Dung beetles: key to healthy pasture? An overvie · 2021. 1. 4. · extremely high (Hanski & Cambefort, 1991a,b). World Scientific News 153(2) (2021) 93-123 -95- Fig. 1. Dung beetle

Available online at www.worldscientificnews.com

( Received 14 December 2020; Accepted 03 January 2021; Date of Publication 04 January 2021 )

WSN 153(2) (2021) 93-123 EISSN 2392-2192

Dung beetles: key to healthy pasture? An overview

Sumana Saha1,a, Arghya Biswas1,b, Avirup Ghosh1,c

and Dinendra Raychaudhuri2,d

1Post Graduate Department of Zoology, Barasat Government College, 10, K.N.C. Road, Barasat, Kolkata – 7000124, India

2IRDM Faculty Centre, Department of Agricultural Biotechnology, Ramakrishna Mission Vivekananda University, Narendrapur, Kolkata – 700103, India

a,b,c,dE-mail address: [email protected], [email protected], [email protected], [email protected]

ABSTRACT

Dung beetles (Coleoptera: Scarabaeidae) do just what their name suggests: they use the manure,

or dung of other animals in some unique ways! Diversity of the coprine members is reflected through

the differences in morphology, resource relocation and foraging activity. They use one of the three broad

nesting strategies for laying eggs (Dwellers, Rollers, Tunnelers and Kleptocoprids) each with

implications for ecological function. These interesting insects fly around in search of manure deposits,

or pats from herbivores like cows and elephants. Through manipulating faeces during the feeding

process, dung beetles initiate a series of ecosystem functions ranging from secondary seed dispersal to

nutrient cycling and parasite suppression. The detritus-feeding beetles play a small but remarkable role

in our ecosystem. They feed on manure, use it to provide housing and food for their young, and improve

nutrient cycling and soil structure. Many of the functions provide valuable ecosystem services such as

biological pest control, soil fertilization. Members of the genus Onthophagus have been widely proposed

as an ideal group for biodiversity inventory and monitoring; they satisfy all of the criteria of an ideal

focal taxon, and they have already been used in ecological research and biodiversity survey and

conservation work in many regions of the world. The existence of distinct communities of coprophagous

beetles, which differ in their species composition, species-abundance relations and efficiency of dung

removal, plays a major role in the breakthrough in pasture and livestock management. They as well play

roles in pollination and trophic management. The dung beetles release strategy group concluded that

they also result in the reduction of greenhouse gas emitted from agricultural sector like many others;

http://www.worldscientificnews.com/

mailto:[email protected]



World Scientific News 153(2) (2021) 93-123

-94-

these insects too have medicinal value. Even after the presence of high endemicity, they are still

somewhat at back place in an adjunct of apt knowledge on them with very few studies done on them,

the need of the hour being to utilise this potential as bioresource in the interest of society and socio-

economic value. Present paper is an attempt to provide an overview on the knowledge generated and

what we could do so far on the dung beetle diversity in the Dairy Farm of Ramakrishna Mission Ashrama

Campus, Narendrapur, South 24 Parganas & buffaloshed Naihati, North 24- Parganas, West Bengal.

Keywords: Dung beetle, Dweller, Oniticellus cinctus (Fabricius), O. spinipes Roth, New record, West

Bengal, Coleoptera, Scarabaeidae

1. INTRODUCTION

Dung beetles do just what their name suggests: they use the manure, or dung, of other

animals in some unique ways! These interesting insects fly around in search of manure deposits,

or pats, from herbivores like cows and elephants. Dung beetles come in a variety of colours,

from dull and glossy black to metallic green and red. Ancient Egyptians thought very high of

the dung beetles, also known as the scarabs (derived from their family name, Scarabaeidae).

The very basic understanding of the links between ecological functions and biodiversity is

needed to assess and predict the true environmental consequences of human activities. Through

manipulating faeces during the feeding process, dung beetles initiate a series of ecosystem

functions ranging from secondary seed dispersal to nutrient cycling and parasite suppression

(Andresen, 2002). They feed on manure, use it to provide housing and food for their young, and

improve nutrient cycling, soil structure, and forage growth in the meantime. Dung beetles are

important enough in manure and nutrient recycling. Many of these ecological functions provide

valuable ecosystem services such as biological pest control and soil fertilization. Members of

the genus Onthophagus (Scarabaeidae) have been widely proposed as an ideal group for

biodiversity inventory and monitoring, they satisfy all of the criteria of an ideal focal taxon, and

they have already been used in ecological research and biodiversity survey and conservation

work in many regions of the world. The existence of distinct communities of coprophagous

beetles, which differ in their species composition, species-abundance relations, and efficiency

of dung removal, plays a major role in the breakthrough in pasture and livestock management.

They as well play roles in pollination and trophic management. The dung beetles release

strategy group concluded that they also result in the reduction of greenhouse gas emitted from

agricultural sector like many others; these insects too have medicinal value. Even after the

presence of high endemicity, they are still somewhat at back place in an adjunct of apt

knowledge on them with very few studies done on them, the need of the hour being to utilise

this potential as bio resource in the interest of society and socio-economic value.

2. GLOBAL DISTRIBUTION (FIG. 1)

Dung beetles are found in all continents except Antarctica and live in farmland, forest,

grassland, prairie and desert habitats. Their diversity in tropical forests and savannas is

extremely high (Hanski & Cambefort, 1991a,b).


-95-

Fig. 1. Dung beetle distribution globally.

Food

The beetles mostly feed on the micro-organism rich liquid component of mammalian

dung and use more fibrous material to brood their larvae (Halffter & Matthews, 1966; Halffter

& Edmonds, 1982).

Types

Based on their nesting strategies dung beetles are broadly classified into three functional

groups viz. rollers (telocoprid), tunnelers (paracoprid) and dwellers (endocoprid) (Figs. 2 & 3).

Three behavioral groups of the beetles are relevant to manure recycling. Probably the best-

known group are the (a) tumble bugs or rollers (e.g., the species Canthon pilularius). The

typical behavioural characteristic of this group is, a male female pair rolls a ball of dung (brood

ball) away from a manure pile in order to bury it. Dung beetles generally work in pairs. Another

group are the (b) tunnelers.

An example of this group is Onthophagus gazella, which typically bury the dung balls

under the manure pat or close to the edge. Piles of soil next to the dung pat are indicators of

tunneler-type dung beetle activity. Collectively, tunnelers and tumblers are classified as nesters

because of their behavior in preparing a home for their young. The third group of beetles that

use dung are the (c) dwellers.

Most dwellers belong to the subfamily Aphodiidae (Fig. 4). They live within the manure

pat, engage in little to no digging, and generally do not form brood balls.


-96-

Fig. 2. Dung beetle types.

Fig. 3. Diagrammatic view of nests of dung beetles


-97-

Fig. 4. Species-specific groups.

3. MORPHOLOGY

The subfamily Scarabaeinae is the large insect group exhibiting extreme diversity in size

(2 mm – 60 mm). Most of them are black, but a few more flamboyant species coming brown to

red and brilliant shades of green or gold. They are usually round with short winged covers

(Elytra) that expose the end of the abdomen. Just below the forehead their exoskeleton forms a

rounded shield-like clypeus, which covers the mouthparts. Some male members possess one or

two impressive horns, which they use as weapons to fend of other male competitors.

The front legs usually have serrated edges, used for power digging. Scarabs are

distinguished from other beetles by appearance of their antenna which are segmented and end

with plate like oval club of 3-7 expansible leaves.

These lamellate antennae create a large surface area for detecting odors. The dung chafers

form manure into a ball using its scooper like head and paddle shaped antennae (Fig. 5).


-98-

Fig. 5. How do dung beetles look?


-99-

Scarabaeine dung beetles have been repeatedly proposed as a useful group for

biodiversity inventory and monitoring.

They possess all of the characteristics of an ideal focal taxon, and have already been used

in ecological research and biodiversity survey work throughout the world (Halffter and Favila,

1993; Favila and Halffter 1997; Rodriguez et al., 1998; Spector and Forsyth, 1998; Davis et al.,

1999; Feer, 1999; Escobar, 2000; Davis, 2001; Jankielsohn et al., 2001; Lobo et al., 2001; Davis

et al., 2002; Roslin and Koivunen, 2001). Dung beetles can be more rapidly and quantitatively

sampled with standardized protocols than the vast majority of other insect groups. Primarily

using simple, inexpensive baited pitfall traps (Halffter and Favila, 1993; Lobo et al., 1988),

even non-specialist researchers can easily and quickly provide quantitative assessments of the

composition, structure, and abundances of dung beetle assemblages. Globally and locally, the

species diversity of the Scarabaeinae is not overwhelmingly large. A global catalog of the

subfamily, the ScarabNet Global Taxon Database, is now available online at

www.scarabnet.org, which lists just over 5,700 valid species of dung beetles grouped into 225

genera (ScarabNet, 2006). At local scales, dung beetle species richness ranges from just a

handful of species in the higher temperate latitudes to 75 species or more in the richest tropical

forests or savannas (Hanski, 1982; Price, 2004).

It is worth comparing this total to birds’ global diversity of roughly 9,700 species and the

richest tropical sites hosting several hundred species. Scarabaeines have a cosmopolitan

distribution, with multi-species communities occurring on every continent (other than

Antarctica), ensuring their broad geographical utility. Dung beetle communities display graded

responses to a variety of ecological factors or anthropogenic disturbances including pollution,

habitat modification, and habitat fragmentation (Howden and Nealis, 1975; Klein 1989; Hanski

and Niemela, 1990; Lobo et al. 1998; Barbero et al. 1999; Davis et al. 1999; Galante and

Cartagena, 1999; Halffter et al. 1995; Krüger and Scholtz, 1997; Davis and Sutton, 1998; Davis,

2000; Jankielsohn et al. 2001; Quintero and Roslin, 2005).

Rapid changes in dung beetle communities at sharp natural habitat ecotones have been

demonstrated in a number of settings (Hill, 1996; Spector and Ayzama, 2003). Shifts in the

diurnal activity, nesting behaviors, and average body size of species in communities have also

been documented due to habitat disturbance (Horgan, 2005).

Behaviour

Behavioural knowledge on these dung beetle presents plastic trait complexes such as

polyphenism, accounting for the bewilding diversity of forms and behaviour play a vastly

unexplored role in the evolution of morphological and behavioural novelties. Males of many

species of dung chafers of the genera Onthophagus and Catharsius express exuberant secondary

trait such as horns. These horns develop in response to larval feeding conditions and are used

as weapons in aggressive interactions and also to gain access and to mate with the female. In

many species, horn expression is discontinuous, and male populations are composed of two

alternative, discrete shapes.

Precisely many species exhibit polyphenism for male morphology. Both horned and

hornless males of Onthophagus taurus are indeed able to recognise the presence or absence of

other male siblings and respond by adjusting their investment into co-operative versus mate

securing behaviour (Moczek & Nijhout, 2003) (Fig. 6A). Such condition-dependent paternal

assistance may represent a mechanism by which males minimize fitness in a social environment

composed of variable degrees of male-male competition for females. Facultative parental


-100-

investment is likely to evolve when a patch and ephemeral resource environment favours co-

operation, while at the same time intraspecific mating competition selects for behaviours that

secure mates and breeding opportunities (Trumbo and Fernandez, 1995).

Many use their poop balls to cool off around noon when the sun is at its peak, they will

routinely climb atop their dung balls to give their feet a break from the hot ground (Fig. 6B).

Thermal imaging also showed that dung balls were measurably cooler than the surrounding

environment probably because of their moisture content. None of the members of scarabaeid

genera relocates food for nesting or prepares a brood ball or mass with provisions for the larva.

The absence of these two behaviours, present in practically all Scarabaeinae, makes one think

that the behaviour of these genera is very primitive. Other species also take advantage of a large

mass of dung gathered by a larger beetle. This behaviour does not require nesting given that the

kleptoparasite takes advantage of the mass of dung gathered by the larger beetle. This is

common in Aphodiini and has also been observed for other small Scarabaeinae. There is no

nidification process (i.e., accumulation or preparation of food by the mother for the larva).

There is no food relocation prior to ovipositing.

The females use food that is available, whether it is a pat of cow dung, the compact mass

of excrement buried by a sloth, the sausage or brood mass made by larger beetles, or even debris

in the nests of Acromyrmex. Neither brood masses nor brood balls are prepared (as occurs in

endocoprid species). The eggs are not deeply laid in the food mass, and there is no preparation

of an oviposition chamber. The larva moves freely through the mass of food, digging only a

few centimetres into it. During this movement, the larva does not prepare a gallery. This is, in

short, the simplest resource utilization process known for a species of the Scarabaeinae, and is

very similar to that observed for the majority of Aphodiini and other Scarabaeoidea that do not

nest. Although it is tempting to interpret the absence of nesting as a plesiomorphic character

that precedes the separation of the two main evolutionary lines of nesting in Scarabaeinae

(Halffter and Edmonds, 1982), this is not corroborated by the phylogenetic analyses available

at present (Vaz-de-Mello, 2003, 2007; Geńier and Kohlmann, 2003). The latter indicates that

the mentioned group is nested within a broader group that includes genera with different types

of nesting behavior: paracoprid and telecoprid of the most important apomorphies are the

presence of nesting behaviour and reduced ovary.

Being well supported as part of this clade and at the same time being the only branch (or

probably one of the few) with no nesting, in addition to having a sister group that do nest, is the

main evidence supporting the hypothesis of the loss of the nesting behaviour. Among the roller

Scarabaeinae there are several species in which nesting is missing in one or several of its

characteristic traits. These behaviours are the formation of a ball of food where the food is

found, and its relocation by rolling. Pachysoma MacLeay is a brachypterous genus that inhabits

the deserts of western South Africa.

The female of this species (nesting process is described by Scholtz et al., 2004) collects

dry pellets and small fragments of plant detritus and digs a gallery. She puts the material at the

bottom of the gallery and adds more that she searches for on the soil surface. The beetle

relocates the material horizontally by grasping the fragment of food with her front legs while

using the other four to walk, a relocation process similar to that of some Scarabaeinae, such as

the Eucraniini and Canthon obliquus Horn, from very arid zones. This behaviour is a clear

adaptation to how food is found under desert conditions. The other key behaviour that has been

lost is the preparation of a nest ball. South African ball-rolling dung beetles exhibit a unique

orientation behaviour to avoid competition for food: after forming a piece of dung into a ball,


-101-

they efficiently escape with it from the dung pile along a straight-line path. To keep track of

their heading, these insects use celestial cues, such as the sun, as an orientation reference. Here

the scientists show that wind can also be used as a guiding cue for the ball-rolling beetles. They

demonstrate that this mechanosensory compass cue is only used when skylight cues are difficult

to read, i.e., when the sun is close to the zenith.

This raises the question of how the beetles combine multimodal orientation input to obtain

a robust heading estimate. To study this, the scientists performed behavioural experiments in a

tightly controlled indoor arena. This revealed that the beetles register directional information

provided by the sun and the wind and can use them in a weighted manner. Moreover, the

directional information can be transferred between these 2 sensory modalities, suggesting that

they are combined in the spatial memory network in the beetle’s brain.

This flexible use of compass cue preferences relative to the prevailing visual and

mechanosensory scenery provides a simple, yet effective, mechanism for enabling precise

compass orientation at any time of the day. And with a keen spatial network, beetles are able to

get dung away from piles as quickly as possible – a resource so important, scientists compare

it to finding a bag of money on the street (Dacke et al. 2019).

Fig. 6. (A): Males of many species of dung chafers with exuberant secondary trait such as

horns, (B): Navigation of dung beetle using milky way as a guide to steer dung.


-102-

Life cycle and nesting behaviour of dung beetle

The life cycle of dung beetles is very interesting and unique with respect to most other

beetles. Most dung beetle species spend 95% of their life in dung or in the soil beneath dung

pats. The only time dung beetles may be observed outside this environment is when they are

searching for a new dung pat to start preparing and laying their eggs. Since each dung beetle

species exhibits different adaptations to their preferred microenvironment, they have somewhat

specific life cycle characteristics.

The life cycle and nesting behaviour of dung beetle is studied under laboratory conditions

(Gaikwad & Bhawane, 2016). The beetle construct simple nest composed of single unbranched

gallery (Fig. 7). At the bottom end of each there may be one or two brood balls lodged. The

depth of brood balls constructed ranges from 6 to 22 mm. The length of brood ball ranges from

22 to 34 mm with average of 27.7 mm.

The diameter of brood ball ranges between 13-18 mm. It is observed that single pair of

this dung beetle has constructed 10 to 35 brood balls in laboratory condition. The female dung

beetle deposits single egg in egg chamber of brood ball. The average size of the eggs is 2.36

mm in length and 1.39 mm in diameter. Embryological developmental period is averaged of

2.38 day. The larval developmental period of the three instar stage is averaged of 31.34 days.

While the average pupae stage is of 13.46 days at laboratory temperature condition ranging

from 22 °C to 25 °C. The adult longevity ranges between 42 to 85 days.

Fig. 7. Life cycle of dung beetle.


-103-

Beetle communities and dipteran flies colonising the beetle community

The faeces of large herbivores constitute a highly specific microhabitat, which is

characterized by discontinuity and a very high rate of micro-succession. Significant physical

and chemical changes occur within short periods of time (Dickinson & Craig, 1990). These

changes result from the activity of the various organisms that colonize animal dung, including

bacteria, protozoa, fungi, nematodes, arachnids, insects and earthworms, as well as local

weather conditions. The species composition of the invertebrates colonizing animal dung are

determined by several factors : physicochemical properties of the dung (moisture content and

size of particles), weather conditions, soil type, exposure to direct sun light, season, the size of

dung piles, interactions between species, the “age” and type of dung (Landin, 1961; Desiere,

1973; Olechowicz, 1974; Breymeyer & Zachareva-Stoilova, 1975; Holter, 2004). The faeces

of ruminants in the palearctic region have been studied extensively as part of ecological studies

on beetles (Koskela, 1972; Koskela & Hanski, 1977; Hanski & Koskela, 1979; Hanski &

Krikken, 1991;Wassmer, 1994; Šlachta et al. 2008), true flies (Hammer, 1941; Olechowicz,

1976), cattle dung (Wassmer, 1994; Šlachta et al. 2008), sheep dung (Olechowicz, 1974, 1976;

Sowig & Wassmer, 1994; Sowig, 1995), horse dung (Psarev, 2002) and others.

These studies encompass the effect of abiotic factors on the community structure of

organisms inhabiting dung at various stages in this micro-succession, their microhabitat

preferences and phenology. In cattle dung the eudominant species was: Cercyonpygmaeus

(Hydrophilidae, 10.4%), the dominants: Athetasordidula (Staphylinidae, 9.1%), Anotylus

tetracarinatus (Staphylinidae, 8.5%), Platystethus arenarius (Staphylinidae, 8.1%) and

Acrotrichis grandicollis (Ptiliidae, 6.6%).

Dung beetles: The phoront

Many mites use other animals to arrive at suitable habitats with enough food resources

that ensure high success of carried mites. The phenomenon in which one animal actively seeks

out and clings to the body surface of another animal in a limited time is called phoresy. During

this phenomenon, the carried animal (termed the phoretic or phoront) stops both feeding and

reproduction. Phoresy results in dispersal from unsuitable habitats to the suitable better

conditions for the development of the phoretic mites or their offsprings. In phoretic mite

species, deutonymphs or adult females are often the phoretic stages. Many groups of predatory

mites have also evolved a specialised habitat association with mammalian dung (Fig. 8).

Because of the temporally and spatially isolated nature of dung pads, these mites often disperse

by phoresy on coprophilous insects that have similar habitat requirements (Krantz, 1983).

Phoresy of this type requires behavioural and life history adaptations to ensure that the

dispersal stage of the mite is available and active at the time when the carrier insect is ready to

leave a local habitat patch.

These adaptations have become especially complex in the case of mites associated with

dung beetles of the subfamily Coprinae, which demonstrate a high level of parental brood care.

In these beetles, the female sometimes together with the male, remains in the underground nest

chamber caring for the brood until the emergence of the progeny. Many species of mites that

breed in these nest chambers disperse by phoresy on the beetles, as is common for other taxa of

dung beetles, but the existence of brood care in the Coprinae means that the mites have the

unusual opportunity of access to both the parental and progeny generations of beetles (Masan

& Halliday, 2009; Bahrami et al. 2011)


-104-

Fig. 8. Mites associated with dung beetle.

Seasonal occurrence (phenology) of coprophilus beetle

Dung is one of the fundamental returns of animal matter and energy into the nutrient and

energy cycles of the planet and supports a biodiverse community, especially arthropods that

contribute to decomposition (Bornemissza, 1960; Waterhouse, 1974; Fincher et al. 1981; Slade

et al. 2007; Nichols et al. 2008; O’Hea et al. 2010; Wu et al. 2010; Beynon et al. 2012;

Kudavidanage and. Lekamge, 2012). Coprophilous beetles are mainly comprised of larvae and

adult beetles from the families Scarabaeidae (subfamilies Scarabaeinae and Aphodiinae) and

Geotrupidae (Pakaluk et al. 1995; Ratcliffe and Jameson, 2005) and adult Hydrophilidae

(subfamily Sphaeridiinae) (Smetana, 1978) whose larvae are predacious. Coprophilous adult

and larval beetles compete with dung inhabiting dipteran larvae for food, whereas the larvae of

hydrophilid beetles are important predators on dipteran larvae. Both competition and predation

by coprophilous beetles reduce the fitness of many pest species that develop within dung

(Valiela, 1974; Fay and Doube, 1983; Kirk, 1992). Another benefit of dung beetles is the

enhancement of soil fertility (Brown et al. 2010; Ishikawa, 2011). Dung beetles increase the

rate of dung decomposition by tunnelling through dung pads and mixing dung with soil when

provisioning for their offspring (Fincher et al., 1981; Miranda et al., 1998; Nichols et al., 2008;

Brown et al. 2010; Liu et al. 2012). Previous research focused on dung beetle communities in

southern North America: Florida (Woodruff, 1973), Georgia (Fincher, 1975; Fincher and

Woodruff, 1979), Texas (Nealis, 1977; Fincher et al. 1986; Howden and Scholtz, 1986;

Howden and Howden, 2001), South Carolina (Harpootlian 2001), North Carolina (Bertone et

al. 2005). Louisiana (Radtke et al. 2008) and Arkansas (Fiene et al., 2011). Dung beetle studies

from northern North America provide distributional and phenological information include

studies in North Dakota (Helgesen and Post, 1967), New York (Valiela 1969; Pimsler 2007),

South Dakota (Kessler et al. 1974), Minnesota (Cervenka and Moon, 1991), southern Québec

(Matheson 1987; Levesque and Levesque 1995), southern Alberta (Floate and Gill, 1998;

Kadiri et al. 2014), New Jersey and Maryland (Price, 2004; Price et al. 2012) and Michigan


-105-

(Rounds and Floate, 2012). Studies that provide natural history data are essential to understand

beetle biodiversity (Braga et al. 2013; Korasaki et al. 2013), monitoring and advancement of

adventive species (Floate and Gill, 1998; Gollan et al. 2011; Kaufman and Wood, 2012; Rounds

and Floate, 2012; Floate and Kadiri et al., 2014), for providing essential ecological information

on local pasture ecosystems (Schulze et al., 2005; Qie et al. 2011; Agoglitta et al. 2012; Liu et

al. 2012; Numa et al. 2012; Campos and Hernandez, 2013), and as a proxy to monitor global

climate change (Menendez and Gutierrez, 2004; Wu and Sun, 2012; Wassmer, 2014).

Importance of dung beetle in nutrient cycle

There are many animals and plants that perform a vital role in the nutrient cycle of

pastures, particularly in humification of cattle dung. The organisms featuring prominently in

this role are insects mainly flies and beetles (Skidmore, 1991). This view is different from

Putman (1983) who asserted that earthworms are the primary invertebrates of cow dung

decomposition. The crucial role of the dung community is demonstrated in Australia where no

native cow dung loving fauna exists. It is estimated that the undegraded dung of five cattle

removes from production one acre of land per year and considerable expense is being incurred

in trying to establish a cow dung community (Skidmore, 1991). Australia, USA and the

ranching areas of South America are seeking to correct the unavailability of native dung fauna

through establishing sustainable cow dung communities mainly beetles. Pasture land covered

by undegraded cow dung is effectively non-productive since research has shown that natural

degradation cannot occur in the absence of insect decomposers since fossil dung pellets have

been frequently found in sedimentary rocks (Skidmore, 1991). Despite the pivotal role played

by beetles in the dung humification process, they are persistently exposed to lethal chemicals

as non-target insects. Pyrethroids pose the greatest danger to beetles given that they are widely

used to control common tropical pests such as ticks and tsetse flies (Regional Tsetse and

trypanosomiasis Control Program Report, 1997). Pyrethroids are known to kill beetles through

disruption of the central nervous system and are known to be more potent at the larval stage.

Nutrient quality of vertebrate dung for the dung beetles

At the basis of a trophic web, coprophagous animals like dung beetles (Scrabaeoidea)

utilise resources that may have advantages as well as drawbacks. Several studies have

characterised the nutrients, eg: C/N ratios and the organic matter content for specific type of

dung. Analysis of water content, C/N ratios, amino acid, neutral lipid fatty acid, free fatty acid

and sterol composition can give us the adequate information on their sustenance. Dung beetles

strongly depend on a resource that is scarce and patchy in occurrence. Also, some dung volatiles

act as nutritional cue for dung beetles. In general, animal droppings vary in nutrient amounts,

even within a species or feeding guild. Whereas higher nutrient concentrations are generally

beneficial, dung beetles may face trade offs that constrain a higher preference of nutrient rich

dung. The C/N ratio abundance is frequently used as an index for quality descriptions of organic

substrates including dung. The C/N ratio is increased over ten- fold from carnivore dung to

herbivore dung. Omnivore dung has intermediate levels. Water content plays an important role

as adult dung beetles mainly use the liquid phase and its nutrients/ particles to feed on, it affects

the occurrence of species and the handling of brood balls. Sterols and cholesterols play key

function as they serve as components of the cell membrane as regulators of developmental

genes and as precursors of different hormones (Frank, 2017).


-106-

Values of dung beetle

Due to the high sensitivity of dung beetles to habitat modification and changing dung

resources, many of these ecological processes have already been disrupted or may be affected

in the future. Prediction of the functional consequences of dung beetle decline demands

functional studies conducted with naturally assembled beetle communities, which broaden the

geographic scope of existing work, assess the spatio-temporal distribution of multiple functions,

and link these ecosystem processes more clearly to ecosystem services. Dung chafers with

different behaviour have different effect on ecosystem services. The maintenance of ecosystem

services by them depends on diverse assemblage of dung beetle species. They are important in

nutrient cycling. They facilitate decomposition and make the nutrients available to other

organism. Studies have found that many soil nutrients are increased when dung beetles are

present. They also help increase nitrogen mineralisation, the process by which nitrogen is

converted from an organic to an inorganic form making it available for use by plants and

subsequently the rest of the food web. Tunnels are specially important for bioturbation i.e.

mixing and redistribution of sediments (Fig. 9).

The process affects soil moisture and soil aeration and thus increases plant productivity.

They also help increase of plant biomass, nitrogen content and grain production. These scarabs

may also contribute to plant productivity by dispersal of seeds found in dung, which lead to

increased plant recruitment. Beetles are important pollinators of decay-scented flowers. Such

beetles can reduce the abundance of parasites and flies that breed in the dung. After laying their

eggs in the ground, the larva hatch and eat gastrointestinal parasites found in the dung, and their

tunnelling action aerates the soil. These processes interrupt the lifecycle of parasite. The dung

beetle destroys the parasites at no charge. The dung burial activity is remarkably disrupted by

land use changes from natural forest to open agricultural area. Their presence elevated about

53% of the total dung removed and reduced about 83% and 63% of fly population and richness,

respectively. Dung beetles are also used as an indicator group, because they reflect structural

differences between habitats caused by forest type or human habitat modification (Klein, 1989;

Nummelin and Hanski, 1989; Hill, 1996; Davis and Sutton, 1998; Davis et al. 2001). The

potential of dung beetles as indicators for disturbance has been reviewed by Halffter and Favila

(1993) and McGeoch et al. (2002). In addition, the body size of dung beetles may be an

important factor interacting with habitat type and may be related to the presence of large

mammals (Hanski and Cambefort, 1991a,b). Smaller dung beetle species are more abundant

and less habitat specific (Hanski, 1982; Hanski and Cambefort, 1991a,b).

On a pasture-management level, dung pat removal is beneficial for forage availability.

Most ruminants will not graze closely to their own species manure pats. Research has shown

that the forage is palatable, but avoided because of the dung pile. Consequently, cattle manure

deposits can make from 5% to 10% per acre per year unavailable. By completely and quickly

removing the manure, dung beetles can significantly enhance grazing efficiency. The tunnelling

behaviour of dung beetles increases the soils capacity to absorb and hold water, and their dung-

handling activities enhance soil nutrient cycling. An adequate population and mix of species

can remove a complete dung pile from the surface within 24 hours. As the adult dung beetles

use the liquid component for nourishment and the roughage for the brood balls, the dung pat

quickly disappears. If left on the surface, up to 80% of manure nitrogen is lost through

volatilization; by quickly incorporating manure into the soil, dung beetles make more of this

nitrogen available for plant use. The larvae use only 40−50% of the brood ball before pupating,

leaving behind the remainder of this nutrient-rich organic matter for soil microbes, fungi and


-107-

bacteria to use in creating humus. The effects of vegetative cover and soil type on the abundance

and distribution of scarabaeine dung beetle resulted in differentiation of the diurnal and

nocturnal species and also the rate of abundance in bushveld less than that of grassland. The

degree of habitat specificity varied widely between species. During overcast weather, both

bushveld and grassveld species became abundant in the transitional zone. It somehow stated

that light intensity is one of the characteristic by which beetles recognise bushveld and

grassveld. Species turnover was highest between primary forest, cropland and secondary forest.

The results of such studies concluded on the idea that regeneration areas and certain land use

system can mitigate the detrimental effects of the clearance of original vegetation (Simmons &

Ridsdill-Smith, 2011).

Fig. 9 Functional consequences of dung beetles.

Factors Affecting Dung Beetle and Their Adaptation

Dung beetles have been the subject of research on the effects of environmental

disturbance caused by human activity in tropical forests (Halffter and Favila, 1993). Thus, if

the structure and composition of a landscape change as a consequence of human disturbance,

those changes would be expected to be reflected in the structure and composition of the dung

beetle assemblages. Nevertheless, landscape configuration does influence the distribution,

abundance and persistence of beetle populations as a result of reduced isolation and this, in turn,

facilitates movement across the landscape matrix (Montes De Oca, 2001, Estrada and Coates-

Estrada, 2002). Most studies have demonstrated that dung beetles respond negatively to the

fragmentation and transformation of natural habitats (Howden and Nealis, 1975; Klein, 1987;

Davis et al. 2001, Medina et al. 2002). The results from the studies referred corroborate this to


-108-

some extent, as most of the forest species are relatively less able to tolerate the microclimatic

conditions typical of open areas (Klein, 1987; Halffter et al. 1992). However, the creation of

new environments such as cropland and pasture favours the presence of the few forest species

that can tolerate the modification of their habitat and also allows for colonization by non-forest

species that arrive from other altitudinal levels. In addition to the reduction in species richness

and abundance, differences in the community species composition between both types of forest

and the areas used for raising cattle and agriculture. In the anthropogenic habitats, the new

component of the dung beetle assemblage is a mixture of species with the capacity to use both

forest and modified areas. Like the findings of Halffter and Arellano (2002), the study has

important implications in terms of the conservation and management of mountain ecosystems,

as they suggest that a certain degree of landscape modification can actually increase the total

number of species at a regional level. Many of the species that inhabit modified areas,

particularly those used for raising cattle and growing crops, can be found at elevations of under

1000 m.s.l., as is the case of Onthophagus acuminatus, O. clypeatus group and Oxysternon

conspicillatum, species typical of lowland forest, and Ontherus trituberculatus, abundant

between 1000 and 1300 m.s.l. (Escobar, unpublished data).

Furthermore, the differences among habitats detected at the guild level are indicative of

the roller species’ sensitivity to changes in vegetation and soil use. On the other hand, the

dominance of large, diurnal species in anthropogenic habitats suggests that this group has a

high competitive ability to exploit available excrement. The great majority of small species are

associated with forest areas, indicating a reduced capacity to colonize new environments. The

environmental matrix and spatial arrangement of various elements that make up the landscape

are crucial for understanding the ways in which organisms respond to changes induced by

humans (Gascon et al. 1999). Many environments act not so much as absolute barriers to

species movement but as selective filters, imposing new trends in the diversity, structure and

composition of species assemblages.

Sustaining Beetle Population in the Ecosystem

Beetles play an important role in maintaining pastures in a productive state. There is need

for constant biological monitoring of beetle populations in pastures and create conditions

conducive for their self-sustenance. Indiscriminate use of broad-spectrum chemicals should be

minimized as this might have negative impacts on dung beetle populations. Sensitivity of beetle

families to chemicals differs hence there is need to identify predominant beetle families in a

given locality. This helps determining location specific pyrethroid concentration levels that

would have less detrimental to the survival of the beetles (Bath et al., 1994).

Retrospect

The section is limited to the taxonomic account of the group, primarily because of the

emphasis laid in the above paragraph. Scaraboidea is one of the largest superfamilies in the

order Coleoptera and is globally known by approximately 2,200 genera and about 31,000

species (Ratcliffe and Jameson, 2013). This superfamily includes three families viz. Passalidae,

Lucanidae and Scarabaeidae. The last named family includes about 91% of all scarabaeoids and

includes about 600 genera and 27,800 species worldwide (Ratcliffe and Jameson, 2013). Of

these 7000 are known as dung beetles. Though a well documented taxonomic work on the

oriental dung beetle fauna had been carried out by Arrow (1931) and Balthasar (1963a & b),


-109-

the important contributions on the oriental scarabaeids have been made by Newton & Malcolm,

1985; Biswas & Chatterjee, 1991; Veenakumari & Veeresh, 1997; Chandra, 2000, 2008, 2009;

Chandra & Ahirwar, 2005, 2007; Priyadarsanan, 2006; Chandra & Gupta, 2012, 2013; Sarkar,

2013; Jadhav & Sharma, 2012; Ali et al., 2015; Sarkar et al., 2015; Sathiandran et al., 2015;

Kalawate, 2018 and Singh et al., 2018).

4. RESULT

Authors being unaware of any study conducted in the dairy farms of West Bengal,

initiated a study to unfold the faunal spectrum of dung beetles in the Dairy Farm of Ramakrishna

Mission Ashrama Campus, Narendrapur, South 24 Parganas & buffaloshed Naihati, North 24-

Parganas, West Bengal. During our survey we have sampled two (2) dung beetles alongwith

four (4) dung loving beetles, eight (8) dung loving flies, 17 dung associated insects of different

groups, one mesostigmatic mite species from cow and buffalo dung community. Out of these,

two coprophagous species, Oniticellus cinctus (Fabricius) and O. spinipes Roth under

subfamily Coprinae are recorded so far from cow dung from dairy farm, Narendrapur, South

24 Parganas and buffalo dung from cattle shed, Naihati, North 24 Parganas respectively.

Oniticellus spinipes is newly recorded from the state and O. cinctus first time reported from the

district (Table 1; Fig. 10).

Table 1. Arthropod Species Encountered In Different Dung Communities.

Sl.

No. Name of Species

Insects Encountered During

Survey State of Dung

In Cow

Dung

In Buffalo

Dung Wet/Dry

A.

1. Dung Beetles Oniticellus cinctus (Fabricius)

√

√

W

2. Oniticellus spinipes Roth x √ D

B.

3. Dung Loving Insects

Sphaeridium scarabaeoides (L.)

√

√

W + D

4. Sphaeridium sp. 2 √ √

W + D

5. Sphaeridium sp. 3 √ √ W + D

6. Clown Beetle [Hister unicolor L.] √ √ W + D

7. Fruit Fly : Drosophilidae : Indet sp. 1 x √ D

8. Dung Loving Fly : Lonchaeidae : Indet

sp. 2 √ √ W + D

9. Dipteran Fly : Opomyzidae Indet sp. 3 √ √ W + D


-110-

10. Dipteran Fly : Opomyzidae Indet sp. 4 x √ D

11. Dipteran Scavenger Fly : Sepsidae : Indet

sp. 5 √ √ W + D

12. Lesser Dung Fly : Sphaeroceridae : Indet

sp. 6 √ √ W + D

13. Lesser Dung Fly : Sphaeroceridae : Indet

sp. 7 x √ D

14. Dipteran Fly : Tachinidae : Indet sp. 8 x √ D

C.

15. Insects Associated with Dung

Ground Beetle [Nebria sp.]

√

√

W + D

16. Ground Beetle [Dromius sp.] √ √ W + D

17. Darkling Beetle [Gonocephalum

tuberculatum Hope] x √ D

18. Scarab Beetle [Aphodius prodromus

(Brahm)] √ x W

19. Staphylinid Beetle [Paederus riparius

(Linnaeus)] √ √ W + D

20. Ant [Camponotus (Tanymyrmex)

compressus (F.)] √ √ W + D

21. Ant [Crematogaster (Acrocoelia)

hodgsoni Forel] √ √ W + D

22. Ant [Diacamma scalpratum (Smith)] √ √ W + D

23. Ant [Myrmicaria brunnea Saunders] √ √ W + D

24. Ant [Pheidole (Pheidole) nietneri Emery] √ √ W + D

25. Ant [Vollenhovia oblonga (Smith)] √ x W

26. Stink Bug [Storthecoris nigriceps

Horvath] x √ D

27. Springtail (Indet sp. 1) x √ D

28. Earwig [Forficula sp. 1] √ √ W + D

29. Earwig [Forficula sp. 2] √ √ W + D

30. Spider : Salticidae (Indet sp. 1) x √ D

31. Spider : Lycosidae [Lycosa carmichaeli

Gravely] √ √ W + D

D.

32.

Mites Associated With Dung Beetles &

Dung Loving Beetles

Glyptholaspis sp.

√

x

W + D


-111-

Fig. 10. Dung beetles encountered during survey.

5. CONCLUSION

Through manipulating faeces during the feeding process, dung beetles instigate a series

of valuable ecosystem services. Even after the presence of high endemicity, they are still

somewhat at back place in an adjunct of apt knowledge on them with very few studies done on

them, the need of the hour being to utilise this potential as bioresource in the interest of society

and socio-economic value.

ACKNOWLEDGEMENTS

The authors express their deep sense of gratitude to The Hon’ble Vice Chancellor, Ramakrishna Mission

Vivekananda Educational and Research Institute (RKMVERI), Narendrapur and The Principal, Barasat

Government College for necessary logistic support. The authors also wish to thank The Secretary & The Dean,

RKMVERI, Narendrapur for necessary permission to carry out the project within the campus.


-112-

References

[1] Agoglitta, R., Moreno, C. E., Zunino, M., Bonsignori, G., and Dellacasa. M. 2012.

Cumulative annual dung beetle diversity in Mediterranean seasonal environments.

Ecological Research, 27(2): 387–395

[2] Ali S.M.S., Naeem, M. Baig, F, Shahzad, 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): 01-06

[3] Andresen, E. 2002a. Effect of forest fragmentation on dung beetle communities and

functional consequences for plant regeneration. Ecography, 26: 87–97

[4] Andresen, E. 2002b.Dung beetles in a Central Amazonian rainforest and their ecological

role as secondary seed dispersers. Ecological Entomology 27: 257–270

[5] Arrow, G. J. 1931. The Fauna of British India including Ceylon and Burma: Coleoptera:

Lamellicornia: Part III Coprinae. Publ. Taylor and Francis, London, 428pp.

[6] Bahrami, F., Arbabi, M. Vafaei Shoushtari, R. and Kazemi, S. 2011. Mesostigmatic

mites associated with Coleoptera and biodiversity calculation of these mites phoretic on

dung beetles in Golestan Province (North of Iran). Middle-East Journal of Scientific

Research, 9 (3): 345-366

[7] Balthasar, V. 1963a. Monographie der Scarabaeidae und Aphodiidae der

Palaearktischen und Orientalischen Region. (Coleoptera: Lamellicornia), Verlag der

Tschechoslowakischen Akademie der Wissenschaften Prag, I: 1-391, 137 figs., 24 pls.

[8] Balthasar, V. (1963b). Monographie der Scarabaeidae und Aphodiidae der

Palaearktischen und Orientalischen Region. (Coleoptera: Lamellicornia), Verlag der

Tschechoslowakischen Akademie der Wissenschaften Prag, I1: 1-627, 226 figs., 16 pls.

[9] Barbero, E., Palestrini, C. and Rolando, A. 1999. Dung beetle conservation: effects of

habitat and resources election (Coleoptera: Scarabaeidae). Journal of Insect

Conservation, 3, 75–84

[10] Bath, D. K. M., Heinzer-Mutz, E. M. and Elster 1994. Colonization and degradation of

cattle dung: Aspects of sampling, faecal composition, and artificially formed parts,

Environmental Entomology, 23: 571-578

[11] .Bertone, M., Green, J., Washburn, S., Poore, M., Sorenson, C. and Watson, W. A.,

2005. Seasonal Activity and Species Composition of Dung Beetles (Coleoptera:

Scarabaeidae and Geotrupidae) Inhabiting Cattle Pastures in North Carolina. Ecology

and Population Biology, 98 (3), 309-321

[12] Beynon, S.A., Mann, D.J., Slade, E.M. & Lewis, O.T. 2012. Species-rich dung beetle

communities buffer ecosystem services in perturbed agro-ecosystems. Journals of

Applied Ecology, 49: 1365-1372

[13] Biswas. S. and Chatterjee, S.K. 1991.Fauna of Orissa. State Fauna Series 1. Zoological

Survey of India, Kolkata, 3: 243-262


-113-

[14] Bornemissza, G.F. 1960. Could dung eating insects improve our pasture? The Journal

of the Australian Institute of Agricultural Science, 26: 54-56

[15] Braga, R. F., Korasaki, V., Andresen, E. and Louzada, J. 2013. Dung beetle community

and functions along a habitat-disturbance gradient in the Amazon: A rapid assessment

of ecological functions associated to biodiversity. Plos One, 8(2): e57786

[16] .Breymayer, A. and Zachaireva-Stoilova, B. 1975. Scarabaeidae in two mountain

pastures in Poland and Bulgaria. Bulletin of the Polish Academy of Science (Sci. Biol.

II), 33: 173–180

[17] Brown, J., Scholtz, C. H., Janeau, J. L., Grellier, S. and Podwojewski, P. 2010. Dung

beetles (Coleoptera: Scarabaeidae) can improve soil hydrological properties. Applied

Soil Ecology, 46(1): 9–16

[18] Campos, R. C., and Hernandez, M. I. M. 2013. Dung beetle assemblages (Coleoptera,

Scarabaeinae) in Atlantic forest fragments in southern Brazil. Revista Brasileira De

Entomologia, 57(1): 47–54

[19] Cervenka, V. J. and Moon, R. D. 1991. Arthropods associated with fresh cattle dung

pats in Minnesota. Journal of the Kansas Entomological Society, 64(2): 131–145

[20] Chandra, K. 2000. Inventory of scarabaeid beetles (Coleoptera) from Madhya Pradesh,

India. Zoos’ Print Journal, 15(11): 359-362

[21] Chandra, K. 2008. Insecta: Coleoptera. In: K. Chandra (Ed.) Faunal Diversity of

Jabalpur District, Madhya Pradesh. Publ. Zoological Survey of India, Kolkata: 159-186

[22] Chandra, K 2009. Insecta: Coleoptera. In: K. Chandra (Ed.) Fauna of Pachmarhi

Biosphere Reserve. Conservation Area Series, 39: 259-299, Publ. Zoological Survey of

India, Kolkata.

[23] Chandra, K. and Ahirwar, S.C. 2005. Scarabaeid Beetles of Bandhavgarh National Park,

Madhya Pradesh. Zoos’ Print Journal, 20(80): 1961-1964

[24] Chandra, K. and Ahirwar, S.C. 2007. Insecta: Coleoptera: Scarabaeidae, Fauna of

Madhya Pradesh (including Chattisgarh). State Fauna Series, 15(Part-1): 273-300

[25] Jerzy Borowski, Adam Byk, Tomasz Mokrzycki, Beetles (Coleoptera) of the Rogów

region. Part IX – superfamily Scarabaeoidea: Bolboceratidae, Geotrupidae, Lucanidae,

Trogidae and Scarabaeidae. World Scientific News 44 (2016) 123-142

[26] Subhankar Kumar Sarkar, Sumana Saha, Dinendra Raychaudhuri, On the Mimela

Kirby, 1823 (Rutelinae: Scarabaeidae) of Buxa Tiger Reserve (a forest under

biodiversity hot spot zone), Dooars, West Bengal, India. World Scientific News 50

(2016) 95-105

[27] Subhankar Kumar Sarkar, Sumana Saha, Dinendra Raychaudhuri, Anomala Samouelle,

1819 (Rutelinae: Scarabaeidae) of Buxa Tiger Reserve, Dooars, West Bengal, India.

Part – I. World Scientific News 65 (2017) 94-122

[28] Subhankar Kumar Sarkar, Sumana Saha, Dinendra Raychaudhuri, Anomala Samouelle,

1819 (Rutelinae: Scarabaeidae) of Buxa Tiger Reserve, Dooars, West Bengal, India.

Part – II. World Scientific News 118 (2019) 17-32


-114-

[29] Chandra, K. & Gupta, D. 2012. Diversity and composition of Dung beetles

(Scarabaeidae: Scarabaeinae and Aphodiinae) assemblages in Singhori Wildlife

Sanctuary, Raisen, Madhya Pradesh (India). Munis Entomology and Zoology, 7(2): 812-

827

[30] Chandra, K. and Gupta, D. 2013. Scarab beetles (Coleoptera: Scarabaeoidea) of

Barnawapara Wildlife Sanctuary, Chattisgarh, India. Journal of Threatened Taxa, 5(12):

4660-4671

[31] Dacke, A., Bell, T.A. Foster, J. J. Baird, E.J., Strube-Bloss, M.F., Byrne, M.J. and

Jundi, B.el. 2019. Multimodal cue integration in the dung beetle compass. Proc Natl

Acad Sci U S A, 116(28): 14248-14253. doi: 10.1073/pnas.1904308116

[32] Davis, A. J. 2000. Does reduced-impact logging help preserve biodiversity in tropical

rainforests? A case study from Borneo using dung beetles (Coleoptera: Scarabaeoidea)

as indicators. Environmental Entomology, 29: 467–475

[33] Davis, A. J., and Sutton, S.L. 1998. A dung beetle that feeds on fig: Implications for the

measurement of species rarity. Journal of Tropical Ecology, 13: 759–766

[34] Davis, A. J., Holloway, J.D., Huijbregts, H., Krikken, J., Kirk-Spriggs, A.H. and Sutton,

S.L. 2001.Dung beetles as indicators of change in the forests of northern Borneo.

Journal of Applied Ecology, 38: 593–616

[35] Davis, A. L. V., Scholtz, C. H. and Chown, S. L. 1999. Species turnover, community

boundaries and biogeographical composition of dung beetle assemblages across an

altitudinal gradient in South Africa. Journal of Biogeography, 26: 1039-1055

[36] Davis, A. L. V., C. H. Scholtz, and Philips, T. K. 2002. Historical biogeography of

scarabaeine dung beetles. Journal of Biogeography, 29: 1217–1256

[37] Desiere, M. 1973. Ecologie des Coléoptères coprophages. Annales de la Societe Royale

Zoologique de Belgiue, 103: 135–145

[38] Dickinson, C.H. and Craig, G. 1990. Effects of water on the composition and release of

nutrients from cow pats. New Phytologist, 115: 139-147.

[39] Escobar, F. 2000. Diversidad de coleopteroscoprofagos (Scarabaeidae: Scarabaeinae) en

unmosaico de habitats en la Reserva Natural Nukak, Guiviare, Colombia. Acta

Zoologica Mexicana, 79: 103–121

[40] Estrada A. and Coates-Estrada R. 2002. Dung beetles in continuous forest, forest

fragments and in agricultural mosaic habitat island at Los Tuxtlas, Mexico. Biodiversity

and Conservation 11: 1903–1918

[41] Favila, M., and Halffter, G. 1997. Indicator groups for measuring biodiversity. Acta

Zoologica Mexicana, 72: 1–25

[42] Fay, H. A. C. and Doube, B.M. 1983. The effect of some coprophagous and predatory

beetles on the survival of immature stages of the African buffalo fly, Hematobia

thirouxi potans, in bovine dung. Zeitschrift Fur Angewandte Entomologie, Journal of

Applied Entomology, 95(5): 460–466


-115-

[43] Feer, F. 1999. Effects of dung beetles (Scarabaeidae) on seeds dispersed by howler

monkeys (Alouatta seniculus) in the French Guianan rain forest. Journal of Tropical

Ecology, 15: 129–142

[44] Fincher, G. T. 1975. Dung beetles of Blackbeard Island (Coleoptera: Scarabaeidae). The

Coleopterists Bulletin, 29(4): 319–320

[45] Fincher, G.T. 1981. The potential value of dung beetles in pasture ecosystems. Journal

of the Georgia Entomological Society, 16(1): 316−333

[46] Fincher, G. T., Monson, W. G. and Burton, G. W. 1981. Effects of cattle feces rapidly

buried by dung beetles on yield and quality of coastal Bermuda grass. Agronomy

Journal, 73(5): 775–779

[47] Fincher, G. T., and Woodruff, R. E. 1979. Dung beetles of Cumberland Island, Georgia

(Coleoptera: Scarabaeidae). The Coleopterists Bulletin, 33(1): 69–70

[48] Fincher, G. T., Blume, R. R., Hunter, J. S. and Beerwinkle, K. R. 1986. Seasonal

distribution and diel flight activity of dung-feeding scarabs (Coleoptera, Scarabaeidae)

in open and wooded pasture in east-central Texas. Southwestern Entomologist

Supplement, 10: 1–35

[49] Fiene, J. G., Connior, M. B., Androw, R., Baldwin, B. and McKay, T. 2011. Surveys of

Arkansas dung beetles (Coleoptera: Scarabaeidae and Geotrupidae): Phenologies, mass

occurrences, state and distributional records. The American Midland Naturalist, 165(2):

319–337.

[50] Floate, K. D., and Gill, B. D. 1998. Seasonal activity of dung beetles (Coleoptera:

Scarabaeidae) associated with cattle dung in southern Alberta and their geographic

distribution in Canada. Canadian Entomologist, 130(2): 131–151

[51] Floate, K. D. and Kadiri, N. 2013. Dung beetles (Coleoptera: Scarabaeidae) associated

with cattle dung on native grasslands of southern Alberta, Canada. The Canadian

Entomologist, 145(06): 647–654

[52] Frank, K., Bruckner, A., Hilpert, A., Heethoff, M. and Bluthgen, N. 2017. Nutrient

quality of vertebrate dung as a diet for dung beetles. Scientific Reports, 7(12141): 1-12

[53] Galante, E. and Cartagena, M.C. 1999. Comparison of Mediterranean Dung Beetles

(Coleoptera: Scarabaeoidea) in Cattle and Rabbit Dung. Environmental Entomology,

Volume 28, Issue 3: 420–424. https://doi.org/10.1093/ee/28.3.420

[54] Gollan, J. R., Reid, C. A. M., Barnes, P. B. and Wilkie, L. 2011. The ratio of exotic-to-

native dung beetles can indicate habitat quality in riparian restoration. Insect

Conservation and Diversity, 4(2): 123–131

[55] Gascon, C., Lovejoy, T. E., Bierregaard, J. R R.O., Malcolm, J. R., Stouffer, P.C.,

Vasconcelos, H. L., Laurence, W. F., Zimmerman, B., Tocher, M. and Borges, S. 1999.

Matrix habitat and species richness in tropical forest remnants. Biological Conservation,

91: 223-229

[56] Gaikwad, A. R. and Bhawane, G. P. 2016. Observation on life cycle and nesting

behaviour of dung beetle Onthophagus catta (Fabricius). International Journal of

Zoology Studies, 1(7): 09-13.

javascript:;

javascript:;

https://doi.org/10.1093/ee/28.3.420


-116-

[57] Geńier, F., and Kohlmann. B. 2003. Revision of the Neotropical dung beetle genera

Scatimus Erichson and Scatrichus gen. nov. (Coleoptera: Scarabaeidae, Scarabaeinae).

Fabreries, 28: 57–111

[58] Halffter, G. and Arellano, L. 2002. Response of dung beetle diversity to human-induced

changes in a tropical landscape. Biotropica, 34: 144-154

[59] Halffter, G. and Edmonds, W. D. 1982. The nesting behavior of dung beetles

(Scarabaeinae): An ecological and evolutive approach. México D.F., Man and the

Biosphere Program UNESCO, 177 pp.

[60] Halffter, G. and Matthews, E.G. 1966. The natural history of dung beetles of the

subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomologica Mexicana,

12/14: 1-312.

[61] Halffter, G., Favila, M. E. and Halffter, V. 1992. A comparative study of the structure

of the scarab guild in Mexican tropical rain forest and derived ecosystems. Folia

Entomologica Mexicana, 84: 131–156

[62] Halffter, G., Favila, M.E. and Arellano, L. 1995. Spatial distribution of three groups of

coleoptera along an altitudinal transect in the Mexican transition zone and its

biogeographical implications. Elytron, 9: 151–185

[63] Halffter, G. and Favila, M. E. 1993. The Scarabaeidae (Insecta: Coleoptera) an animal

group for analyzing, inventorying and monitoring biodiversity in tropical rainforest and

modified landscapes. Biology International, 27: 15-21

[64] Hammer, O. 1941. Biological and ecological investigations on flies associated with

pasturing cattle and their excrement. Vidensk. Meddel. Dansk Naturhist. Foren, 105:

143–393

[65] Hanski, I. 1982. Dynamics of regional distribution: the core and satellite species

hypothesis. Oikos, 38: 210–221

[66] Hanski, I. 1989. Dung beetles. In Tropical rain forest ecosystems (eds. H. Leith and J.

A. Werner), Elsevier Publishers, Amsterdam, Holland: 489–511

[67] Hanski, I. and Cambefort, Y. 1991a. Spatial processes. In: I Hanski & Y. Cambefort

(Eds.). Dung beetle ecology. Princeton University Press, Princeton: 283-304

[68] Hanski, I and Cambefort, Y. 1991b. Competition in dung beetles. In: Hanski, I. & Y.

Cambefort, Y. (Eds.). Dung beetle ecology. Princeton University Press, Princeton: 305-

329

[69] Hanski, I. and Koskela, H. 1979: Resource partitioning in six guilds of dung-inhabiting

beetles (Coleoptera). Annals Entomologica Fennica, 45: 1–12

[70] Hanski, I. and Krikken, J. 1991. Dung beetles in tropical forests in south-east Asia. In: I.

Hanski, & Y. Cambefort, Y. (Eds.). Dung beetle ecology. Princeton University Press,

Princeton, 179-197

[71] Hanski, I. and Niemela, J. 1990. Elevational distributions of dung and carrion beetles in

northern Sulawesi, 145–152. In: Rain Forest Insects of Wallacea. W. J. Knight and J. D.

Holloway (Eds.). Royal Entomological Society, London.


-117-

[72] Harpootlian, P. J. 2001. Scarab Beetles (Coleoptera: Scarabaeidae) of South Carolina.

Clemson University Public Service Publication, Clemson, SC.

[73] Helgesen, R. G. and Post. R. L. 1967. Saprophagous Scarabaeidae (Coleoptera) of

North Dakota. Available online at www.ndsu.edu/fileadmin/

entomology/Publications/007a.pdf (Accessed on 15.5.2019)

[74] Hill, C. J. 1996. Habitat specificity and food preferences of an assemblage of tropical

Australian dung beetles. Journal of Tropical Ecology, 12: 449–460

[75] Holter, P. 2004: Dung feeding in hydrophilid, geotrupid and scarabaeid beetles:

Examples of parallel evolution. European Journal of Entomology, 101: 365–372

[76] Horgan, F. G. 2005. Effects of deforestation on diversity, biomass and function of dung

beetles on the eastern slope of the Peruvian Andes. Forest Ecology and Management,

216: 117–133

[77] Howden, H. F. and Nealis, V. G. 1975. Effects of clearing in a tropical rain forest on the

composition of the coprophagous scarab beetle fauna (Coleoptera). Biotropica, 7: 77–83

[78] Howden, H. and Howden, A. 2001. Change Through Time: a Third Survey of the

Scarabaeinae (Coleoptera: Scarabaeidae) at Welder Wildlife Refuge. The Coleopterists

Bulletin, 55 (3): 356-362

[79] Howden, H. F. and Scholtz, C. H. 1986. Changes in a Texas dung beetle community

between 1975 and 1985 (Coleoptera: Scarabaeidae, Scarabaeinae). The Coleopterists

Bulletin, 40(4): 313–316

[80] Ishikawa, H. 2011. Effects of dung beetles on seedling emergence from herbaceous

seeds in the dung of sika deer (Cervus nippon) in a temperate Japanese grassland

ecosystem. Ecological Research, 26(4): 725-734

[81] Jadhav, M.J. and Sharma, R.M. 2012. Insecta: Coleoptera: Scarabaeidae: Scaeabaeid

beetles. Fauna of Maharashtra, State Fauna Series 20 (Part-2), Zoological Survey of

India, Kolkata: 489-494.

[82] Jankielsohn, A., Scholtz, C. H. and Louw, S. V. D. M. 2001. Effect of habitat

transformation on dung beetle assemblages: A comparison between a South African

nature reserve and neighbouring farms. Environmental Entomology, 30: 474–483

[83] Kadiri, N., Lumaret, J. P. and Floate, K. D. 2014. Functional diversity and seasonal

activity of dung beetles (Coleoptera: Scarabaeoidea) on native grasslands in southern

Alberta, Canada. The Canadian Entomologist, 146 (3), 291–305

[84] 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

[85] Kaufman, P. E. and Wood, L. A. 2012. Indigenous and exotic dung beetles (Coleoptera:

Scarabaeidae and Geotrupidae) collected in Florida cattle pastures. Annals of the

Entomological Society of America, 105(2): 225–231

[86] Kessler, H., Balsbaugh, E. U. J. and McDaniel, B.1974. Faunistic comparison of adult

coleoptera recovered from cattle and sheep manure in east central South Dakota.

Entomological News, 85: 67–71


-118-

[87] Kirk, A. A. 1992. The effect of the dung pad fauna on the emergence of Musca

tempestiva (Dipt, Muscidae) from dung pads in southern France. Entomophaga, 37(4):

507–514

[88] Klein, B. C.1987. Effects of forest fragmentation on dung and carrion beetle

communities in central Amazonia. Ecology, 70: 1715-1725

[89] Klein, B.1989. Effects of forest fragmentation on dung and carrion beetle communities

in central Amazonia. Ecology, 70: 1715–1725

[90] Koskela, H. 1972: Habitat selection of dung-inhabiting Staphylinids (Coleoptera) in

relation to age of the dung. Annales Zoologici Fennici, 9: 156–171

[91] Koskela, H. and Hanski, I. 1977. Structure and succession in a beetle community

inhabiting cow dung. Annales Zoologici Fennici, 14: 204-223

[92] Korasaki, V., Lopes, J., Brown, G. and Louzada, J. 2013. Using dung beetles to evaluate

the effects of urbanization on Atlantic Forest biodiversity. Insect Science, 20(3): 393–

406

[93] Krantz, G.W. 1983. Mites as biological control agents for dung-breeding flies, with

special reference to the Macrochelidae. In: Biological Control of Pests by Mites (Eds.

M.A. Hoy, G.L. Cunningham & L. Knutson), University of California, Berkeley, pp.

91-98.

[94] Krüger, K. and Scholtz, C. H. 1997. Possible risk posed by drift of the insect growth

regulator pyriproxyfen to the rare dung beetle Circellium bacchus (F.) in a national

park. Journal of insect Conservation, 1: 215–220

[95] Kudavidanage, E.P. and Lekamge, D. 2012. A provisional checklist of dung beetles

(Coleoptera; Scarabaeidae) in Sri Lanka, pp.438-444. In The National Red List 2012 of

Sri Lanka; Conservation Status of the Fauna and Flora (Eds. D.K. Weerakoon and S.

Wijesundara). Ministry of Environment, Colombo, Sri Lanka.

[96] Landin, B.O. 1961. Ecological studies on dung-beetles (Col.Scarabaeidae). Opuscula

Entomologica Supplement, 19: 1–227

[97] Levesque, C. and Levesque, G. Y. 1995. Abundance and flight activity of some

Histeridae, Hydrophilidae and Scarabaeidae (Coleoptera) in southern Quebec, Canada.

Great Lakes Entomologist, 28(1): 71–80

[98] Liu, R. T., Zhao, H. L. and Zhao, X. Y. 2012. Influence of grazing exclusion on soil

macroinvertebrate diversity in degraded sandy grassland (Inner Mongolia, China).

Polish Journal of Ecology, 60(2): 375–385

[99] Lobo, J. M., Martín-Piera, F. and Veiga, C. M. 1988. Las trampas pitfall con sebo, sus

posibilidades en el estudio de lascomunidadescoprófagas de Scarabaeoidea (Col.). I.

Características determinantes de sucapacidad de captura. Revue d`ècologie et de

biologie du sol, 25: 77-100

[100] Lobo, J. M., Lumaret, J. P. and Jay-Robert, P. 1998. Sampling dung beetles in the

French Mediterranean area: effects of abiotic factors and farm practices. Pedobiologia,

42: 252–266


-119-

[101] Lobo, J. M., Lumaret, J. P. and Jay-Robert, P. 2001. Diversity, distinctiveness and

conservation status of the Mediterranean coastal dung beetle assemblage in the Regional

Natural Park of the Camargue (France). Diversity and Distributions, 7: 257–270

[102] Matheson, M. M. 1987. Insects associated with cattle dung in southern Québec. M.Sc.

thesis, McGill University, Montreal, Canada.

[103] Masan, P. and Halliday, B. 2009. Mesostigmatid mites associated with the dung beetle

Copris lunaris (Coleoptera: Scarabaeidae). European Journal of Entomology, 106: 545–

550

[104] McGeoch, M.A., Rensburg, B.J.V, and Botes, A. 2002.The verification and application

of bioindicators: a case study of dung beetles in a savanna ecosystem. Journal of

Applied Ecology, 39: 661-672

[105] Medina, C. A., Escobar, F. and Kattan, G.H. 2002. Diversity and habitat use of dung

beetles in a restored Andean landscape. Biotropica, 34(1): 181-187

[106] Menendez, R., and D. Gutierrez. 2004. Shifts in habitat associations of dung beetles in

northern Spain: Climate change implications. Ecoscience, 11(3): 329–337

[107] Miranda, C. H. B., Dos Santos, J. C. C. and Bianchin, I. 1998. Contribution of

Onthophagus gazella to soil fertility improvement by bovine fecal mass incorporation

into the soil. Greenhouse studies. Revista Brasileira De Zootecnia - Brazilian Journal of

Animal Science, 27 (4): 681–685

[108] Moczek, A.P. and Niijhout, H.F. 2003. Rapid evolution of a polyphenic threshold.

Evolution & Development, 5: 259-268

[109] Montes de Oca E. 2001. Escarabajos coprófagos de un escenario ganadero típico de la

región de los Tuxtlas, Veracruz, México: importancia del paisaje en la composición de

un gremio functional. Acta Zool. Mex. (n.s.) 82: 111-132 (2001)

[110] Newton, P.N. and Malcolm, J.C. 1985. Dung and dung beetles in Kanha Tiger Reserve,

Central Indian Highlands. Journal Bombay Natural History Society, 85: 218-220

[111] Nichols, E., Spector, S., Louzada, J., Larsen, T., Amezquita S. and Favila, M. E. 2008.

Ecological functions and ecosystem services provided by Scarabaeinae dung beetles.

Biological Conservation, 141: 1461-1474

[112] Nealis, V. G. 1977. Habitat associations and community analysis of south Texas dung

beetles (Coleoptera: Scarabaeinae). Canadian Journal of Zoology, 5: 138-147

[113] Nummelin, M. and Hanski, I. 1989. Dung beetles of the Kibale Forest, Uganda: a

comparison between virgin and managed forests. Journal of Tropical Ecology, 5: 349–

352

[114] Numa, C., Verdu, J. R., Rueda, and Galante, E. 2012. Comparing dung beetle species

assemblages between protected areas and adjacent pasturelands in a Mediterranean

savanna landscape. Rangeland Ecology & Management, 65(2): 137–143

[115] O’Hea, N. M., Kirwan, L. and Finn, J. A. 2010. Experimental mixtures of dung fauna

affect dung decomposition through complex effects of species interactions. Oikos,

119(7): 1081–1088


-120-

[116] Olechowicz, E. 1974. Analysis of a sheep pasture ecosystem in the Pienieny mountains

(the Carpathians). Ekologia Polska, 22: 589–616

[117] Olechowicz E. 1976. The role of coprophagous dipterans in a mountain pasture

ecosystem. Ekologia Polska, 24: 125–165

[118] Pakaluk, J., Slipinski, S. A. and Crowson, R. A.1995. Biology, Phylogeny, and

Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson.

Muzeum i Instytut Zoologii PAN, Warszawa, Poland.

[119] Pimsler, M. L. 2007. A survey of the dung beetles in cattle manure on pastures of an

organic and a conventional dairy farm in New York State. Honors Thesis, Cornell

University, Ithaca, NY.

[120] Price, D. L. 2004. Notes on the Scarabaeoid dung beetles (Coleoptera: Scarabaeidae,

Geotrupidae, and Trogidae) of Hutcheson Memorial Forest. New Jersey Entomological

News, 117: 347–350

[121] Price, D. L., Brenneman, L. M. and Johnston, R. E. 2012. Dung beetle (Coleoptera:

Scarabaeidae and Geotrupidae) communities of eastern Maryland. Proceedings of the

Entomological Society of Washington, 114(1): 142–151.

[122] Priyadarsanan, D.R. 2006. Insecta: Coleoptera: Scarabaeoidea: Scarabaeidae (Dung

beetles), pp 91-135. Thirumalai, G. & E. Krishnan (Eds.). Fauna of Biligiri

Rangaswamy Temple Wildlife Sanctuary, Conservation Area Series 27, Zoological

Survey of India, 263 pp.

[123] Psarev, A. M. 2002. Succession in an insect community inhabiting horse dung. Russian

Entomological Journal, 11: 287–290

[124] Putman, R. J.1983. Carrion and Dung: The decomposition of animal waste, Studies in

Biology, 156, Institute of Biology, London, UK: 62.

[125] Qie, L., Lee, T. M., Sodhi, N. S. and Lim. S. L. H. 2011. Dung beetle assemblages on

tropical landbridge islands: small island effect and vulnerable species. Journal of

Biogeography, 38(4): 792–804

[126] Quintero, I. and Roslin, T. 2005. Rapid recovery of dung beetle communities following

habitat fragmentation in central Amazonia. Ecology, 12: 3303–3311

[127] Ratcliffe, B. C., and Jameson, M. L. 2005. Generic guide to New World scarab

beetles.Availablefrom:museum.unl.edu/research/entomology/Guide/Guide-

introduction/Guideintro.html (Accessed on 15.12.2020)

[128] Ratcliffe, B. C., and Jameson, M.L. 2013. Generic guide to New World scarab beetles.

Availablefrom:museum.unl.edu/research/entomology/Guide/Guideintroduction/Guidein

tro.html (Accessed on 15.12.2020)

[129] Radtke, M.G., Carlton, C. E. G., Williamson, B. 2008. A Dung Beetle Assemblage in an

Urban Park in Louisiana. Southeastern Naturalist, 7 (1): 101-110

[130] Raychaudhuri, D. and Saha, S. (Eds.) 2014. Atlas of Insects and Spiders of BuxaTiger

Reserve. Publ. West Bengal Biodiversity Board and Nature Books India, Kolkata: 357

pp.


-121-

[131] Rodriguez, J. P., Pearson, D. L. and Barrera, R. 1998. A test for the adequacy of

bioindicator taxa: are tiger beetles (Coleoptera: Cicindelidae) appropriate indicators for

monitoring degradation of tropical forests in Venezuela. Biological Conservation, 83:

69–76

[132] Roslin, T. and Koivunen, A. 2001. Distribution and abundance of dung beetles in

fragmented landscapes. Oecologia, 127: 69–77

[133] Rounds, R. J., and Floate, K. D. 2012. Diversity and seasonal phenology of

coprophagous beetles at Lake City, Michigan, USA, with a new state record for

Onthophagus taurus (Schreber) (Coleoptera: Scarabaeidae). The Coleopterists Bulletin,

66(2): 169–172

[134] Sarkar, S. 2013. Scarabaeidae (Coleoptera) of Buxa Tiger Reserve. Ph.D. Thesis (Sc.),

Univ of Calcutta, 263pp.

[135] Sarkar, S., Saha, S. and Raychaudhuri, D. 2015. On the taxonomy of scarabaeine fauna

(Coleoptera: Scarabaeidae) of Buxa Tiger Reserve (BTR), West Bengal, India. Munis

Entomology & Zoology, 10(1): 18-48

[136] Sathiandran, N., Thomas, S.K. and Flemmimg, 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

[137] ScarabNet, 2009. ScarabNet Global Taxon Database Version 1.5 online at

http://www.scarabnet.org (accessed on 15.12.2020)

[138] Scholtz, C., J. du G. Harrison, and V. V. Grebennikov. 2004. Dung beetle [Scarabaeus

(Pachysoma)] biology and immature stages: reversal to ancestral states under desert

conditions (Coleoptera: Scarabaeidae)? Biological Journal of the Linnean Society, 83:

453–460

[139] Schulze, C.H., Shahabuddin and Tscharntke, T. 2005.Changes of dung beetle

communities from rainforests towards agroforestry systems and annual cultures in

Sulawesi (Indonesia). Biodiversity Conservation, 14(4): 863–77

[140] Shahabuddin. 2011. Effect of land use change on ecosystem function of dung beetles;

experimental evidence from Wallacea Region in Sulawesi, Indonesia. Biodiversities,

12(3): 177-181

[141] Simmons L.W. and Ridsdill-Smith, T.J. 2011. Ecology and evolution of dung beetles.

Publ. Blackwell Publishing Ltd., 347pp.

[142] Singh, A.P., Mahajan, S., De, K., Uniyal, V.P. and Bhardwaj, A.K. 2018. An

Assessment of Coleopteran Fauna of the forest of Siswan, Punjab. Indian Forester,

144(11): 1107-1113

[143] Skidmore, P. 1991. Insects of the British cow dung community: Field Studies Council.

Occasional Publications, 21: 1-166

[144] Šlachta, M., Frelich, J. and Svoboda, I. 2008. Seasonal biomass distribution of dung

beetles (Scarabaeidae, Geotrupidae, Hydrophilidae) in mountain pastures of South-West

Bohemia. Journal of Agrobiology, 25: 163–176

http://www.scarabnet.org/


-122-

[145] Slade, E. M., Mann, D. J., Villanueva, J. F. and Lewis, O. T. 2007. Experimental

evidence for the effects of dung beetle functional group richness and composition on

ecosystem function in a tropical forest. Journal of Animal Ecology 76(6): 1094–1104

[146] Smetana, A. 1978. Revision of the subfamily Sphaeridiinae of America north of Mexico

(Coleoptera: Hydrophilidae). Memoirs of the Entomological Society of Canada, 110: 1–

292

[147] Sowig, P.1995. Habitat selection and offspring survival rate in three paracoprid dung

beetles: the influence of soil type and soil moisture. Ecograpahy, 18:147–154.

[148] Sowig, P. and Wassmer, T. 1994. Resource partitioning in coprophagous beetles from

sheep dung: Phenology and microhabitat preferences. Zoologische Jahrbucher

Abtheilung für Systematik, 121: 171-192

[149] Spector, S. and Ayzama, S. 2003. Rapid turnover and edge effects in dung beetle

assemblages (Scarabaeidae) at a Bolivian Neotropical forest–savanna ecotone.

Biotropica, 35: 394–404

[150] Spector, S. and Forsyth, A. B. 1998. Indicator taxa for biodiversity assessment in the

vanishing tropics.Conservation in a Changing World (Eds G. Mace, A. Balmford & R.

Ginsberg), 181–209.

[151] Trumbo, S. T. and Fernandez, A. G. 1995. Regulation of Broodsize by male parents and

cues employed to access resource size by burying beetles. Ethology, Ecology &

Evolution, 7: 313-322

[152] Valiela, I. 1969. Composition, food webs and population limitation in dung arthropod

communities during invasion and succession. American Midland Naturalist, 92(2): 370–

385

[153] Valiela, I. 1974. Composition, food webs and population limitation in dung arthropod

communities during invasion and succession. American Midland Naturalist, 92(2): 370–

385

[154] Vaz-de-Mello, F. Z. 2003. Species formerly in the genera Trichillum Harold, 1868 and

Pedaridium Harold, 1868 (Coleoptera: Scarabaeidae). M.Sc. Thesis, Universidade

Federal de Lavras, Lavras, Brazil.

[155] Vaz-de-Mello, F. Z. 2007. Revisiońtaxono´mica y ana´ lisisfilogene´tico de la tribu

Ateuchini (Coleoptera: Scarabaeidae: Scarabaeinae). D.Sc. Thesis, Instituto de

Ecologıá, A.C., Xalapa, Mexico

[156] Veenakumari, K & Veeresh, G.K. 1997. Dung beetles (Coleoptera: Scarabaeidae:

Scarabaeinae) fauna of Bangalore, Karnataka. Journal of the Bombay Natural History

Society, 94(1): 171-173

[157] Wassmer, T. 2014. Seasonal occurrence (Phenology) of coprophilous beetles

(Coleoptera: Scarabaeidae and Hydrophilidae) from cattle and sheep farms in

southeastern Michigan, USA. The Coleopterists Bulletin, 68(3): 603–618

[158] Waterhouse, D. F. 1974. Biological control of dung. Scientific American, 230(4): 100–

109


-123-

[159] Woodruff, R. E.1973.The Scarab Beetles of Florida (Coleoptera: Scarabaeidae).

Volume 8. Florida Dept. of Agriculture and Consumer Services, Division of Plant

Industry, Gainesville, FL.

[160] Wu, X., Duffy, J. E, Reich, P. B. and Sun, S. 2010. A brown-world cascade in the dung

decomposer food web of an alpine meadow: effects of predator interactions and

warming. Ecological Monographs 81(2): 313–328

[161] Wu, X.W., and Sun, S. C. 2012. Artificial warming advances egg-laying and decreases

larval size in the dung beetle Aphodius erraticus (Coleoptera: Scarabaeidae) in a Tibetan

alpine meadow. Annales Zoologici Fennici, 49(3): 174–180

Dung beetles: key to healthy pasture? An overvie · 2021. 1. 4. · extremely high (Hanski & Cambefort, 1991a,b). World Scientific News 153(2) (2021) 93-123 -95- Fig. 1. Dung beetle

Documents