Recolonisation patterns of orthopteran species in successional stages of revegetated coal mine sites

2013·3 MB·

by  Shbbir R Khan

HALTERES,Volume4, 1-11, 2013
ISSN0973-1555©SHBBIR RKHANANDNEELKAMALRASTOGI
Recolonisation patterns of orthopteran species in successional stages
of revegetated coal mine sites
Shbbir R Khan and Neelkamal Rastogi*
InsectBehaviouralEcologyLaboratory, DepartmentofZoology, Centre ofAdvancedStudy,
Banaras Hindu University, Varanasi-221 005, U.P., India
(e-mail: [email protected])
Abstract
The diversity and abundance ofinsects belonging to the order Orthoptera was studied in coal mine spoils of
different ages (0- 10year old) located in Singrauli and the adjoining border area between UttarPradesh and
Madhya Pradesh in India. The abundance of the 21 orthopteran species (belonging to 16 genera) showed
significant variations in 0-8 year old mine sites relative to the 10 year old reference site. Orthopteran alpha
diversity increased sharply in the 2 year old site. The gradual increase in alpha diversity in the 4-10 year old
sites did not demonstrate any significant differences. However, abundance and number of grasshopper
species showed a clear linear increase with increase with mine site rehabilitation. Overlap in orthopteran
rank abundance curves obtained for 4-10 year old sites is probably due to differences in the occurrence,
abundance and diversity patterns ofgrasshopper and cricket species with increase in mine site rehabilitation
age. Our results demonstrate that grasshopper species assemblages are highly sensitive to variations in the
level of anthropogenically-caused disturbance, due to mining activities. Thus, the abundance patterns of
grasshoppers, including those of habitat specialists, may be of significance in evaluation of the restoration
progressindegraded ecosystems, such as mine sites.
Keywords: Grasshoppers, insectdiversity, degradedecosystems, recolonisationpatterns.
Introduction
Anthropogenically-caused environmental the re-establishment ofa functioning ecosystem
degradation and the consequent adverse impact (Majer, 2009).
on biodiversity are of worldwide concern Arthropods are found to occupy a wide
(Bengtsson et al, 2000; Folke et al, 2004; diversity ofmicrohabitats and niches. Moreover,
Domelas, 2010), The restoration of degraded they play more diverse ecological roles than any
^ lands such as mine sites is a priority area in other group of animals. Their short generation
conservation biology (see recent review by times and their small size makes arthropods,
Suding, 2011), since mine spoils (the piles of particularly insects, ideal organisms for
accumulated over-burden excavated during the monitoring spatial and temporal changes in
miningprocess) cause ecosystem degradationby habitat quality and restoration success
destruction of flora and fauna. Though (Longcore, 2003). Hence, among the mine site
succession and recolonisation by vegetation and fauna, insects need to be targeted for
vertebrate wildlife on reclaimed mine lands has assessment ofthe recolonisingpatterns.
been studied (Majer, 1989), diversity patterns of In revegetated mines, the colonising
early colonising invertebrate species needs patterns of insect herbivores assume special
greater attention since they play a vital role in significance, since these may influence
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,
ShbbirRKhan andNeelkamalRastogi
ecosystem succession via herbivory and may sites ofNorthern Coalfields Ltd., are situated at
also reflect the rehabilitation process. Insects Singrauli and the adjoining border area lying
belonging to the order Orthoptera play a critical between Uttar Pradesh and Madhya Pradesh in
N
role in grassland ecosystems, since they India (latitudes 23° 47' to 24° 12' and
constitute the major proportion of arthropod longitudes 81° 48' to 82° 52' E). The vegetation
biomass (Shure andPhillips, 1991). Grasshopper in the mine sites is dominated by Eragrostis
assemblages appear to play the main roles in unioloides, Saccharum spontaneum, Bambusa
forming and maintainingthebiodiversity andthe bamboo, Euphorbia hirta. Acacia catechu.
stability of various kinds of grassland Acacia nilotica, Acacia mangium, Dalbergia
ecosystems (Guo et ah, 2006). Moreover, sissoo and Prosopis juliflora, plants (Singh,
orthopteran nymphs are highly mobile (Floren et 2011,2012).
al., 2001), allowing orthopteran communities to
SamplingMethods
closely follow plant communities with , v , , . u. >
The orthopteran populations were sampled
progressive adjustments during the season
(Joem, 1982; Parmenter et al, 1991; Bonnet et from February to March, 2012 and again during
October - December, 2012. Two main methods
ah, 1997). Grasshopper biodiversity is not only
considered to be a product of the evolution of were used to record the diversity and abundance
grassland ecosystems; a close relationship is also of orthopterans at the six mine sites. These
found between grasshopper biodiversity and the included: (1) the visual scarming, and (2) the
health of grassland ecosystems (Guo et ah, sweep netting, methods. In the visual scan
method, a metal quadrat frame wasusedand, the
2006).
number and diversity of orthopterans were
Hence, the present study focuses on
orthopteran diversity and abundance in coal recorded in the quadrats (area = 1 m^, n = 125
mine spoils of different ages to elucidate the per dayper site) whichwere randomlyplaced on
recolonisation patterns of grasshoppers and the ground surface, over a period offive days (n
crickets following anthropogenic disturbances = 625 per site). The sweep netting method
caused by surface mining activities. Information involved random sweeping (n = 20 per site) of
related to insect recolonisation and successional the ground, bushes and shrubs to capture the
patterns shouldbe useful to land managers inthe orthopterans, by using an insect net. Samples
were preserved in 75% alcohol and taken to the
development of suitable strategies for
laboratory for sorting and identification to
reconstruction ofdisturbedecosystems.
genus/species level. Identification of the
Materials and Methods orthopteran specimens was donewiththehelp of
,
experts from the Department of Zoology,
Study Site AligarhMuslimUniversity.
Investigation of the diversity and
abundance of insects belonging to the order DataAnalysis
Orthoptera was conducted at six coal mine Species diversity (alpha andbeta diversity)
dump sites ofvarious ages (0, 2, 4, 6, 8 and 10 at eachmine site was calculated accordingtothe
years) established from 2001 to 2011. The Shannon-Weiner and Whittaker index methods,
original vegetation is that of a tropical dry respectively.
deciduous forest (Singh, 2012). Forty - six plant Changes in the overall orthopteran
species were planted by the mining company at abundance with mine site rehabilitation was
each dump site about 1.5 years after analysed by carrying out one-way analysis of
establishment ofthe mine dumps. The coal mine variance (ANOVA) followed by Dunnett's and
!iio,t lis:K-tKirw;•; -jjeto, 2
Recolonisationpatternsoforthopteran speciesin revegetated coal mines
Duncan's Multiple Range post hoc Tests fuscovittata, Atractomorpha psittacina
(DMRT) (p < 0.05), tests for analysis of Camarvonella sp., Pterophylla sp., Metioche
abundance and of alpha diversity, respectively. sp., Catantops pinguis, Atractomorpha
Variance followed by DMRT post-hoc test was crenulata, Chrotogonus trachypterus,
also applied to analyse in the abundance of Sphingonotus savignyi, Bryodema luctuosa,
grasshoppers and crickets in the 0 - 10 year old Oedaleus abruptus (One-way ANOVA: F =
sites. Since no pristine areas were available, the 52.823; 21.798; 33.607; 24.252; 18.672; 16.725;
dump site established in 2001 (10 years old) was 5.253; 12.851; 42.073; 14.896; 30.889; 37.003;
taken as the reference site for determination of 7.626; 11.808; 8.584; 9.339; 16.933; 13.384;
beta diversity and for the Duimett's post-hoc 16.170; 18.617; 17.806, respectively, df = 5,
test. 3749; p < 0.001 in each case) (Fig. 1 a-c).
The orthopteran diversity at different sites Dunnett's Post hoc test revealed
was compared using the rank abundance plot in significantly lower abundance ofA. exaltata, G.
which percentage cumulative abundance (log) africanus, L. cruciata, S. prasiniferum, and
wasplottedagainstspeciesrank. Pterophylla sp. in the 0-6 year old sites (p <
All orthopteran species restricted to 0.001, in each case) relative to the reference site.
specific succession stages were classified as The abundance of A. conica, Pyrgomorpha sp.,
habitat specialists. All others consistently Chrotogonus sp., G. musicus, A. crenulata, C.
recorded in 2-10 year old sites were referred to trachypterus and S. savignyi in the early-
ashabitatgeneralists (Table. 1). intermediate successional stages, (0-4 year old)
All statistical analyses were done by using was significantly lower (p < 0.001, in each case)
SPSS (16.0) statistical package (SPSS Inc, relative to their abundance in the 10 year old
Chicago, USA; 1997) andMS Excel 2007. site. In the newly established dump sites (0-2
year old), abundance of C. armatus and C.
Results pinguis was significantly lower (p < 0.001, in
Species diversity and abundance each case) in comparison to that found in the
A
total of 21 orthopteran species reference site. However, B. luctuosa, O.
belonging to 16 genera were recorded in this abruptus, Camarvonella sp, and Metioche sp.
study. The five orthopteran families present found in the 4 and 6 year old sites were
comprised Acrididae (10 species and 8 genera), completely absent in the reference site. The
Pyrgomorphidae (7 species and 4 genera), abundance of O. fuscovittata and A. psittacina,
Gryllidae (2 species, 2 genera), Eumastacidae was significantly lower (p < 0.001, in each case)
andTettigoniidae (1 species and 1 genus, ineach in all the five (0-8 yearold) mine sites relative to
case) (Table 1). the climax site. Both A. exaltata andL. cruciata
Abundance of the 21 orthopteran species exhibited lower abundance (respective
showed significantvariations at all the five mine significance values being p < 0.01 and p < 0.05)
sites (0-8 year old), relative to the reference site: inthe 8 year old site. Althoughthe abundance of
Acrida conica, Acrida exaltata, Gastrimargus Pyrgomorpha sp. Chrotogonus sp., A.
africanus, Leva cruciata, Pyrgomorpha sp., crenulata, S. savignyi and G. musicus was
Chrotogonus sp., Lepidogryllus sp., progressively higher on moving from 2 - to 8-
Chrotogonns armatus, Spathosternum
year old sites, significantly lower values relative
prasiniferum, Gastrimargus musicus, Oxya to the reference site were obtained only in the
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ShbbirRKhan andNeelkamalRastogi
Table-1: Orthopteran diversity,feeding guilds,habitatspecificityand stagesrecordedinthe0-10yearold
minesitesbyusingthevisualscanningandsweep netting methods (H=HerbivoresandD=Detritivores).
Samplingmethods
Habitat Feeding Visual Scanning SweepNetting
Family Species
specificity guild Adult
^1^vTiInInIlUiUcS
s
Acrididae Acrida conica Generalist H + + + +
Acrididae Acridaexaltata Specialist H + + +
Gastrimargus
Acrididae afiicanus Generalist H - + + +
Spathostemum
Acrididae prasiniferum Specialist H + + - -
Gastrimargus
Acrididae musicus Generalist H + + - +
Acrididae Oxyafiiscovittata Specialist H + + +
Acrididae Catantopspinguis Generalist H + -
Sphingonotus
Acrididae savignyi Generalist H + + - +
Acrididae Bryodema luctuosa Specialist H +
Acrididae Oedaleusabruptus Specialist H + + - +
Pyrgomorphidae Levacruciata Generalist H + + +
Pyrgomorphidae Pyrgomorpha sp. Generalist H + - + +
Pyrgomorphidae Chrotogonus sp. Generalist H + + + +
Chrotogonus
Pyrgomorphidae armatus Generalist H + - + -
Atractomorpha
Pyrgomorphidae psittacina Specialist H + +
Atractomorpha
Pyrgomorphidae crenulata Generalist H + +
Chrotogonus
Pyrgomorphidae trachypterus Generalist H + + +
Gryllidae Lepidogryllus sp. Generalist D + + + +
GryUidae Metiochesp. Specialist D + + + +
Tettigoniidae Pterophylla sp. Specialist H + +
Eumastacidae Camarvonella sp. Specialist H + +
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Recolonisationpatternsoforthopteran speciesin revegetated coal mines
DMRT
case ofthe 2-4yearoldsites. The abundance of diversity. However, the post hoc test
C. pinguis was significantly lower in the 0-2 (p revealed highly significant differences between
< 0.001) and 4 (p < 0.01) year old sites relative alpha diversity found in the 0 and 2 year old site
to the reference site. Acrida conica, G. in comparison to the 6 and 8 year old and the
africanus, Pyrgomorpha sp., Chrotogonus sp.,S. reference, sites. Significant differences in
prasiniferum, G. musicus, Camarvonella sp., diversity were also obtained between the 0, 2
Pterophylla sp., A. crenulata, S. savignyi and B. and 4 year old sites. No significant differences
luctuosa, did not demonstrate any significant were obtained between 6, 8 and 10 year old
differences in their abundance at the 8 year old sites.
dump site in comparison to the climax site. Beta diversity was highest in the 4 and 6
Significant differences were also not obtained in year old sites and lowest in the 2 year old dump
the abundance ofLepidogryllus sp., C. armatus, site, while there was no difference in the species
C. pinguis and C. trachypterus at the 6 and 8 composition ofthe 8 and 10 year old dump sites
year old sites. Metioche sp. and O. abruptus (Fig. 3). While grasshopper abundance and
abundance did not vary significantly at the 0, 2 number ofspecies showed a linearincrease, (the
and 8 year old dump sites in comparison to the curve was steep initially and more gradual in the
climax, 10 year old site. Camarvonella sp. was late successional stages) in the case ofcrickets,
recorded only in 4 and 6 year old sites. Since all both parameters demonstrated a decrease in the
the species were absent in the 0 year old dump later, 8-10 yearold mine sites (Fig. 4). One-way
ANOVA
site, the abundance of each revealed highly of grasshopper abundance showed
significant values (p < 0.001, in each case) with highly significant differences (F5,24=^198.843,
respecttothatforthereference site. p < 0.001) in the 0-10 year old sites, although
While twelve orthopteran species were abundance of crickets demonstrated a lower
found to be generalist species progressively significance value (F 24= 3.186, p < 0.05). The
5.
DMRT
increasing in abundance in the 2-10 year old post hoc test revealed significant
sites, an exception was Lepidogryllus sp. which differences in grasshopper abundance among the
did not show any specific pattern with increase 0-10 year old sites, while the abundance of
inrestoration. crickets showed significant differences between
There were nine habitat specialist 0 and 2-6 year old sites. No significant
orthopteran species. Camarvonella sp., differences were found between 2-10 year old
Metioche sp., Bryodema luctuosa and Oedaleus sites (Fig. 4)
abruptus were found only in 4 - 6 year old sites Rank abundance patterns revealed that
which had intermediate level of disturbance. In lowest orthopteran diversity is found in the 2
contrast, Acrida exaltata, Spathostemum year old site (the uppermost curve). There was
prasiniferum. Oxyafuscovittata, Atractomorpha overlap between the rank abundance curves
psittacina and Pterophylla sp. were restricted to obtained forthe4-10 yearoldsites (Fig. 5).
themaximallyrestored, 8 and 10yearoldsites.
Alpha diversity increased with the age of Discussion
the coal mine sites firom 0 to 10 years (Fig. 2). Our results reveal that grasshoppers are
One-way ANOVA indicates significant early colonisers ofrevegetated mine sites, since
differences in the orthopteran alpha diversity at out ofthe 12 orthopteran species recorded at the
the five (2-10 year old) study sites {F $. 24 = 2 yearold site, 11 species were ofgrasshoppers
344.465, p < 0.001). The 4-10 year old mine and only a solitary species of cricket
sites showed a very gradual increase in alpha (Lepidogryllus sp.) was found. Acrididae and
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ShbbirRKhan andNeelkamalRastogi
Pyrgomorphidae were species rich families species ofcrickets (i.e. decomposers) decreased
dominant at the mine sites. While G. africanus while those of herbivorous grasshoppers
remained consistently abundant in the 2-10 year increased with increase in mine site
old spoils, A. conica abundance increased rehabilitation age. Moreover, while some species
gradually, reaching peak values only in the 10 are habitat specialists, characteristic of specific
year old, climax, successional stage (Fig. 1). successional stages (either with low or high
Three species of grasshoppers, Camarvonella disturbance levels), others are generalists which
sp., B. luctuosa and O. abruptus preferred gradually increase in abundance along the
habitat conditions prevalent in 4 and 6 year old descendingdisturbance gradient.
dump sites and were totally absent in the late The abundance and diversity of crickets
successional stages. Some grasshopper species (decomposer feeding guild, GuUan and
appear to be highly sensitive to environmental Cranston, 2000) showeda steep increase initially
disturbances caused due to mining activities. but decreased in the 8-10 year old mine sites.
Thus, A. exaltata, O. fuscovittata and While Metioche sp. showed an increasing
Pterophylla sp. appear to be late- colonising abundance trend in the 4 to 6 year oldmine sites
grasshopper species, arriving much later i.e. in and was absent in later stages, Lepidogrylltis sp.
the almost restored, 8 year old mine site, did not demonstrate any abundance pattern in
onwards. Ourresults thus support earlierstudies, the 2-10 year old sites. Thus, decomposers are
carried out in rangelands, which show that found to be more abundant in the early
grasshopper species assemblages are highly successional stages and gradually decrease and
^ sensitive to the level of disturbance (Kemp et give way to the herbivores in the late
al., 1990). successional 8 year old site and the climax, 10
Orthopterans were totally absent in the year old mine site. Earlier studies on newly
newly established mine sites. However, alpha abandoned arable land, meadows with large
diversity of orthopterans progressively but mammal grazing pressure and protected semi-
gradually increased (although not significantly) natural steppe meadow, report orthopteran
in the 4-10 year old sites. Beta diversity assemblages to be more stable on the
exhibited a gradual increase, reaching highest undisturbed and intermediate disturbance level
values in the 4-6 year old sites, there being no sites (Baldi and Kisbenedek, 1997). In the
difference in orthopteran diversitybetween the 8 complete absence of significant grazing by
and 10 year old sites. Alpha diversity and rank mammalian herbivores at the mine sites, the
abundance patterns indicate that orthopteran herbivorous grasshoppers and decomposer
diversity is low in the highly disturbed site (2 crickets provide an important ecological
year old) but after increasing sharply in 4 year pathwayfor energyflow andnutrient cycling.
old site, at intermediate levels of disturbance Being a major source offood forpredatory
does not increase significantly in the more arthropods (Schmitz, 2008), orthopteran species
disturbed, 6-10 year old sites. The lackofa clear may influence other organisms during the re-
pattern inrank abundance curves obtained for4- colonisation process. They are also important
10 year old sites may be due to the fact that components of the food chain for many
while the alpha diversity of orthopterans does insectivorous birds and mammals (Capinera et
not differ significantly between sites with al, 1997; Mayya et al, 2005), Alteration of
intermediate and low levels of disturbance, orthopteran population dynamics is expected to
abundance levels are significantly higher in the affect several trophic levels in the food chain
less disturbed sites. Secondly, the number of (Capinera a/., 1997).
6
Recolonisationpatternsoforthopteran speciesin revegetated coalmines
ShbbirRKhan andNeelkamalRastogi
Fig.l: Abundance of orthopteran species belonging to different families: (a) Aciididae, (b) Pyrgomorphidae, (c)
Gryllidae,Tettigoniidae andEumastacidae, in coal minespoUsestablishedin 2011 (0yearold),2009(2yearold),2007(4
year old), 2005, (6 year old), 2003 (8 year old) and 2001 (10 year old, reference site). One-way ANOVA followed by
Dunnett'sposthoctest: *p<0.05,**p<0.01 and*** p<0.001,ns-Notsignificant.
45 1
Rehabilitationage(yrs.)
Fig.2:Alphadiversityoforthopteranspeciesin0to10yearoldcoalminesites(establishedin2011,2009,2007,2005,2003
and 2001, respectively). One way ANOVA followed by Duncan's Multiple Range Test (DMRT) (p < 0.05) were used to
compare the calculated orthopteran alpha diversity at different coal mine sites. Different letters denote significant
differences(p<0.05)amongorthopteranalphadiversityatdifferentsites(0-10yearold).
8
Recolonisationpatternsoforthopteran speciesin revegetatedcoalmines
1.4 n
Rehabilitationage(yrs.)
FigJ: Beta diversity of orthopteran species in 2-8 year old coal mine sites (established in 2009, 2007, 2005 and 2003,
respectively).
Rehabilitationage(yrs.)
Fig.4: The diversity and abundance ofgrasshoppers and cricket species in 0-10 year old coal mine sites (established in
2011,2009,2007,2005,2003 and2001,respectively).
=Abundanceofgrasshoppers, "=Abundanceofcricliets,
--
=Diversityofgrasshoppers, =Diversityofcrickets
9
ShbbirRKhan andNeelkamalRastogi
0 2 4 6 8 10 12 14 16 18
Fig.5: Speciesrankabundanceplotoforthopterancommunityin2to10yearoldcoalminesites(establishedin2009,
2007,2005,2003 and2001,respectively).
The results of the present study clearly We thank the reviewer for his
demonstrate a strong relationship between constructive suggestions. Funding support from
anthropogenic disturbances due to mining the Ministry of Environment and Forests, New
We
activities and orthopteran assemblage Delhi, is gratefully acknowledged. thank
compositions in mine sites of different ages. Northern Coalfields Ltd. for providing
Grasshopper diversity and abundance varied permission for conducting the field study at the
linearly with degraded ecosystem rehabilitation, coal mine site of NCL, Singrauli, in Madhya
We
suggesting the important role ofgrasshoppers in Pradesh, India. are very grateful to Dr. K.
food webs operating in the mine sites. Since Usmani, Department of Zoology of Aligarh
grasshopper assemblages are documented to Muslim University for his help in the
show stronger differentiation among disturbed identification of specimens and to Dr. Ram
sites not evident even on the basis of floristic Sagar ofDepartment ofBotany, Banaras Hindu
data (Andersen et al, 2001), the abundance University, for his helpful suggestions related to
patterns of grasshoppers, particularly those of diversityanalysis.
habitat specialists, may be ofsignificance in the
evaluation of restoration progress in degraded References
ecosystems. Andersen, A. N., Ludwig, J.A,, Lowe, L. M. and
Rentz, D.C.F. 2001, Grasshopper
Acknowledgements biodiversity and bioindicators in Australian
tropical savannas: Responses to disturbance
10