or more branches, which ends into brood cells as in O. gazelle and O. 
rectecornutus (Veenakumari and Veeresh, 1996c); O. nuchicornis and O. fracticornis 
(Burmeister, 1930); O .medorensis (Hunter et al., 1991); O. incensus (Huerta et al., 
2010); and O. stylocerus (Romero and Piera 1995). 
                 The current study provides the first-time data on the nesting behaviour and 
life history of Onthophagus fasciatus and also enabled comparison of data with other 
Onthohpagus species (details listed out in Table.7). Comparison of the life cycle and 
brood mass construction, brood ball size, fecundity, duration of egg,  larval, pupal and 
adult mortality, duration of adult phase, and life span of various  Onthophagus species 
52 
 
revealed the presence of the following categories. Based on the brood mass production 
Onthophagus species can be categorized as high and low brood mass producers. 
Onthophagus fasciatus, O. catta (Gaikwad and Bhawane, 2016), O. rectecornutus 
(Veenakumari and Veeresh, 1996c), O. stylocerus (Romero and Piera,1995)  and O. 
lentolti (Perez-cogollo et al.,2015)  comes under the category of high brood mass 
producers with a brood mass range of 1- 40. O. gazelle (Veenakumari and 
Veeresh,1996c), O. incensus (Huerta et al., 2013),  O. lecontei (Arellano et al., 2017),  
and  O. hirculus (González-Vainer and Morelli, 1999) falls under the category of mass 
producers with a brood mass range of 1-10. Similarly based on the size of brood ball 
two categories of  Onthophagus species are recognizable with a large-sized brood ball 
category consisting of O. gazella, O. rectecornutus (Veenkumari and Veeresh, 1996c),  
O. stylocerus (Romero and Peira, 1995),  O. catta (Gaikwad and Bhawane, 2016)  and 
small brood ball category consisting of  O. lecontei (Arellano et al., 2017),  O. 
depressus (Hunter et al.,1996),  O. fasciatus, O. hirculus (González-Vainer and 
Morelli, 1999) and O. medorensis (Hunter et al., 1991). 
Based on the duration of egg incubation, a pattern of longer egg incubation 
period as in O. stylocerus (Romero and Peira, 1995),  O. rectecornutus (Veenakumari 
and Veeresh, 1996c),  O. gazella (Veenakumari and Veeresh, 1996b), O. incensus 
(Huerta et al., 2010), O. medorensis (HunterIII et al., 1991),  O. depressus (Hunter et 
al., 1996),  O. fasciatus, and  O. hirculus (González-Vainer and Morelli, 1999) and 
short egg incubation period in O. lecontei (Arellano et al., 2017),  O. catta (Gaikwad 
and Bhawane, 2016),  and in O. landolti (Pérez-Cogollo et al., 2015) is distinct. 
Comparison of larval duration shows that O. fasciatus and O. rectecornutus 
(Veenakumari and Veeresh, 1996c)  belongs to the shorter larval duration category 
compared to O. catta (Gaikwad and Bhawane, 2016),  O. lecontei (Arellano et al., 
53 
 
2017),  O. incensus (Huerta et al., 2010),  O. landolti (Pérez –Cogollo et al.,  2015),  
O. gazelle (VeenaKumari and Veeresh, 1996),  O. insensus (Huerta et al., 2015),  O. 
medorensis (Hunter et al.,1991,  O. stylocerus (Romero and Peira,1995) and O. 
depressus (Hunter et al.,1996)  with long larval duration. 
Comparison of pupal duration of various Onthophagus species shows that a 
short pupal period is present in O. landolti (Pérez-Cogollo et al., 2015)  compared to 
the longer pupal duration of O. stylocerus (Romero and Peira, 1995),   O. catta 
(Gaikwad and Bhawane, 2016),  O .gazella (Veenakumari and Veeresh, 1996),  O. 
lecontei (Arellano et al., 2009),  O. medorensis (Hunter et al., (1991),  O. depressus 
(Hunter et al.,1996), O. fasciatus and O. rectecornutus (Veenakumari and 
Veeresh,1996c). High pupal mortality shows that the pupal phase as the crucial phase 
in the life cycle of O. fasciatus. 
The developmental period of O. fasciatus (egg to a teneral adult) is closer to 
the developmental period in O. incensus (Huerta et al., 2010), O. lecontei (Arellano et 
al.,2017),  O. rectecornutus (Veenakumari and Veeresh, 1996) and the mexican 
species O. landolti (Perez cogollo et al., 2015)  is having the shortest developmental 
period. Comparison of adult duration shows that O. incensus (Huerta et al., 2010), O. 
rectecornutus (Veenakumari and Veeresh, 1996c), O. catta (Gaikwad and Bhawane, 
2016), O. stylocerus (Romero and Piera,1995), and O. gazelle (Veenakumari and 
Veeresh, 1996c) are species with longer adult duration and O. landolti (Pérez –
Cogollo et al., 2015),  O. lecontei (Arellano et al., 2017),  O. medorensis (Hunter et 
al.,1991),  O. depressus (Hunter et al., 1996)  are species with short adult longevity.  
Higher longevity of females and sex ratio biased towards females seen in O. 
fasciatus. Why females live longer than male is generally unknown, either metabolic 
54 
 
differences or differences in patterns of resource allocation between males and females 
probably account for the gender difference in lifespan (Fox et al., 2003).  
The female-biased sex ratio noticed in O. fasciatus indicates that mating 
competition takes place between male offsprings and the high cost of producing males 
might have led to the reduction in the ratio of males to females in O. fasciatus. Traits  
such as low egg mortality, shorter larval duration, and shorter developmental period, 
short generation time, female-biased sex ratio, longer duration of females favouring high 
egg production lead to the higher abundance of O. fasciatus in the agribelts of the 
Malabar coast region.  
5.3 .  Tiniocellus spinipes (Roth, 1851). 
The reproductive biology and life span of a member species Tiniocellus 
spinipes of the genus Tiniocellus of soil tunnelling sub-tribe Oniticellina of tribe 
Onitcinellini and dominant in the regional landscape in South India is analyzed.  
Additionally, the present study enabled a comparison of the life biology of 
representative species of the related soil tunneling genera of the subtribe Oniticellina 
namely, Tiniocellus, Euoniticellus, and Liatongus. Compared the present data on the 
brood mass production,  fecundity, egg, larval, pupal,  adult duration, adult mortality 
and life span of  Tiniocellus spinipes with available data on the soil tunneling genera 
of the subtribe Oniticellina species Euoniticellus intermedius (Reiche,1848) and 
Liatongus rhadamistus (Fabricius,1775).  The present study revealed that Tiniocellus 
spinipes is having the shortest larval, pupal, teneral adult, mature adult duration 
(details listed out in Table.8). Short egg to teneral adult duration and high brood mass 
production compared to Liatongus rhadamistus (Gaikwad and Bhawane, 2015) and 
Euoniticellus intermedius (Martinez et al., 2019) are taken as part of the reproductive 
strategy of Tiniocellus spinipes and is favouring the population build-up of the 
55 
 
species. Advantages of the longer egg incubation period in Liatongus rhadamistus 
(Gaikwad and Bhawane, 2015) and Tiniocellus spinipes, and the short egg incubation 
period in Euoniticellus intermedius (Martinez et al., 2019) are not understood. Egg 
incubation period ranging from 2-5 days is observed as a general pattern in many 
highly abundant dung beetles such as Onthophagus gazella, O. rectecornutus 
(Veenakumari and Veeresh.,1996c), and Oniticellus cinctus  (Singh et al., 2019). 
Among the various life cycle stages the high pupal mortality of Tiniocellus 
spinipes is the crucial phase adversely affecting the life cycle of T. spinipes. Present 
data shows that T. spinipes is a soil-tunneling species producing a simple, shallow 
tunnel with bottom containing brood masses (Cambefort and Lumaret, 1983) and have 
high fecundity and is a multivoltine species. Differential mortality is common in dung 
beetles with both the sexes having distinct nutritional requirements and energy 
expenditures as a result of differential mobility and investment in parental care (Veran 
and Beissinger, 2009). The longevity of females and males in T. spinipes indicates the 
absence of differential mobility and the possible presence of an investment in parental 
care by both sexes. 
 Sex ratio biased towards females is seen in Tiniocellus spinipes. Evaluation of 
the cost of male production studies in other groups (Jokela et al., 1997; Wolinska et 
al., 2008; Macagno et al., 2019), have suggested a cost of producing males. Also, as 
per LMC (Local mate competition), a female-biased sex ratio is favoured, if the 
mating competition takes place between male offsprings, whereas an equal sex ratio is 
expected under random mating (Fisher et al., 1930). Hence, the female-biased sex 
ratio noticed in T. spinipes indicates that mating competition occurs between male 
offspring. 
 
56 
 
 
 
5.4.   Sisyphus longipes (Olivier, 1789).  
The present study showed that Sisyphus longipes make spherical brood balls 
as reported for  South African species S. tibialis Raffray 1877 (Paschalidis, 1974). 
Spherical brood balls of  S.longipes differs from the dome and sphere-shaped brood 
ball in S. seminulum, Gerstaecker 1871, S. costatus Hunberg, 1818,  S. sordidus 
Boheman, 1857, S. caffer Boheman, 1857, the pear-shaped brood ball in S. 
impressipennis Lansberge, 1886 and the sphere-shaped with a flat dome in  S. rubrus 
and S. macrorubrus (Paschalidis, 1974),  sphere-shaped with a very short tip in S. 
mirabilis Arrow 1927 and sphere with the long tip in S. spinipes Thunberg 1818 
(Paschalidis, 1974).  The diameter of brood ball of S. longipes is similar to the brood 
ball in other Sisyphus species, S. rubrus, S. infusticatus and S. calcaratus. 
 Comparison of data (details listed in Table. 9) on brood mass production in 
Sisyphus longipes showed that a lower number of brood balls are produced compared 
to other species,  S. rubrus and S. calcaratus (Paschalidis, 1974). Duration of egg 
incubation revealed a pattern of longer egg incubation period in S. fortuitus Peringuey 
1901, S. infuscatus Klugg, 1855 and S. mirabilis (Paschalidis, 1974) and short egg 
incubation period in S. sordidus (Boheman, 1857), S. seminulum (Gerstaecker, 1871), 
S. calcaratus (Klug, 1855), (Paschalidis, 1974) and in S. longipes. In S. seminulum 
(Gerstaecker, 1871) and S. sordidus (Boheman, 1857), one nest contains a single egg 
(Paschalidis, 1974) and the same pattern was observed in the present study. 
Paschalisidis (1974) reported that in Sisyphus species three larval instars can take less 
than 30 days to complete in the summer season and the same pattern with similar 
larval duration was observed in Sisyphus longipes. 
57 
 
Compared to the pupal duration in other Sisyphus species (Sisyphus sordidus, 
S. mirabilis, S. fortuitus and S. fasciculatus), the pupa transformed into an adult within 
2-3 weeks as reported in the previous studies of (Paschalidis, 1974). High pupal 
mortality was observed in the present study. Based on their developmental period, 
Sisyphus species are categorized into two categories, species with a long 
developmental period (Sisyphus sordidus, S. mirabilis, S. fortuitus, and S. 
fasciculatus) and species with a shorter developmental period (S. seminulum, S. 
spinipes, S. infuscatus, S. calcaratus, S. muricatus, and S. longipes). 
The sex ratio of 1:1 observed in the present study was reported in other species 
of the genus (S. mirabilis, S. muricatus, and S.barbarossa)   also (Paschalidis, 1974). 
It is best explained by Fisher’s principle (Hamilton 1967) and it says that in most 
sexually reproducing species, the ratio tends to be 1:1, given the assumption of equal 
parental expenditure on offspring of both sexes. Male births are less common than 
females and the new born male then has better mating prospects than a newborn 
female and therefore can expect to have more offspring. Therefore parents genetically 
disposed to produce males tend to have more than average numbers of grandchildren 
born to them. The genes for male-producing tendencies spread, and male births 
become more common. As the 1:1 sex ratio is approached, the advantage associated 
with producing males dies away. The same reasoning holds if females are substituted 
for males throughout. Therefore 1:1 is the equilibrium ratio and is the evolutionarily 
stable strategy (ESS). 
  In the current study, the female longevity is long compared to males, the same 
trend has been observed in the previous studies of other Sisyphus species viz.,   
Sisyphus sordidus, S. mirabilis, S. fortuitus, S. fasciculatus, S. seminulum, S. spinipes, 
S. infuscatus, and S.calcaratus  (Paschalidis, 1974). Why females live longer than 
58 
 
males is generally unknown, it would be the metabolic differences or difference in 
patterns of resource allocation between males and females probably account for the 
gender difference in life span (Fox et al., 2003).  
  The type-2 pattern of nesting is the more typical pattern amongst rollers and 
usually some participation by both sexes (Halffter 1989) are involved.  Several 
patterns of nesting, brood ball shape and burial are reported in the Sisyphus species 
(Paschalidis, 1974). Our results show that in Sisyphus longipes, shorter developmental 
period and female longevity contribute to the abundance of the species and it also 
helps to accelerate the population growth of the species in the moist belts of south 
India. 
5.5.   Tibiodrepanus setosus (Wiedemann, 1823).  
The nesting biology of Tibiodrepanus setosus showed a Type 4 nesting 
pattern. T. setosus exploits comparatively drier dung for constructing brood masses 
with each brood mass containing one egg. Among the various life cycle stages of T. 
setosus pupal phase was the crucial phase adversely affecting the life cycle of T. 
setosus. Sex ratio biased towards females is seen in T. setosus. Also, as per LMC 
(Local mate competition), a female-biased sex ratio is favored, if mating competition 
takes place between male offsprings. Whereas an equal sex ratio is expected under 
random mating (Fisher et al., 1930). Hence the female-biased sex ratio noticed in 
T.setosus indicates that the females have relatively high fecundity, with the females 
laying more eggs which are usually spread out in dung pats through their lifespan and 
multivoltine species. Relatively small size and high fecundity of inferior competitors 
help such species to avoid exclusion from the local species pool, even at high levels of 
competition (Hanski and Cambefort, 1991a). The present analysis of the life cycle 
indicates that  T. setosus with traits that are common in an r-selective species 
59 
 
(MacArthur and Wilson 1967) such as high fecundity, small body size, low egg 
mortality, shorter larval duration, early onset of maturity, shorter developmental 
period, short generation time (multivoltine) together with female-biased sex ratio, 
contributed to the higher abundance of T. setosus in the region. 
 
  
    
 
  
60 
 

                                                 CONCLUSION 
 
The current study provides information on the life biology and nesting 
behaviour of the five prominent dung beetle species, Onthophagus cervus, O. 
fasciatus, Tiniocellus spinipes, Sisyphus longipes and Tibiodrepanus setosus in the 
agribelts of the Malabar Coast region.  
1.   Onthophagus cervus 
Shorter larval duration, early onset of maturity, shorter developmental period, 
female-biased sex ratio, and longer duration of females (favouring high egg 
production) contribute to the higher abundance of tunneler species, Onthophagus 
cervus, in the Malabar Coast region. Type 1 nesting behaviour with vertical and 
horizontal tunnels extending to the deep soil was recorded.  
2.   Onthophagus    fasciatus  
Shorter larval duration, longer duration of females and female-biased sex ratio 
contribute to their abundance. Type 1 nesting behaviour with the vertical and 
horizontal tunnels reaching deep soil layer was recorded.  
 3.   Tiniocellus spinipes. 
Shorter larval and pupal developmental period, longer duration of females, 
female-biased sex ratio and Type 1 nesting behaviour with shallow tunnels in the top 
soil layer that enabled easier and fast tunnelling process lead to their abundance in the 
region. 
4.    Sisyphus longipes.  
Short larval and pupal developmental periods and more extended female 
longevity contribute to the higher abundance of the roller species. Sisyphus longipes 
61 
 
rolled the dung ball away from the dung pat and Type 2 nesting pattern with dung ball 
burial in the top soil layer recorded.  
5.   Tibiodrepanus setosus. 
The relatively small size, female-biased sex ratio, shorter larval-pupal 
duration, and early maturity onset lead to the high abundance of Tibiodrepanus 
setosus. Type 4 nesting behaviour is observed with the whole development from egg 
to adults occurring within the dung pat itself recorded. Beetles used moderately wet 
dung for building brood mass construction and laid one egg per brood mass. 
 
62 
 

Ali, S. M. S.,Naeem, M., Baig, F., Shazad, A., & Zia, A.(2015). New records, 
distributional notes and species diversity of dung beetles (Coleoptera: 
Scarabaeidae: Scarabaeinae) from Pothohar Plateau of Punjab. Pakistan. 
Journal of Entomology and Zoology Studies, 3(3), 1–6. 
Andresen, E. (2005). Effects of season and vegetation type on community organization 
of dung beetles in a tropical dry forest 1. Biotropica: The Journal of 
Biology and Conservation, 37(2), 291–300. 
Andresen, E. (2008). Dung beetle assemblages in primary forest and disturbed habitats 
in a tropical dry forest landscape in western Mexico. Journal of Insect 
Conservation, 12(6), 639–650. 
Arellano, L., Castillo-Guevara, C., Huerta, C., Germán-García, A., & Lara, C. (2017). 
Nesting biology and life history of the dung beetle Onthophagus lecontei 
(Coleoptera: Scarabaeinae). Animal biology, 67(1), 41–52. 
Arrow, G. J. (1931). The Fauna of British India: Including Ceylon and Burma (Vol. 
13). Taylor & Francis. London. 
Balthasar, V. (1955). Scarabaeidae per Aphghanistan-Expendition (1952-53), 1. 
Klapperichs' Sbor. Entomol Odd. nar. Mus., Praha, 30, 409 – 439. 
Balthasar, V. (1963a). Monographie der Scarabaeidae und Aphodiidae der 
Palaearktischen und Orientalischen Region (Coleoptera: Lamellicornia). 
Verlagder TschechoslowakischenAkademie der Wissenschaften, Prague,1,1 
– 391. 
 Balthasar, V. (1963b).Monographic der Scarabaeidae und Aphodiidae der 
Palaearktischen und Orientalischen Region (Coleoptera: Lamellicornia) 
.Verlag der Tschechoslowakischen Akademie der Wissenschaften. Prague,1 
,1 – 627.  
 Balthasar, V. (1964).Monographie der Scarabaeidae und Aphodiidae der 
Palaearktischen und Orientalischen Region.(Coleoptera: Lamellicornia). 
63 
 
Verlag der Tschechoslowakischen Akademie der Wissenschaften Prague 
,III, 1– 652. 
Bang, H. S., Crespo, C. H., Na, Y. E., Han, M. S., & Lee, J. H. (2008). Reproductive 
development and seasonal activity of two Korean native Coprini species 
(Coleoptera: Scarabaeidae). Journal of Asia-Pacific Entomology, 11(4), 
195– 199. 
Baraud, J. (1992). Coleopteres Scarabaeoidea d'Europe, Faune deFrance 78. Federation 
Francaise des Societes de Sciences Naturelles, Paris, and Societe 
Linneenne de Lyon, Lyon, 856 pp . 
Barretto, J. W., Cultid-Medina, C. A., & Escobar, F.(2019). Annual abundance and 
population structure of two dung beetle species in a human-modified 
landscape. Insects,10(1), 2. 
Biswas, S., & Chatterjee, S. K. (1985). Insecta: Coleoptera: Scarabaeidae: 
Coprinae. Records of the Zoological Survey India, 82(1-4), 1471–77. 
Biswas, S., & Chatterjee, S. K. (1986). Scarabaeidae (Coleoptera) of Silent Valley, 
Kerala, India, with a description of three new species. Silent  Valley special 
issue. Records of the Zoological Survey of India, 82(1-4), 79 –96. 
Biswas, S., & Mulay, S. V. (2001). Insecta: Scarabid (Coleoptera), Fauna of Nilgiri 
Biosphere Reserve: Fauna of Conservation Area. Zoological Survey of 
India, 11, 129 –142. 
Blackwelder, R. E. (1944). Checklist of the coleopterous insects of México, Central 
America, The West Indies, and South America (No. 185). US Government 
Printing Office. 
Bornemissza, G. F., & Williams , C. H. (1970). An effect of dung beetle activity on 
plant yield. Pedobiologia, 10, 1– 7. 
Boucomont, A. (1914). Les coprophages de l'archipel malais. In Annales de la Société 
Entomologique de France . 83, pp. 238 – 350. 
64 
 
Branco, T.(2010) .Revision of the genera Tiniocellus Péringuey, 1901 and Nitiocellus 
gen. n.(Coleoptera, Scarabaeidae, Oniticellini). Boletín de la Sociedad 
Entomológica Aragonesa, 47, 71 – 126. 
Cambefort, Y. & Hanski, I. (1991). Dung beetle population biology. In: Hanski I. and 
Cambefort Y. (eds.). Dung beetle ecology, 36 –50.  
Cambefort, Y., & Lumaret, J. P. (1983). Nidification et larves des Oniticellini afro-
tropicaux [Col. Scarabaeidae]. Bulletin de la Société entomologique de 
France, 88 (7), 542– 569. 
Cambefort, Y., & Walter, P. (1991). Dung beetles in tropical forests in Africa.[pp. 198–
210.]. In. Dung beetle ecology (I. Hanski and Y. Cambefort, editors). 
Chandra, K. (2000). Inventory of scarabaeid beetles (Coleoptera) from Madhya 
Pradesh, India. Zoo’s Print Journal, 15(11), 359– 362. 
Chandra, K. (2005). Fauna of Western Himalaya (Part-2). Insecta: Coleoptera: 
Scarabaeidae. Zoological Survey of India, 141 –155. 
Chandra, K., & Ahirwar, S. C. (2007). Insecta: Coleoptera: Scarabaeidae. Zoological 
Survey of India, Fauna of Madhya Pradesh (including Chhattisgarh), State 
Fauna Series, 15(1), 273 –300. 
Chatterjee, S. K., & Biswas, S. (2000). Insecta: Coleoptera: Scarabaeidae. Zoological 
Survey of India, Fauna of Tripura, State Fauna Series, 7(3), 87– 98. 
 Chauhan, M., & Uniyal, V.P. (2020). A checklist of Dung beetles of Uttarakhand, 
Western Himalaya India. Indian Forester, 146 (11), 1059– 1064. 
Chown, S. L., Scholtz, C. H., Klok, C. J., Joubert, F. J., & Coles, K. S. (1995). 
Ecophysiology, range contraction and survival of a geographically 
restricted African dung beetle (Coleoptera: Scarabaeidae). Functional 
Ecology, 30– 39.  
Creutzer, C. (1799). Entomologische Versuche.(Kleine Beiträge zur nahen    
Berichtigungen eineger Käferferarten mit 3 ausgemahlten Kupfertafeln von 
Sturm J.).   Schaumburg und comp.Vien, 142pp.  
65 
 
Damborsky, M. P., Bohle, M. A., Polesel, M. I., Porcel, E. A., & Fontana, J. L. (2015). 
Spatial and temporal variation of dung beetle assemblages in a fragmented 
landscape at Eastern Humid Chaco. Neotropical Entomology, 44 (1), 30 –
39. 
Daniel, G. M., Sole, C. L., Davis, A. L., Strümpher, W. P., & Scholtz, C. H. (2020). 
Systematics of the dung beetle tribe Sisyphini Mulsant (Scarabaeidae: 
Scarabaeinae) inferred from molecular phylogeny and biogeography of 
southern African species. Systematic Entomology, 45(1), 73 –84. 
Davis, A. J. (2000). Species richness of dung-feeding beetles (Coleoptera: Aphodiidae, 
Scarabaeidae, Hybosoridae) in tropical rainforest at Danum Valley, Sabah, 
Malaysia. The Coleopterists Bulletin, 54(2), 221– 231. 
Davis, A. L. (1996). Seasonal dung beetle activity and dung dispersal in selected South 
African habitats: implications for pasture improvement in 
Australia. Agriculture, ecosystems  & environment, 58 (2-3), 157 –169. 
Davis, A. L. V. (2002). Dung beetle diversity in South Africa: influential factors, 
conservation status, data inadequacies and survey design. African 
Entomology, 10 (1), 53 –65. 
Delgado, L. (1997). Distribución estatal de la diversidad y nuevos registros de 
Scarabaeidae (Coleoptera) mexicanos. Folia Entomologica Mexicana, 99, 
37–56. 
Edwards, P. B., &  Aschenborn, H. H. (1987). Patterns of nesting and dung burial in 
Onitis dung beetles: implications for pasture productivity and fly 
control. Journal of Applied Ecology, 837– 851. 
Espinoza, V. R., & Noriega, J. A. (2018). Diversity of the dung beetles (Coleoptera: 
Scarabaeinae) in an altitudinal gradient in the east slope of Los Andes, 
Napo province, Ecuador. Neotropical Biodiversity, 4 (1), 145 –151. 
Estrada, A., & Coates-Estrada, R. (1991). Howler monkeys (Alouatta palliata), dung 
beetles (Scarabaeidae) and seed dispersal: ecological interactions in the 
66 
 
tropical rain forest of Los Tuxtlas, Mexico. Journal of Tropical 
Ecology, 7(4), 459 –474. 
Fabre, J.H. (1918). The Sacred Beetle and Others, Hodder and Stoughton, London. 
296pp. 
Fisher, R. A. (1930). The genetical theory of natural selection. Clarendon. London. 
272pp. 
Forgie, S. A. (2009). Reproductive activity of Onthophagus granulatus Boheman 
(Coleoptera: Scarabaeinae) in New Zealand: Implications for its 
effectiveness in the control of pastoral dung. New Zealand Entomologist, 
32, 76 –84. 
Fourcroy, A. F. (1785). Entomologia  parisiensis. Via et Aedibus Serpentineis, Paris. 
 
Fox, C. W.,Dublin, L., & Pollitt, S. J. (2003). Gender differences in lifespan and 
mortality rates in two seed beetle species. Functional Ecology, 17(5), 619– 
626. 
Gaikwad, A. R., & Bhawane,G. P. (2016). Observation on life cycle and nesting 
behavior of dung beetle Onthophagus catta (Fabricius). International 
Journal of Zoology Studies, 1(7), 09 –13. 
Gaikwad, A. R., & Bhawane,  G. P. (2015). Study of nidification behavior in three dung 
beetle species (Scarabaeidae: Scarabaeinae) from south-western 
Maharashtra, India. International Journal of Science and Research, 4, 1538 
–1542.  
Gajendra, N., & Prasad, S.K.(2016). A Review of Coleoptera diversity of Chhattisgarh: 
Updated checklist 2015. International Journal of Science and 
Research, 5(4), 710 –714. 
Gillard, P. (1967). Coprophagous beetles in pasture ecosystems. Journal of the 
Australian Institute of Agricultural Science, 33, 30–34. 
67 
 
Gillet, J. J. E. (1908). Coprides d’ Afrique tropicalc.Memoires de la Société 
entomologique de Belgique, 16, 63–81. 
Gillet, J. J. E. (1911). Coprophaga africana. Descriptions d’espèces nouvelles et 
remarques diverses.In Annales de la Société entomologique de Belgique ,. 
55, (11), 308 –312. 
Gillet, J. J. E. (1924). Scarabaeides coprophages de Sumatra. Annales de la Société 
entomologique  de Belgique, 64, 102–104. 
Gillet, J. J. E. (1935). Schwedisch-chinesische wissensch. Expedition nach den 
nordwestl. Provinzen Chinas. Scarabaeidae. Arkiv For Zoology, 27 (3),  1–
12. 
Gittings, T., Giller, P. S., & Stakelum, G. (1994). Dung decomposition in contrasting 
temperate pastures in relation to dung beetle and earthworm 
activity. Pedobiologia (Germany).  
Gómez-Cifuentes, A., Munevar, A., Gimenez, V., Gatti, M., & Zurita, G. (2017). 
Influence of land use on the taxonomic and functional diversity of dung 
beetles (Coleoptera: Scarabaeinae) in the southern Atlantic forest of 
Argentina.  Journal of Insect Conservation, 21(1),147 –156.  
González-Alvarado, A., & Vaz-de-Mello, F. Z. (2021). Towards a comprehensive 
taxonomic revision of the Neotropical dung beetle subgenus Deltochilum 
(Deltohyboma) Lane, 1946 (Coleoptera: Scarabaeidae: Scarabaeinae): 
Division into species-groups. Plos one,16 (1), e0244657. 
González-Maya, J. F., & Mata-Lorenzen, J. (2008). Dung-beetles (Coleoptera: 
Scarabeidae) from the Zona Protectora Las Tablas, Talamanca, Costa 
Rica. Check List, 4 (4), 458 – 463. 
González-Vainer, P. (2015). Feeding, reproductive, and nesting behavior of Canthon 
bispinus Germar (Coleoptera: Scarabaeidae: Scarabaeinae). The 
Coleopterists Bulletin, 69(1), 61 – 72. 
68 
 
Gonzalez-Vainer, P., & Morelli, E. (1999). Phenology and biology of the dung beetle 
Onthophagus hirculus Mannerheim (Coleoptera: Scarabaeidae). The 
Coleopterists' Bulletin, 53(4),303– 309. 
Halffter, G., & Edmonds, W. D. (1982). The nesting behavior of dung beetles 
(Scarabaeinae). An ecological and evolutive approach. The nesting 
behavior of dung beetles (Scarabaeinae). An ecological and evolutive 
approach.Journal of the  New York Entomological society, 91,512–515. 
   Halffter, G., & Matthews, E. G. (1966). The natural history of dung beetles of the 
subfamily Scarabaeinae (Coleoptera, Scarabaeidae). Folia Entomologica 
Mexicana, 12–14, 1–132. 
    Hamilton, W. D.(1967). Extraordinary sex ratios. Science, 156(3774), 477– 488. 
    Han, G. Y., Choi, J. B., & Park, J. K. (2021). A checklist of subfamily Scarabaeinae 
(Coleoptera: Scarabaeidae) from Cambodia. Journal of Asia-Pacific 
Biodiversity, 14 (1), 132– 139. 
  Hanski, I. (1991a). The dung insect community. In: Hanski I. and Cambefort Y. (eds.). 
Dung beetle ecology, 5–21. Princeton University Press, Princeton. 
  Hanski, I. (1991b). North Temperate Dung beetles. In: Hanski I. and Cambefort Y. 
(eds.). Dung beetle ecology, 75–96. Princeton University Press, Princeton. 
  Hanski, I., & Cambefort, Y. (1991a). Index of the genera in Scarabaeidae. Dung beetle 
ecology, 465-473. Princeton University Press. 
  Hernández,  M. I., Monteiro, L.R., & Favila, M. E. (2011).  The role of body size and 
shape in understanding competitive interactions within a community of 
Neotropical dung beetles. Journal of Insect Science,11 (1), 1 –13. 
HO, B. H. (2018). Two Dung Beetles of the Genus Onthophagus (Coleoptera: 
Scarabaeidae: Scarabaeinae) in Kinmen Islands: New Distribution 
Record., 3(2), 43 –45. 
Holter, P. (1975). Energy budget of a natural population of Aphodius rufipes larvae 
(Scarabaeidae). Oikos, 177 –186. 
69 
 
Hope, F. W. (1837). The Coleopterist's Manual, containing the lamellicorn insects of 
Linneus and Fabricius. The Coleopterist's Manual, containing the 
lamellicorn insects of Linneus and Fabricius,1837,Bohn,London.136pp. 
Horgan, F. G. (2001). Burial of bovine dung by coprophagous beetles (Coleoptera: 
Scarabaeidae) from horse and cow grazing sites in El Salvador. European 
Journal of Soil Biology, 37(2), 103– 111. 
House, C. M., Simmons, L. W., Kotiaho, J. S., Tomkins, J. L., & Hunt, J. (2011). Sex 
ratio bias in the dung beetle Onthophagus taurus: Adaptive allocation or 
sex-specific offspring mortality?. Evolutionary ecology, 25(2), 363 –372.  
Howden, H. F., & Gill, B. D.(1987).  New species and new records of Panamanian and 
Costa Rican Scarabaeinae (Coleoptera: Scarabaeidae). The Coleopterists' 
Bulletin, 41(3), 201 – 224. 
Howden, H. F., & Young, O. P. (1981). Panamanian Scarabaeinae: taxonomy, 
distribution,and habits (Coleoptera, Scarabaeidae) [New taxa]. 
 Contributions of the American Entomological Institute (USA),18 (1),1– 24. 
Huerta,  C., & Bang, H. S. (2004). Fecundity and offspring survival of Copris tripartitus 
Waterhouse (Coleoptera, Scarabaeidae: Scarabaeinae) under laboratory 
rearing conditions. The Coleopterists Bulletin, 58(4), 501 –507. 
Huerta, C., & García-Hernández, M. (2013) . Nesting Behavior of Onthophagus 
incensus Say, 1835 (Coleoptera: Scarabaeidae: Scarabaeinae). The 
Coleopterists Bulletin, 67(2), 161 –166.  
Huerta, C., Halffter, G., Halffter, V., & López, R. (2003). Comparative analysis of 
reproductive and nesting behavior in several species of Eurysternus Dalman 
(Coleoptera: Scarabaeinae: Eurysternini). Acta zoológica mexicana, 88, 01 
–41. 
Huerta, C., Martínez, I., & García- Hernández,  M. (2010). Preimaginal development of 
Onthophagus incensus Say, 1835 (Coleoptera: Scarabaeidae: 
Scarabaeinae). The Coleopterists Bulletin, 64 (4), 365 –371.  
70 
 
Hunter III, J. S., Fincher, G. T., & Lancaster Jr, J. L. (1991). Observations on the life 
history of Onthophagus medorensis. The Southwestern entomologist 
(USA),16(3),205– 213. 
Hunter III, J. S., Fincher, G. T., & Sheppard, D. C. (1996). Observations on the life 
history of Onthophagous depressus (Coleoptera: Scarabaeidae). Journal of 
entomological science, 31(1),  63– 71. 
Illiger, J. C. W. (1803). Verzeichniss der in Portugall einheimischen Käfer. Erste 
Lieferung. Magazin für Insektenkunde, 2(1802), 186 –258. 
Illiger, J.C.W. (1798). In J. Kugelann,Verzeichniss der Kafer Preussens, ausgearbeitet 
von Illiger. Halle. 
Janssens, A. (1949). Contribution a l’ etude des Coleopteres Lamellicornes. Table 
synoptique et essai de classification practique des Coleopteres 
Scarabaeidae. Bulletin de l'Institut Royal des Sciences Naturelles de 
Belgique Entomologie, 25, 1–30. 
Jokela,  J., Lively, C. M., Dybdahl, M. F., & Fox, J. A. (1997). Evidence for a cost of 
sex in the freshwater snail Potamopyrgus antipodarum. Ecology, 78(2), 
452-460. 
Joseph, K. J. (1994). Sexual dimorphism and intra-sex variations in the elephant dung 
beetle Heliocopris dominus (Coprinae: Scarabaeidae). Entomon-
Trivandrum-, 19, 165– 165.  
Joseph, K. J. (1998). Biology and breeding behaviour of the elephant dung beetle, 
Heliocopris dominus Bates (Coprinae: Scarabaeidae). Entomon-
Trivandrum-, 23(4), 325– 330. 
Joseph, K. J. (2003).  The life cycle, ecological role and biology of immature stages of 
Heliocopris dominus Bates (Coleoptera:Scarabaeidae: Coprinae). Entomon-
Trivandrum-, 28(3), 247– 252.  
Kaartinen, R., Hardwick, B., & Roslin, T.  (2013). Using citizen scientists to measure an 
ecosystem service nationwide. Ecology, 94(11), 2645– 2652. 
71 
 
Kakkar, N., & Gupta, S. K.(2010). Diversity and seasonal fluctuations in dung beetle 
(Coleoptera) community in Kurukshetra, India. Entomological 
Research, 40(3), 189 –192. 
Kalawate, A. S. (2018). A preliminary study on the dung beetles of the northern 
Western Ghats, Maharashtra, India. Journal of Threatened Taxa, 10(2), 
11316 –11331. 
Karimbumkara, S. N., & Priyadarsanan, D. R. (2016). Report of dung beetles 
(Scarabaeidae: Scarabaeinae) attracted to unconventional resources, with 
the description of three new species. Entomon, 41(4), 265 –282. 
Karimbumkara, S. N., & Priyadarsanan, D. R. (2013). Fauna of Karnataka, State Fauna 
Series. Zoological Survey of India, 21, 173 –178. 
 Khadakkar, S. S., Tiple, A. D., & Khurad, A. M. (2018). Description of life stages of 
dung beetle Scaptodera rhadamistus (Fabricius, 1775) (Coleoptera: 
Scarabaeidae: Scarabaeinae) with notes on nesting and biology. Journal of 
Threatened Taxa, 10(15), 12990 –12994. 
Khadakkar, S. S., Tiple, A. D., & Khurad, A.M. (2019). Scarab Beetles (Coleoptera: 
Scarabaeoidea: Scarabaeidae) of Vidarbha, India, with Notes on 
Distribution. Proceedings of the National Academy of Sciences, India 
Section B: Biological Sciences, 89 (4), 1239– 1249. 
Kharel, B. P., Schoolmeester, P.,  & Sarkar, S. K. (2020). A  first faunistic account on 
the Onthophagus Latreille, 1802 (Coleoptera, Scarabaeidae, Scarabaeinae) 
of the Nadia district, West Bengal, with a preliminary checklist from 
India. Check List, 16(2), 361– 381. 
Kirby, W. (1828). A description of some coleopterous insects in the collection of the 
Rev. F.W. Hope, F.L.S. Journal of Zoology, 3,520–525. 
Klemperer, H. G. (1983). Subsocial behaviour in Oniticellus cinctus (Coleoptera, 
Scarabaeidae): effect of the brood on parental care and 
oviposition. Physiological Entomology, 8(4), 393 –402. 
72 
 
Kotiaho, J. S., & Simmons, L.W. (2003). Longevity cost of reproduction for males but 
no longevity cost of mating or courtship for females in the male-dimorphic 
dung beetle Onthophagus binodis. Journal of Insect Physiology, 49 (9), 817 
–822. 
 Krajcik, M.(2006).Checklist of Scarabaeoidea of the world, 1. Scarabaeinae 
(Coleoptera: Scarabaeidae: Scarabaeinae). Anima. X, Supplement, 3, 1 –
190. 
Krikken, J. (2009). Drepanocerine dung beetles: a group overview, with a description 
of new taxa (Coleoptera: Scarabaeidae: Scarabaeinae). Haroldius Press. 
Lacordaire, M.T. (1856). Histoire Naturelle des Insectes-Genera des Coleopteres. 
Librairie Encyclopedique de Roret, Paris. Lamellicornien des 
palaearctischen Faunengebietes. Verhandlungen des. 
Latha, M. (2011). ‘Systematics and ecology of dung beetles (Coleoptera: Scarabaeidae: 
Scarabaeinae) in the Nelliampathi region of south Western Ghats’, Ph.D. 
Thesis, submitted to the University of Calicut. 
Latha, M., Cuccodoro,G., Sabu, T. K., and Vinod, K. V. (2011). Taxonomy of the dung 
beetle genus Ochicanthon Vaz-de-Mello (Coleoptera: Scarabaeidae: 
Scarabaeinae) of the Indian subcontinent, with notes on distribution 
patterns and flightlessness. Zootaxa, 2745 (1), 1 –29. 
Latreille, P.A. (1796). Precis de caracteres generiques des insectes, disposes dans un 
order naturel. Bordeaux. 
Latreille, P.A. (1802). Histoire naturelle generale et particuliere des crustacees et des 
insectes 3. Paris :F. Dufart,An X-XIII [1802-1805]. 
Latreille, P.A. (1807). Genera Crustaceorum et Insectorum secundum ordinem 
naturalem in familias disposita, iconibus exemplisque. Parisiis et 
Argentorate. Koenig,1806-09. 
 Lawrence, J. F. & Newton, A. F. (1995). Families and sub families of Coleoptera (with 
selective genera, notes, reference and data on family-group names). In: 
73 
 
Pakaluk A.F.J. and Lipiski S.A. (eds.).Biology, Phylogeny and 
Classification of Coleoptera,779–1006. Muzeum i Instytut Zoologii PAN, 
Warszawa, Poland.  
   Lee, C. M., & Wall, R. (2006). Cow-dung colonization and decomposition following 
insect exclusion. Bulletin of entomological research, 96 (3), 315 –322. 
Linnaeus, C. V. (1758). Systema Naturae per regna tria naturae. Secundum classes, 
ordines, genera, species, cum characteribus, differentiis, synonymis, 
locis. Editio, 1(10), 823.  
  Löbl, I. & Smetana, A.  (2006).  Catalogue of   Palaearctic Coleoptera. Apollo Books 
Stenstrup, 3, 1–690. 
Löbl, I. & Löbl, D. (Eds.). (2016). Scarabaeoidea–Scirtoidea–Dascilloidea–
Buprestoidea-Byrrhoidea: Revised and Updated Edition.  Brill. 
Lopera, A. (1996). Distribución y diversidad de escarabajos coprófagos (Scarabaeidae: 
Coleoptera) en tres relictos de bosque altoandino (Cordillera Oriental, 
Vertiente Occidental,Colombia). Trabajo de grado. Pontificia Universidad 
Javeriana. Bogotá. 
Lugon, A. P., Boutefeu, M., Bovy, E., Vaz‐de‐Mello, F. Z.,  Huynen, M. C., Galetti, 
M., & Culot, L. (2017). Persistence of the effect of frugivore identity on 
post‐dispersal seed fate: consequences for the assessment of functional 
redundancy. Biotropica, 49 (3), 293 –302.  
     Lutz, F. E. (1931). Insect vs. the people. Natural History, 31, 49–57. 
  Macagno, A., Moczek, L.A. P. & Pizzo, A.(2016). Rapid divergence of nesting depth 
and digging appendages among tunneling dung beetle populations and 
species, The American Naturalist, 187 (5), E143 – E151. 
Main, H.(1922).Notes on the metamorphosis of Onthophagus taurus L. In  Proceedings 
of the Entomological Society of   London (Vol. 1922, pp. 14 –16). 
Martínez, M.I., Montes de Oca, E., & Cruz, R. M. (1998).Contribución al conocimiento 
de la biología del escarabajo coprófago Onthophagus incensus Say 
74 
 
(Coleoptera: Scarabaeidae: Scarabaeinae): Datos ecológicos y 
reproductivos en relación a su fenología. Folia Entomológica 
Mexicana, 103, 1-13. 
Martínez-M, I., Lumaret, J. P., Diego, A. K. M., & Cano, B. M. (2019). The 
reproductive biology of Euoniticellus intermedius (Reiche)(Coleoptera: 
Scarabaeinae: Oniticellini). Proceedings of the Entomological Society of 
Washington, 121 (4), 642 –656. 
Mathew, G. (2004). Biodiversity Documentation for Kerala.Part.7 Insects. KFRI 
Handbook No.17. Kerala Forest Research Institute. 
Medina Hernández,  M. I., Niero, M. M., Schumacher, F., & Wuerges, M. (2020). 
Feeding and reproductive behavior of the dung beetle Canthon rutilans 
cyanescens (Coleoptera: Scarabaeinae). Revista Brasileira de 
Entomologia, 64 (2),1 –10. 
Menéndez, R.,Webb, P., & Orwin, K. H. (2016). Complementarity of dung beetle 
species with different functional behaviours influence dung–soil carbon 
cycling. Soil Biology and Biochemistry, 92, 142– 148. 
Miranda, C. H. B., Santos, J. D., & Bianchin, I. (1998). Contribuição de Onthophagus 
gazella à melhoria da fertilidade do solo pelo enterrio de massa fecal 
bovina fresca. Revista Braileira de. Zootecnia, 27(4), 681 –685. 
Mittal, I. C. (1993). Natural manuring and soil conditioning by dung beetles. Tropical 
Ecology, 34(2), 150 –159. 
Mittal, I. C. (2005). Diversity and conservation status of dung beetles (Laparosticti: 
Scarabaeidae: Coleoptera) in North India. Bulletin of the National Institute 
of Ecology, 15, 43 –51.  
Mittal, I.C. & Jain, R.(2015). A checklist of Indian dung beetles (Coleoptera: 
Scarabaeidae). Indian journal of Entomology, 77 (4), 383– 404. 
75 
 
Nakane, T. & Shirahata, K. (1957). A list of coprophagous Lamellicornia collected 
from Shansi, North China. Scientific reports of the Saikyo University, 2, 
229–232. 
 Nakane, T. & Tsukamoto K. (1956). On the Scarabaeidae of Japan III. (Sub family 
Scaraabeinae). Entomological Review of Japan,7(1), 23–27. 
  Nealis, V. G. (1977). Habitat associations and community analysis of south Texas dung 
beetles (Coleoptera: Scarabaeinae). Canadian Journal of Zoology, 55(1), 
138 –147. 
Nervo, B., Caprio, E., Celi, L.,Lonati, M., Lombardi, G., Falsone, G. & Rolando, A. 
(2017). Ecological functions provided by dung beetles are interlinked 
across space and time: evidence from 15N isotope tracing. Ecology, 98(2), 
433 –446. 
Nithya, S. (2012).‘Taxonomy and ecology of forest dung beetles (Coleoptera: 
Scarabaeidae: Scarabaeinae) in the moist south Western Ghats’, Ph.D. 
Thesis, University of Madras. 
Nithya, S., Thomas, S. K., & Flemming, A. T. (2015). Forest vegetation types related 
variation in the diversity and community structure of dung beetles 
(Coleoptera: Scarabaeidae: Scarabaeinae) in the moist South Western 
Ghats. Entomon, 40(4), 199 –208. 
Novais, S., Evangelista, L. A., Reis-Júnior, R., & Neves, F. S. (2016). How does dung 
beetle (Coleoptera: Scarabaeidae) diversity vary along a rainy season in a 
tropical dry forest. Journal of Insect Science, 16(1), 1 –6. 
Ocampo, F. C., & Philips, T. K. (2017). Food relocation and nesting behavior of the 
Argentinian dung beetle genus Eucranium and comparison with the 
southwest African Scarabaeus (Pachysoma) (Coleoptera: Scarabaeidae: 
Scarabaeinae). Revista de la Sociedad Entomológica Argentina, 64(1-
2),53– 59. 
76 
 
Pacheco, T. L., & Vaz-de-Mello, F. Z. (2019). New dung beetle genus and species from 
a cave in the Espinhaço mountain range, Brazil (Coleoptera: Scarabaeidae: 
Scarabaeinae). Journal of Natural History, 53(19-20), 1247 –1253. 
Palestrini, C., Barbero, E.,Trevisan, E., & Borghesio, L. (2002). Reproductive biology 
in three species of the genus Onitis Fabricius 1798 (Coleoptera 
Scarabaeidae Onitini). Tropical Zoology, 15(1), 97 –103. 
Paschalidis, K. M. (1974). The  genus Sisyphus latr. Coleoptera: scarabaeidae) in 
southern  Africa. Master’s degree Thesis. 
Patole, S. (2019). Studies on diversity and relative abundance of dung beetles 
(Coleoptera:  Scarabaeidae) from Sakri Tahsil, District, Dhulia (M.S) India. 
Indian Journal of Applied Research, 9 (10), 57–59. 
Paukku, S., & Kotiaho, J. S. (2005). Cost of reproduction in Callosobruchus maculatus: 
effects of mating on male longevity and the effect of male mating status on 
female longevity. Journal of Insect Physiology, 51(11), 1220– 1226. 
 Paulian, R. (1938). Expedition to southwest Arabia (1937-1338). Coleoptera: 
Scarabaeidae: Troginae, Geotrupinae, Dynamopinae, Hybosorinae, 
Coprinae, Aphodiinae. British Museum, I (12), 141–155. 
Pérez-Cogollo, L. C.,Rodríguez-Vivas, R. I., Delfín-González, H., Reyes-Novelo, E., & 
Morón, M. Á. (2015). Life history of Onthophagus landolti Harold, 1880 
(Coleoptera: Scarabaeidae), with descriptions of the preimaginal stages. The 
Coleopterists Bulletin, 69(2), 255– 263. 
Peringuey, L. (1900). Descriptive Catalogue of the Coleoptera of the South Africa 
(Lucanidae and Scarabaeidae). Transactions of the South African 
Philosophical Society, 12, 1–563. 
Philips, T. K.(2016). Phylogeny of the Oniticellini and Onthophagini dung beetles 
(Scarabaeidae,Scarabaeinae) from morphological evidence. ZooKeys, (579), 
9 –57. 
77 
 
Piccini, I., Arnieri, F., Caprio, E., Nervo, B., Pelissetti, S.,Palestrini, C. & Rolando, A. 
(2017). Greenhouse gas emissions from dung pats vary with dung beetle 
species and with assemblage composition. PLoS One, 12(7), e0178077. 
 Pulido-Herrera, L. A., & Zunino, M. (2007). Catálogo preliminar de los Onthophagini 
de América (Coleoptera: Scarabaeinae). Escarabajos, diversidad y 
conservación biológica. Ensayos en homenaje a Gonzalo Halffter, 7, 93 –
129. 
 Ratcliffe, B. C. (1991). The scarab beetles of Nebraska. Bulletin of the University of 
Nebraska  State Museum (USA),22,1 –570. 
 Reitter, E. (1892). Bestimmungs-tabelle der Lucaniden und coprophagen,    
Lamellicornen des palaearctischen Faunengebietes. Separate publication, 
Brunn :3-230. (1893).Catalogue of life check list. 
 Romero-Samper, J. E. S. U. S., & Martin-Piera, F. E. M. I. N. (1995). Nesting 
behaviour, ontogeny and life-cycle of Onthophagus stylocerus (Coleoptera: 
Scarabaeidae). European Journal of Entomology, 92 (4), 667– 679. 
Rougon, D., & Rougon, C. (2014). Dung beetles of the Sahel region. In Dung beetle 
ecology (pp. 230 –241). Princeton University Press. 
Sabu, T. K. (2011). Guild Structure, Taxonomic Diversity and Biosystematics of Dung 
Beetles (Coleoptera: Scarabaeinae) in the Agriculture and Forest Habitats 
of South Western Ghats. Project report submitted to University Grant 
Commission, India. 
Sabu,  T. K., Vinod, K.V., Latha, M., Nithya, S., & Boby, J. (2011). Cloud forest dung 
beetles (Coleoptera: Scarabaeinae) in the Western Ghats, a global 
biodiversity hotspot in southwestern India. Tropical Conservation 
Science, 4(1), 12– 24. 
Sabu, T.K. (2012). Dung specificity, guild structure, seasonality and species 
composition of dung beetles (Coleoptera: Scarabaeinae) associated with the 
dung droppings of major mammals (elephant, gaur, wild boar, deer, and 
78 
 
macaque) and composition of arboreal dung beetles in the wet and dry 
forests of the Western Ghats. Project report submitted to Ministry of 
Environment and Forests, India. 
Salomão, R. P., Gonçalves, L. K. S., de Morais, R. F., & Lima, L. R. C. (2019). Dung 
beetles (Coleoptera: Scarabaeinae) in a mosaic habitat at the ecotone 
between two savanna ecosystems in the Neotropical region. International 
Journal of Tropical Insect Science, 39 (3), 249 –256. 
Sánchez-Hernández, G., Gómez, B., Chamé-Vázquez, E. R., Dávila-Sánchez, R. A., 
Rodríguez-López, M. E., & Delgado, L. (2020). Current status of dung 
beetles (Coleoptera, Scarabaeidae, Scarabaeinae) diversity and conservation 
in Natural Protected Areas in Chiapas (Mexico). Neotropical Biology and 
Conservation, 15 (3), 219 –244.  
Sarkar,   S. K., & Kharel, B. P.  (2020). A first faunistic study on the tribe Oniticellini 
Kolbe, 1905 (Coleoptera: Scarabaeidae) of Baikunthapur Tropical Forest of 
the Himalayan foothills, West Bengal, India. Biodiversity Data Journal,8, 
e5744 
 Sarkar, S. K, Saha, S. & Raychaudhuri, D. (2015). On the taxonomy of Scar-abaeine 
fauna (Coleoptera: Scarabaeidae) of Buxa Tiger Reserve (BTR), West 
Bengal, India. Munis Entomology  & Zoology, 10 (1), 18– 48. 
Sarkar, S., K, Saha, S & Raychaudhuri, D.(2010). Further additions to the scarab 
beetles of  Buxa Tiger Reserve,  Jalpaiguri, West Bengal. Bionotes, 12 (4), 
131–132. 
Sathiandran, N.,Thomas, S. K., &  Flemming, A. T. (2015). An illustrated checklist of 
dung beetles (Coleoptera: Scarabaeinae) from the Periyar Tiger Reserve, 
Kerala, India. Journal of Threatened Taxa, 7(15), 8250 –8258. 
Sato, H. (1997). Two nesting behaviours and life history of a subsocial African 
dungrolling beetle, Scarabaeus catenatus (Coleoptera: 
Scarabaeidae). Journal of Natural History, 31(3), 457 –469. 
79 
 
  Scheffler, P. Y. (2002). Dung beetle (Coleoptera: Scarabaeidae) ecology in the intact 
and modified landscape of Eastern Amazonia. The Pennsylvania State 
University. 
  Scholtz, C.H., Davis, A. L.V., & Kryger, U. (2009). Evolutionary biology and 
conservation of dung beetles (pp.147-154).  Sofia-Moscow: Pensoft.  
 Schoolmeesters, P. (2019). World   Scarabaeidae Database in the Catalogue of Life. 
 Schoolmeesters, P., &  Sabu, S. K. (2006). A new Onthophagus species from Kerala, 
India. Phegea, 34(2), 73–75. 
Serville, J.G.A. (1825). In    Encyclopedie Methodique. Entomologie, 10, 1–344. 
Sewak, R. (2006). Insecta: Coleoptera: Scarabaeidae: Coprinae (Dung 
beetles). Zoological Survey of India, Fauna of Arunachal Pradesh, State 
Fauna Series, 13, 191 –224.  
 Shahabuddin, S.(2010). Diversity and community structure of dung beetles (Coleoptera: 
Scarabaeidae) across a habitat disturbance gradient in Lore Lindu National 
Park, Central Sulawesi. Biodiversitas Journal of Biological Diversity, 11 
(1), 29 –33. 
Shobhana, K.A. (2016).‘Taxonomy and Ecology of Dung Beetles 
(Coleoptera:Scarabaeidae: Scarabaeinae) in a Thorny Forest in the south 
Western Ghats’, Ph.D. Thesis, submitted to the University of Calicut. 
Siddiqui, H., Ahmed, A., &   Khatri, I. (2014). Distributional Notes and New Records 
for the Dung Beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of Pakistan, 
Pakistan Journal of Zoology, 46 (2), 295 –307. 
Silva, P. G. (2017).Dung beetles (Coleoptera, Scarabaeinae) from high-altitude 
grasslands in São Joaquim National Park, Santa Catarina, southern 
Brazil. Check List, 13(6), 817 –830. 
Simi, V.K., Sabu, K.T. & Albin, T.F. (2012). Diversity and community structure of 
dung beetles (Coleoptera: Scarabaeinae) associated with semi-urban 
80 
 
fragmented agricultural land in the Malabar Coast in southern India, 
Journal of Threatened Taxa, 4(7), 2685–2692. 
Simmons, L.W., & Ridsdill‐Smith, J. (2011). Reproductive competition and its impact 
on the evolution and ecology of dung beetles. Ecology and evolution of 
dung beetles, 1 –20. 
Singh, A. P., De, K., Mahajan, S., Mondal, R., & Uniyal, V. P. (2019). Observations on 
nesting activity, life cycle, and brood ball morphometry of the Bordered 
Dung Beetle Oniticellus cinctus (Fabricius, 1775) (Coleoptera: 
Scarabaeidae) under laboratory conditions. Journal of Threatened 
Taxa, 11(9), 14137 –14143. 
Slade, E. M., Mann, D. J., & Lewis, O. T.(2011). Biodiversity and ecosystem function of 
tropical forest dung beetles under contrasting logging regimes. Biological 
Conservation, 144 (1), 166 –174. 
 Smith, A. B. T. (2003). Checklist of the Scarabaeoidea of the Nearctic Realm. 
University of Nebraska,  Lincoln. 
Sowig, P. (1996). Brood care in the dung beetle Onthophagus vacca (Coleoptera: 
Scarabaeidae): the effect of soil moisture on-time budget, nest structure, 
and reproductive success. Ecography, 19 (3), 254 –258. 
Subha, B.J., & Sabu, K. T. (2017). Bioindicator Dung beetles of a shaded coffee 
plantation in the Nilgiri Biosphere  Reserve of the south Western Ghats” 
Environmental Information System (ENVIS) Newsletter,envis Zoological 
Survey of India, 23 (4),8 –10.  
Subha, M. (2018). ‘Taxonomy, guild structure and dung specificity of dung beetles 
(Coleoptera: Scarabaeidae: Scarabaeinae) in a coffee plantation belt in South 
Wayanad’, Ph. D. Thesis, submitted to the University of Calicut. 
Thomas, S. K., Nithya, S. & Vinod, K.V. (2011). Faunal survey, endemism and 
possible species loss of Scarabaeinae (Coleoptera: Scarabaeidae) in the 
81 
 
western slopes of the moist South Western Ghats, south India, Zootaxa, 
2830(1), 29–38. 
Thomson, C. G. (1863). Skandinaviens coleoptera, synoptiskt bearbetade. (Vol. 5). 
Tryckt uti  Lundbergska boktryckeriet. Lund,Tryckt uti Lundbergska 
boktryckereit,1859-68.vol 7. 
Vaz-de-Mello, F.Z. (2003). Ochicanthon, a new name for Phacosoma Boucomont 
(Coleoptera, Scarabaeidae), preoccupied with Phacosoma Jukes-Browne 
(Mollusca). Coleopterist's Bulletin, 57(1),25–26.  
Veenakumari, K., & Veeresh G.K. (1996a). Dung beetle (Coleoptera: Scarabaeidae: 
Scarabaeinae) fauna of Banglore, Karnataka. Journal of the Bombay 
Natural History Society, 94, 171–173. 
Veenakumari, K., & Veeresh, G. K. (1996b). Notes on the feeding and breeding 
behaviour of Gymnopleurus gemmatus Harold and Gymnopleurus miliaris 
(F.)(Coleoptera: Scarabaeidae). Journal-Bombay Natural History 
Society, 93, 13– 19.  
 Veenakumari , K., & Veeresh G.K. (1966c). Some aspects of the reproductive biology 
of Onthophagus gazella (F.) and Onthophagus rectecornutus Lansb. 
(Coleoptera: Scarabaeidae), Journal of the Bombay Natural History 
Society, 93, 250–256. 
Veenakumari, K., & Veeresh, G. K. (1997). Subsociality in Dung Beetles Copris 
repertus Walker and Copris indicus Gill.(Coleoptera: Scarabaeidae)(With 
one plate). Journal-Bombay Natural History Society, 94, 530 –535. 
Venugopal, K. S., Thomas,S. K., & Flemming, A.T. (2012). Diversity and community 
structure of dung beetles (Coleoptera: Scarabaeinae) associated with semi-
urban fragmented agricultural land in the Malabar Coast in southern 
India. Journal of Threatened taxa, 4(7),2685– 2692. 
82 
 
Veran, S., & Beissinger, S.R.  (2009). Demographic origins of skewed operational and 
adult sex ratios: perturbation analyses of two‐sex models. Ecology 
Letters, 12 (2), 129– 143. 
Vinod, K. V. (2009). ‘Studies on the Systematics and Distribution of Dung Beetles 
(Scarabaeinae: Coleoptera) in the Forests and Agricultural Fields of 
Wayanad’, Ph.D. Thesis, Forest Research Institute University, Dehradun. 
Wagner, P.M., Abagandura, G.O., Mamo, M., Weissling, T., Wingeyer, A., & 
Bradshaw, J. D. (2021). Abundance and diversity of dung beetles 
(Coleoptera: Scarabaeoidea) as affected by grazing management in the 
Nebraska sandhills ecosystem. Environmental Entomology, 50(1), 222 –
231. 
Weber, F.(1801). Observationes entomologicae, continentes novorum quae condidit 
generum characteres, et nuper detectarum specierum descriptiones. 
Bibliopolium Academici. Novi. Kiliae,impensis Bibliopolii Academici 
Novi,1801.https://doi.org/10.5962/bhl.title.8639. 
Wolinska, J., & Lively, C. M. (2008). The cost of males in Daphnia pulex. 
 Oikos, 117(11), 1637 –1646.  
Woodruff, R.E. (1973). The scarab beetles of Florida. Part I, the Laparosticti 
(subfamilies Scarabaeinae, Aphodiinae, H ybosorinae, Ochodaeinae, 
Geotrupinae, Acanthocerinae). Arthropods of Florida and Neighboring 
Land Areas, Florida Department of Agriculture and Consumer    Services, 
8. Gainesville,Florida. 
Ziani, S. (2021). Subsequent spelling ritchiei for Mnematium ritchii Macleay, 1821 
(Coleoptera: Scarabaeoidea: Scarabaeidae: Scarabaeini) to be maintained 
for prevailing usage. Insecta Mundi, 0856, 1 –5. 
Zunino, M. (1983). Essai pre´liminaire sur l’e´volution des armatures genitales des 
Scarabaeinae, par rapport a` la taxonomie du groupe et a` l’e´volution du 
83 
 
comportement de nidification (Col. Scarabaeidae). Bulletin de la Societe 
Entomologique de France,88, 531–542. 
Zunino, M.(1984). Analisi sistematica e zoogeografica della sottofamiglia 
Taurocerastinae Germain (Coleoptera, Scarabaeoidea: Geotrupidae). 
Bollettino del Museo Regionale di Scienze Naturali - Torino, 2, 445–464. 
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LIST OF FIGURES 
 
  Figure 1: A) Onthophagus cervus (Fabricius, 1798); B) Tiniocellus spinipes 
(Roth, 1851); C) Sisyphus longipes (Olivier, 1789); D) Onthophagus 
fasciatus Boucomont,1914; E) Tibiodrepanus setosus (Wiedemann, 
1823). 
  Figure 2: Laboratory cultural setup of dung beetles; A) Rearing of roller 
species; B) Tunneler species C) Dweller species; D) Earthern pot 
topped with fresh cow dung and covered with mesh net; E) 
Maintenance of cultural setup in a controlled room condition; F)  
Experimental setup sprayed with mist sprayer .  
 
Figure 3: Nest architecture study of dung beetles; A, B & C) Plastic pot cut in 
half lengthwise; D) Rejoined with masking tape; E) Rejoined plastic 
pot tied with mesh net. 
Figure 4: A) Brood ball of Onthophagus cervus; B) Egg glued to the wall of 
brood mass; C)  First instar larva; D) Second instar larva; E) Third 
instar larva;  F) Pupal cell G) Pupa early phase ; H) Pupa -late phase ;  
I ) Teneral adult; J) Emergence of adult from pupal cell; K) Adult 
male;L) Adult female;M&N) Nesting behaviour vertical and 
horizontal tunnels. 
Figure 5: A) Egg of Onthophagus fasciatus ; B) Brood ball ; C) First instar larva 
;D) Second Instar larva; E) Third instar larva ; F) Pupa; G) Pupal cell; 
H) Male;  I) Female. 
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Figure 6:  A) Egg of Tiniocellus spinipes ; B) First instar larva ;  C) Third instar 
larva;  E) Pupa ; F) Adult emerged cutting a hole in the pupal cell ; G) 
Female; H) Male.  
Figure  7: Brood ball of Sisyphus longipes; B) Egg ; C) First instar larva; D ) 
Second instar larva; E) Third instar larva; F) Pupa ;  G) Pupal cell ; H) 
Emerging adult; I) Male ; J) Female. 
Figure  8: A) Egg of Tibiodrepanus setosus; B) First instar larva; C) Second 
instar larva; D) Third instar larva;  E) Pupal cell; G) Teneral adult ; H) 
Male ;I) Female.  
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
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LIST OF TABLES 
Table 1: Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Onthophagus cervus in the 
agribelts of Malabar Coast region. 
Table 2: Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Onthophagus fasciatus in the 
agribelts of the Malabar Coast region. 
Table 3: Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Tiniocellus spinipes in the 
agribelts of   Malabar Coast region. 
Table 4: Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Sisyphus longipes in the agribelts 
of   Malabar Coast region. 
Table 5: Fecundity, egg mortality, egg hatchability, larval survivability, pupal 
survivability, and adult mortality of Tibiodrepanus setosus in the 
agribelts of Malabar Coast region. 
Table 6: Number and size of brood balls and duration (days) of different life 
cycle stages of various Onthophagus species (Author details are 
provided in parenthesis; ND: no data available). 
Table 7: Number and size of brood balls and duration (days) of different life 
cycle stages of   various   Onthophagus species. 
(Author details are provided in parenthesis; ND: no data available). 
 
87 
 
Table 8: Number  and  size  of  brood  balls  and  duration  (days)  of  different  
life cycle  stages  of  various sub-tribe Oniticellina species. 
 
Table 9: Number,shape, and size of brood balls and duration (days) of different 
life stages of Sisyphus species. 
 
 
 
 
 
 
 
88