Diversity and
community structure of dung beetles (Coleoptera: Scarabaeinae) associated with
semi-urban fragmented agricultural land in the Malabar coastin southern India
K. Simi Venugopal 1, Sabu K. Thomas 2& Albin T. Flemming 3
1,3Post Graduate and Research Department of Zoology, Loyola College,
Chennai, Tamil Nadu 600034, India
2 Post
Graduate and Research Department of Zoology, St. Joseph’s College, Devagiri,
Kozhikode, Kerala 673008, India
Email: 1 simisachin@gmail.com, 2 sabukthomas@gmail.com
(corresponding author), 3 dratfleming@gmail.com
Date of publication
(online): 26 July 2012
Date of publication (print):
26 July 2012
ISSN 0974-7907 (online) |
0974-7893 (print)
Editor: B.B.
Hosetti
Manuscript details:
Ms # o3074
Received 20 January 2012
Final received 09 May 2012
Finally accepted 29 May 2012
Citation:Venugopal, K.S., S.K. Thomas & A.T. Flemming (2012). Diversity and
community structure of dung beetles (Coleoptera: Scarabaeinae) associated with
semi-urban fragmented agricultural land in the Malabar coastin southern India. Journal of Threatened Taxa 4(7):
2685–2692.
Copyright: © K.
Simi Venugopal, Sabu K. Thomas & Albin T. Flemming 2012. Creative
Commons Attribution 3.0 Unported License. JoTT allows unrestricted use
of this article in any medium for non-profit purposes, reproduction and
distribution by providing adequate credit to the authors and the source of
publication.
Author Details: Mrs. K. Simi Venugopal is pursuing her PhD
programme on the ecology and systematics of dung beetles at Post Graduate and
Research Department of Zoology, Loyola College, Chennai, Tamil Nadu, India.
Dr. Sabu K. Thomas is an
associate professor attached to Post Graduate and Research Department of
Zoology, St. Joseph’s College, Devagiri, Kozhikode, Kerala and systematics and
ecology of ground beetles in the moist south Western Ghats is his thrust area.
Dr. Albin T. Fleming,
associate professor and head of the Post Graduate and Research Department of
Zoology, Loyola College, Chennai, Tamil Nadu and is actively engaged with the
systematic of various insect groups in peninsular India.
Author Contribution:Taxonomic analysis, sampling and data analysis by the first and second authors; discussion by all the three authors.
Acknowledgments: The
financial assistance provided by UGC (University Grants Commission, India), is
gratefully acknow-ledged.
Abstract:An evaluation
of the diversity and community structure of dung beetles associated with
semiurban agricultural land in the Malabar coast of southern India revealed
that urbanization has led to decreased diversity compared to regional forests,
and has affected the community status of dung beetles. However, contrary to expectations,
species richness was observed to be equivalent to rural agricultural fields in
the region. Low abundance of prominent
agricultural habitat species indicates that the study area has changed as a
result of habitat modification/urbanization, and the prevailing conditions are
not ideal for the establishment of the most common species in agriculture
belts. Prominence of two less
common species, Tiniocellus
spinipes and Caccobius
vulcanus, indicates these generalist urban adaptable (synanthropic) species
have become increasingly widespread and locally abundant. The low abundance of tunnelers and
rollers is attributed to fragmentation of the urban agricultural belt, low
mammalian diversity and dung availability, and the hard nature of the laterite
soil in the Malabar coast region.
Keywords: Dung beetles, fragmentation, Malabar coast, southern India, Tiniocellus spinipes, urbanization.
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INTRODUCTION
Destruction and deterioration of natural
habitats associated with urbanization has led to dramatic changes in the biotic
structure and composition of ecological communities. Observations include decreased abundance
and diversity, disappearance or replacement of indigenous species by
non-natives (Blair 1996, 2004; La Sorte & Boecklen 2005) and habitat
specialists (Magura et al. 2010), and local extinctions (Raupp et al. 2010). In many places, although
highly modified and disturbed, small urban fragments of agricultural lands in
the midst of urban environments have been identified as an important source of
native biodiversity (Gaston et al. 2004). The different fauna found in small urban fragments may be a consequence
of any of a number of pressures associated with fragmentation and urbanization,
including increased anthropogenic disturbance, reduced area, loss of hosts,
invasion of new species and release of natural enemies (Yahner 1988). Such areas can provide ephemeral or more
permanent habitats for species, dispersal corridors or resting places for
migrating organisms (Gaston et al. 2005). Therefore, it is important to document the status of biodiversity
prevailing in other areas to identify the level of biodiversity still left in
urban areas and characterize the remaining elements of
the original biota (e.g. are they specialist or generalist). In the present effort we aim to
determine the community structure of dung beetles in a small isolated
agricultural land in the midst of urban settlements in the coastal Malabar
region.
The Malabar coast moist deciduous forests ecoregion—hereafter
referred as MCF—was a swath of lush tropical evergreen forest that
extends along the Western Ghats mountains and the Arabian Sea. MCF
represents an extreme example of deforestation in the Western Ghats, having
undergone major ecological transformations over the last 100 years (Nair 1991;
Wikramanayake et al. 2002). MCF has
lost more than 95% of its original vegetation to deforestation during the
British rule (Colonial period), cash crop cultivation during the post Colonial
period and recent urbanization. Currently,
due to the recent wave of urbanization, the agricultural lands are being transferred into urban
jungles at alarming rates in the MCF. It is certain that the remaining
original biota that took shelter in these pockets will be lost soon. No records exist about the impact of
anthropogenic activity on biodiversity in the region and hence we lack crucial
historical documentation of the natural communities in MCF
which would remain as an important source of information for measuring
species extinctions in the area (Brook et al. 2003). The present effort aims to gather data
on the composition and guild structure of dung beetle assemblage associated
with a fragmented agricultural landscape in the midst of an urban environment
in Kozhikode region in the MCF. We selected dung
beetles because they showed significant changes in species composition and
community assemblage following forest fragmentation and habitat disturbances
(Nichols et al. 2007), making them excellent biodiversity indicators for
examining the responses of species communities to anthropogenic disturbance
(Gardner et al. 2008a,b). We
propose that the regional dung beetle fauna might not have been affected by
urbanisation, disappearance of native mammals and an unchanged native
assemblage with high diversity and abundance exists in the region.
MATERIALS AND METHODS
Study area
Selected
study site was an open agricultural field (11015’N & 75048’E)
of predominantly coconut plantation with the intervening grasslands close to
Devagiri College campus, Kozhikode used for cattle and sheep grazing. Annual temperature 24–32 0C;
relative humidity 40–80 %; average rainfall 750–1500 mm/year which
occurs mostly in the wet months of June to November (CWRDM 2008-09).
Sampling
Dung beetles were collected using dung
baited pitfall traps of the bait-surface-grid type on a seasonal basis during
southwest monsoon (June–August), northeast monsoon
(September–November), presummer (December–February) and summer
(March–May) periods from June 2008 to May 2009. Pit fall traps were made of plastic
basins, 10cm in diameter and 15cm deep and a mixture of water-formalin-liquid
soap mixture were used as preservative. The basins were buried with their rim in level with the surrounding
substrate and each trap was topped with a plastic plate supported on iron bars
to prevent desiccation during sunny days and inundation during the periods of
rain. Two hundred grams of fresh cow dung was placed on a wire grid between the
basin and the tray. Ten such traps
at 50m intervals along a linear transect were placed following the standardized
dung-beetle sampling design of maintaining a minimum distance of 50m between
traps to minimize trap interference (Larsen & Forsyth 2005). Beetles were collected at 0600 and 1600
hr each day. Both diurnal and
nocturnal collections were made separately.
Beetles were identified to species levels
using taxonomic keys available in Arrow (1931), Balthasar (1963) and by
comparing with the verified specimens. After
identification, specimens were deposited in the insect collections of St.
Joseph’s College, Devagiri, Kozhikode. The species were sorted into three
functional guilds - dwellers (endocoprids), rollers (telecoprids) and tunnelers
(paracoprids) following Cambefort & Hanski (1991) and three temporal guilds
(noctural/diurnal/generalists) following Krell et al. (2003). Species that were present during all
seasons with >10% abundance were treated as major groups, <10% were
treated as minor groups and singletons as rare groups. Based on size, dung beetles are
classified into small beetles (with size 5mm and less) and large beetles (5.1
and 30 mm).
Data
analysis
Diversity (Magurran 2004) was estimated using Shannon–Weaver
diversity index (Shannon & Weaver 1949), evenness with Simpson’s evenness
index (Simpson 1949) and richness with Margalef’s species richness index (d). Data used for statistical analysis were
tested for normality with GRETL open source software version 1.1 (Cottrell
2006). Significant levels of
variation in the overall and species-wise abundance with seasons were tested with Kruskal-Wallis followed
by Mann-Whitney U test (Weiss 2007). All statistical data analyses were done
with Mega Stat Version 10.0 software (Orris 2005) and diversity analysis with Primer v5.2.9 software.
RESULTS
Species richness and diversity
A total of 519 dung beetles representing
26 species, belonging to eight genera and five tribes were recorded. Assemblage consisted of three major
species, 17 minor and six rare species. Tiniocellus spinipes (44.89%)
and Caccobius vulcanus (17.92%) dominated the assemblage (Image 1). Large and small size beetles varied in
abundance (Kruskal-Wallis test, H = 8.64, df = 1, P<0.05). Small sized
beetles represented by 10 species (67.83% of total abundance) dominated the
assemblage over the large sized beetles represented by 15 species (Table 1). The diversity (H) was 2.015, Margalef
species richness index value (d) was 9.75, dominance index value (λ) was
0.24 and evenness (1-λ) was 0.751. Rank of each species based on relative abundance is represented in Fig.
1.
Functional and temporal guild composition
Dwellers were the most abundant guild
(53.75%) with T. spinipes and Tibiodrepanus setosus as dominant
species followed by tunnelers (41.42%) and rollers (3.85%) (H=53.46, df=2, P<0.05). Tunnelers, represented by 20 species (76.92%) were the most speciose
guild followed by rollers and dwellers represented by three species each
(11.53%) (Table 1). Six species were nocturnal, five were
diurnal and 15 were generalists. Diurnal guild dominated the assemblage followed by nocturnal and
generalist (H=20.01, df=2, P <0.05) (Table 1).
Seasonality
Highest abundance was recorded during
southwest season followed by northeast and summer season (H=45.33, df=3,
P<0.05). Among the three guilds,
tunnelers were seasonal (H=3.81, df=3, P <0.05) with high abundance during
southwest monsoon period followed by northeast monsoon, summer and presummer
periods. Dwellers (H=4.39, df=3, P >0.05) and rollers (H=11.42, df=3, P
>0.05) did not show any seasonal variation. Eight species (one roller, one dweller
and six tunnelers) showed seasonal variation in abundance (Table 2).
DISCUSSION
Evaluation of the diversity and community
structure of the dung beetles associated with the semi-urban agricultural lands
revealed that urbanization does cause a decrease in dung beetle diversity
compared to regional forests and has affected the community status of dung
beetles. However, contrary to the
expectations, species richness was in par with the rural agrifields in the
region. Following patterns, namely,
(1) dominance of the community by two species (T. spinipes andC.vulcanus) recorded rarely from the moist forests and agrifields, and the
record of five rare forest specialist species Ochicanthon murthyi, Onthophagus
insignicollis, O. kchatriya, O. duporti and Oniticellus cinctus(Arrow 1931; Sabu 2011) and (2) low abundance of prominent species in the
regional agrilands and forests namely Caccobius meridionalis, Onthophagus
dama, O. turbatus, O. falsus and O. fasciatus (Sabu 2011) were
observed.
Community structure of the assemblage was
highly uneven with the dominance of two species (T. spinipes and C.
vulcanus). Uneven distribution
of species is relatively common in unstable environments and point towards
extreme disturbance (Magurran 2004). Tiniocellus spinipes and C. vulcanus are rare in the
agriculture and forest belts of the Western Ghats, and the most abundant
species are Caccobius meridionalis, Onthophagus dama, O. turbatus, O. falsusand O. fasciatus (Sabu 2011). Low abundance of these prominent agri habitat species indicates that the
study region has changed as a result of the habitat modification/urbanization
and the prevailing conditions in the study region is not ideal for the
establishment of even the most common and hence the most adapted species in the
agriculture belts. Two dominant
species in the study region namely T. spinipes and C. vulcanusare the prominent species in the open semi-arid dry north western part of the
Indian subcontinent with a long history of habitat modification (Arrow 1931;
Balthasar 1963; Mittal 2005; Kakkar & Gupta 2010). Higher abundance of two species common
in the open and dry belts in northern India and low abundance in the moist
regional forest and agri belts indicate them to be heliophilic, open habitats
adapted and disturbance tolerant species. Dominance of such heliophilic, dry habitats species in the moist study
region indicates that the study region underwent extreme modifications and the
utility of these two species can be seen as indicators of habitat change in the
MCF in general. Cosmopolitan
distribution and common occurance of T. setosus in the forests and
agribelts in the moist Western Ghats (Sabu 2011) and in the present study MCF site which is
an isolated urban agribelt indicates that T. setosus is a generalist
species with capacity to persist in a wide array of habitat types. Distinctly low representation of forest
specialist species supports the ‘habitat specialist hypotheses’ (Magura et al.2008), which suggested that ‘species richness of forest specialists should
decrease from the rural areas to the urban ones and different elements of an
assemblage will react differently, because the specialists are expected to
decrease with urbanization while the generalist species could be favored (Magura
et al. 2004). Results
indicate that T. spinipes and C. vulcanus the generalist urban
adaptable (synanthropic) species adapted to the modified urban habitats that
have become increasingly widespread and locally abundant. It is likely that these two species may
have become increasingly wide spread and locally
abundant in different urban regions in the Malabar coast region and are likely
to proliferate further in the region at the cost of other species. This type of homogenization of the
taxonomic composition is a major negative consequence of urbanization world wide (Magura et al. 2010).
In total contrast to the dry habitat
dwelling species, occurrence of Ochicanthon species, a rare primitive
old world dung beetle species present in moist forest patches ( Krikken & Huijbregts 2007; Latha et al. 2011) is
unexpected. Its presence indicates
that the recent habitat modifications in the Western Ghats have not wiped out
these relict old world dung beetles (primitive groups) from the agrilands.
Similar record of the following rare species namely Onthophagus
insignicollis, O. kchatriya, O. duporti and Oniticellus cinctus in
the moist Western Ghats indicate that these species represent the sink
population of a larger pool of the source, i.e. the native dung beetle
population. It is possible that the
fragmented agri habitats in the region is a safe
microhabitat for such rare species (Onthophagus malabariensis, O.
unifasciatus, O. pygmaeus, Ochicanthon murthyi) (Magura et al. 2004;
Elek & Lövei 2007) till the gradual disappearance of such habitats.
No record of the large species belonging
to the genera Gymnopleurus, Catharsius and Heliocopris in the
assemblage indicates that they might have vanished from the region. It could be due to the inability of
large dung beetles to withstand dessication and the low survival chance of
their larvae in dry soil conditions (Chown 2001). Since the present study was
confined to a single site, studies in other similar sites in the region are
necessary to establish whether the disappearance of large beetles is a widely
applicable pattern.
Dominance of tunnelers in forest and agri
habitats, and the higher abundance of the cosmopolitian T. setosus is typical of dung beetle assemblages in the Western Ghats
(Sabu et al. 2006, 2007; Vinod & Sabu 2007) and across the globe (Cambefort
& Walter 1991; Andresen 2005). Disappearance leads to the question whether dominance of dweller guild
with T. spinipes as prominent species is a feature of extremely
disturbed habitats in the moist western slopes of the Western Ghats and Malabarcoast region. Distinctively high abundance of dwellers over tunnelers and rollers is
arising from the unequal abundance of T. spinipes. Similar dominance of dwellers with
another species (T. setosus) is reported from regions with high dung pad
availability as in the elephant dung rich Wayanad forests in the Western Ghats
(Vinod 2009) and is attributed to the abundance of elephant/gaur and the
resulting dung pad abundance. However there is no record of dominance of dwellers in the agri belts in
the Western Ghats. High abundance
of dwellers in an agribelt with low dung availability and mammalian diversity
is attributed to the low abundance of other guilds (tunnelers/rollers) and the
hard nature of the laterite soil in the Malabar coast region which do not
favour the population build up of other guilds.
Occurrence of two dominant species T.
spinipes (diurnal) and C. vulcanus (nocturnal) with
distinctly contrasting pattern of temporal resource utlilisation patterns as
prominent species could be an adaptive strategy for efficient resource
partitioning to avoid competition for resources. Such perfect temporal resource partition
of the two prominent species could be a major factor that leads to the decline
in the abundance of other species. Overall abundance showed distinct seasonality with high abundance during
southwest monsoon followed by northeast and summer season. Peak in abundance is linked to the
single species dominance of T. spinipes which was the most abundant
species during southwest monsoon. It is likely that softening of the lateritic soil during the rainy
periods could be favouring the abundance of both tunnelers and dwellers. Seasonality of tunnelers, which was the
only guild showing seasonality in the region, is additional proof to this
assumption.
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