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RA
U
RICHNESS
AND
COMPOSITIONAL
RESPONSES
OF
ANTS
TO
LAND
USE
CHANGE
Thesis submitted
for
the award
of
MASTER'S
DEGREE
in
WILDLIFE
SCIENCE
by
MOHAMMAD
ABDUS
SHAKUR
to
Saurashtra
University
Rajkot-360005
Under the Guidance of
Dr.
Gautam Talukdar
Dr.
Navendu
Page
Dr.
M.
Jadegowda
Wildlife Institute of India
August, 2021
RICHNESS AND COMPOSITIONAL RESPONSES
OF ANTS TO LAND USE CHANGE
Thesis submitted for the award of
MASTER’S DEGREE
in
WILDLIFE SCIENCE
by
MOHAMMAD ABDUS SHAKUR
to
Saurashtra University
Rajkot- 360005
Under the guidance of
Dr. Gautam Talukdar
Dr. Navendu Page
Dr. M. Jadegowda
August, 2021
ii
Wildlife
Institute
of
India
DECLARATION
I,
Mohammad
Abdus
Shakur,
hercby declarc that
the
rescarch
work
entitled
"Richness
and
compositional
responses
of
ants
to
land
use
change",
carricd
out
in
partial
fulfilment
of
M.Sc.
(Wildlife
Science)
degree
of
Saurashtra
University,
Rajkot
is
an
original
piece
of
research
work.
This
research
work
was
caried
out
under
the
supervision
of
Dr.
Gautam
Talukdar,
Dr.
Navendu
Page
and
Dr.
Jadegowda
at
the
Wildlife
Institute
of
India
from
January
2021
to
July
2021.
I
hercby declare that this work has not been submitted for any other degrec
of
any
university.
Mr.
Mohammad
Abdus
Shakur
(XVII M.Sc. Wildlife Sciencc
Date: 13h August,
2021
Place:
Dehradun
t .
18,
Tadt,
àrarF-248
001,
3FTE,
HTZa
Post
Box
No.
18,
Chandrabani,
Dehradun
-
248
001,
Utarakhand,
INDIA
di.vE:
+91-135-2640100,
2640114,
2640115,
:0135-2640117
EPABX:+91-135-2640100,
2640114,
2640115,
a:
0135-2640117
-H/E-mail
:
wii@wii.gov.in,
4/Website:
www.wi.gov.in
lP
Wildlife
Institute
of
India
CERTIFICATE
This
is
to certify that
Mr.
Mohammad
Abdus
Shakur,
has carried out an
original picce of rescarch
in
partial fulfilment
of
Master's Degree
in
Wildlife Scicnce
of
the
Saurashtra
University, Rajkot, Gujarat.
The
topic
of
her
dissertationis
"Richness
and
compositional
responses
of
ants
to
land
use
change".
The
study
was
camied
out
under
our
supervision from
January
2021
to
July 2021.
We
hereby
certify that
this
work
has
not
been
submitted
for
any
degree
to
any
university.
aukala
Ir.
Gautam
Talukdar
Dr.
Navendu Page]
Dr.
M.
Jadeyegowda
Scientist
E Scientist C Associate Professor
Supervisor
Co-Supervisor
Co-
Supervisor
Date:
13th
August,
2021
Place: Dehradun
Post
Box
No.18.
Chandrabani, Dehradun
248001.
India
EPABX
91-135
640111
to
640115
640990
FAX:
640117
GRAM: WiLDLIFE
E-Mail:
wi@w.gov
in
Wildlife
Institute
of
India
CERTIFICATE
OF
PLAGIARISM
CHECK
It
is
certified that the M.Sc thesis titled
"Richness
and
compositional
responses
of
ants
to
land
use
change"
submitted
by
Mr.
Mohammad Abdus
Shakur
has bcen
Cxamined
by
us
for plagiarism check
as
per UGC (Promotion
of
Academic Integrity and
Prevention of Plagiarism
in
ligher liducational Institutions) Rcgulations.
The
following
inferences
arc
drawn
from
this
check:
a)
Thesis has significant new work/knowledgc as comparcd
to
alrcady publishcd work
or
work
under
considcration
for
publication
clsewherc.
b)
No
sentence, cquation, diagram, table, paragraph or section
is
found
to
havc been
copicd
verbatim
from
previous
work
unless
it
was
placcd
under
quotation
marks
and
the source
was
duly citcd.
c)
The work presented
is
original work
of
the author (i.c. there
is
no plagiarism) and
there
is
no
fabrication
of
data
or
result
by
manipulating rescarch material, cquipment
or
processors
or
by
changing
or
by
omitting
data
or
results
such
that
the
research
is
not
accuratcly reprcscnted.
The
similarity
indices for the
individual
chapters
as
reported
by
the softwarc
iThenticatc
are
as follows:
Chapter
iThenticatc"
similarity
index
S.No
1/%
.
Introduction
9/0
Study
arca
5%
Methodology
1%
Results
4.
2/%
Discussions
0%
Conclusions
and
Implications
.
5%
Overall
lahder
Dr.
autam
Talukdar
Smt. Sunita
Agarwal
Supervisor
Librarian
geTbictqEZTeT
Librarian
Idlife Instit
at
Ho
18.
a1,
ENTEI-
248
001,
TYTEus,
ATVA
Post
Box
No.
18,
Chandrabani,
Dehradun
248
001,
Utarakhand,
INDIA
.g.a.grm:
491-135-2640114,
2640115,
2646100
0135-2640117
EPABX:
+91-135-2640114,
2640115,
2646100
Fax:0135-2640117
/
Email:
wii@wil.gov.in
àa/Website:
www.wi.gov.in
v
Dedicated to my Parents
&
All the Queens that were generous enough to donate
their daughters for this study
vi
Painting by: Poonam pal
Species in the painting from top to bottom -
1. Oecophylla smaragdina, (Asian Weaver Ant) weaving its nest
2. Crematogaster sp. feeding on a caterpillar
3. Strumigenys sp. foraging amidst the leaf litter
vii
ACKNOWLEDGEMENTS
I would like to thank me for believing in me and motivating myself through-
out the course of study.
I’m thankful to the Karnataka Forest department for the permissions to work
and collect specimens for the study. I would like to thank Prabhakaran sir, IFoS for
the permissions and the trip to Pushpagiri Wildlife Sanctuary which made me familiar
with the landscape.
I’m grateful to my supervisors Dr. Gautam Talukdar and Dr. Navendu Page
for guiding me through this period. I would like to thank Gautam sir for motivating
me and the support and suggestions throughout the study. I thank Navendu sir for his
constant support, technical inputs and all the help with getting me in touch with the
right people during the field and in IISc which made my work flow a lot easier during
the course of study. It will be un justful if I don’t thank Navendu sir for introducing
me to plant identification.
I would like to extend thanks to the administration of College of Forestry,
Ponnampet for offering us facilities during the initial course of the field which was
very helpful. I’m thankful to Dr. Kushalappa for help in selection of sites for the study
and the stay. I’m grateful to Dr. Jadegowda and Dr. Satish from Forestry college for
their help and suggestions related to site selection and obtaining permissions during
the field work. I thank Arjun, Ashik, Niaz, Syed Ali from Forestry college for their
help during my stay at the college and Vinay for accompanying and helping me in the
field. I am thankful to Manju anna, forest guard for accompanying me and helping
during the field work. It’s worth remembering the field with Ashish and his friend
Vinay and Andrew. The trip to Kukke in the rain at night was worth the introduction
to a landscape like Western Ghats.
I thank Dr. Sumanta Bagchi for allowing me to use the facilities in Eco systems
lab at IISc, Bangalore. I’m indebted to Pronoy Baidya for introducing me to the art of
ant taxonomy and helping me throughout the course of the study. I thank the people
of the Eco systems lab for helping me during the lab work.
viii
I’m lucky to have friends like Saanjbati, Jithin, Satwik, Bhavya, Unnati,
Lavanya who are more like a family. I thank Hashim bhai for his guidance and help
whenever I’m in Bangalore.
I would like to thank all the faculty for aiding me to increase my understanding
and appreciation of wildlife. I thank Dr. Suresh Kumar for the bird watching and mist
netting sessions and Dr. Bindu Raghavan for the treats during the first lockdown. I
thank Negi ji for all the things he did for us both formally and informally.
I thank my MSc batchmates for their role in improving me as a person and for
all the discussions that increased my understanding of science and wildlife. I will miss
eating biryani from Andhra mess along with Yukti, Patel and Chakma. I thank
Anubhuti, Anjitha, Gitima, Divya, Jithin, Lovy bhai and Patel for all the snacks I used
to eat when they get stuff from home. The tours wouldn’t have been memorable if not
for the whole pack staying together.
I thank Kalzang bhai, Mohit bhai, Sohom bhai and Vishnu who are always
ready to play football. I thank Bhawana di and Pankaj bhai for motivating me and
supporting me throughout my stay in the hostel. I thank Poonam di for the painting of
ants and for all the fish we ate. I thank all the hostel inmates for tolerating me in the
mess when the power goes off during dinners.
I am indebted to my family for giving me the freedom to pursue my interests.
I thank Dr. Y. Kalyan Kumar and Dr. Michael David for their constant support and
motivating me to go after my dreams.
Lastly, I would like to thank those firefighters who doused the fire when the
train I was travelling caught fire.
I thank Allah for the grateful time I had during the course and throughout the
field.
********
ix
Table of Contents
DECLARATION ........................................................................................................ ii
CERTIFICATE ........................................................................................................ iii
CERTIFICATE OF PLAGIARISM CHECK ........................................................ iv
ACKNOWLEDGEMENTS ..................................................................................... vii
LIST OF FIGURES ................................................................................................... x
LIST OF TABLES ................................................................................................... xii
SUMMARY ................................................................................................................ 1
INTRODUCTION ...................................................................................................... 1
AIM ............................................................................................................................. 4
STUDY AREA ............................................................................................................ 5
Continuous and undisturbed forests ..................................................................... 7
Forest fragments/ Sacred grove ............................................................................ 8
Coffee Plantations ................................................................................................. 10
METHODOLOGY ................................................................................................... 12
Structural attributes ............................................................................................. 12
Ant sampling ......................................................................................................... 12
Statistical methods ................................................................................................ 15
RESULTS ................................................................................................................. 17
Overview ................................................................................................................ 17
Structural complexity ........................................................................................... 17
Species Composition ............................................................................................. 22
DISCUSSION ........................................................................................................... 28
Structural complexity ........................................................................................... 28
Species richness ..................................................................................................... 28
Beta diversity ........................................................................................................ 29
Insecticides and Human influence ...................................................................... 30
Species Composition ............................................................................................. 31
Limitations ............................................................................................................ 32
CONCLUSION ......................................................................................................... 33
REFERENCES ......................................................................................................... 34
Appendix 1 ................................................................................................................ 41
Appendix- II- ............................................................................................................ 45
x
LIST OF FIGURES
Figure 1: Map of the study area ............................................................................................... 6
Figure 2: The different land use types considered in this study, each different from the other.
(Taken in: Feb-2021) ................................................................................................................ 7
Figure 3:: A) A Devarakadu in a matrix of Paddy. B) A fragment that is encroached for
resources (here red soil for construction). (Taken on: 16-Mar-2021) ..................................... 8
Figure 4:The path leading to deity. A) A narrow and gravelled road preserving natural
environment. B) A wide metalled road at the cost of the native trees. (Taken on: 16-Mar-
2021) ........................................................................................................................................ 9
Figure 5: Sampling method of ants at a site. ......................................................................... 13
Figure 6: Structural complexity of land use with respect to various habitat characters. A)
Canopy closure & B) Litter weight decreased from Forest to coffee plantations, while C) Tree
basal area was highest in fragments and least in coffee plantations D) Height of vegetation
decreased from forests to coffee plantations and was least in fragments ........................... 18
Figure 7: Diversity in Girth and height classes of vegetation didn’t show any trend ............ 18
Figure 8: Observed Species richness (Mean & SE) and the line is just to denote the trend. (A-
D) The trend observed at sample level, transects vanishes at sites and reverses at land use,
D) Rarified and Estimated species (Chao I) richness across land use, (E- F) The contribution of
beta diversity increased from forests and fragments to coffee plantations at both transect
and sites ................................................................................................................................. 20
xi
Figure 9: NMDS plot of various land use types. The vectors represented here shows the
direction of their increase in the plot and thus the contribution to the sites (vectors represented
are those with p ≤ 0.02) .......................................................................................................... 23
Figure 10: Rank abundance curves across land use types (The top five most abundant species
are labelled). Although some species are dominant across other land use their dominance
order changed. There is an increase of rare species in coffee plantations. ......................... 25
Figure 11: The contribution of replacement and nestedness to the total beta diversity for each
land use. There is an increase in contribution by replacement from forest to coffee plantations.
............................................................................................................................................... 27
Figure 12: Sample based accumulation curves for each land use. None of the land use
reached saturation. ................................................................................................................ 28
Figure 13: A Venn diagram of species shared between each land use (Total species = 70) . 32
xii
LIST OF TABLES
Table 1: Description of the study sites in Kodagu district, Karnataka, India. ....................... 14
Table 2: Intercept only model. The estimates on intercept are significant as they are not
overlapping zero. .................................................................................................................... 21
Table 3: Although the best model explained little variation (marginal R2 = 0.065), it was
within 2 delta AIC units from the intercept only model (Δ AIC=1.5) .................................. 21
Table 4: Table of results of Beta diversity of individual land use and for the landscape ...... 26
Table 5: Species by site matrix .............................................................................................. 41
1
SUMMARY
Human-made changes to the ecosystem are leading to the loss of biodiversity
across the world. Western Ghats (WG) known for its rich biodiversity has historically
undergone fragmentation and loss of its primary forest cover which has significant
negative effects on biodiversity. Studies have shown that fragmentation almost
invariably leads to a decrease in species richness, a pattern consistently observed
across different taxonomic groups. Ants, one of the numerically dominant groups in
forest ecosystems, play a vital role in many ecosystem processes and yet is one of the
least studies groups within the WG landscape. The effect of land use change on ants
in forests, fragments, native shaded coffee plantations and silver oak shaded coffee
plantations was studied in the Kodagu district of Karnataka, India. We hypothesized
that habitat heterogeneity would decrease from forests to fragments to native shaded
coffee plantations and silver oak shaded coffee plantations and this would
consequently lead to a decline in the richness and change in species composition of
ants along this gradient. To test this prediction, ants were sampled along transects in
the four-land use types using both pitfall traps and modified wrinkler extractors. As
expected, there is a reduction in structural complexity across forest, fragments, native
shade coffee and silver shade coffee plantations. We found that at the level of a
transect, species richness reduced from forests to coffee plantations. Interestingly, the
trend reversed when viewed at the scale of the land use, where all the forest sites
together were the least species rich while the coffee plantations collectively showed
the highest species richness. With respect to species composition, we found that coffee
plantations had species adapted to drier environments whereas species inhabiting leaf
litter were primarily found in forests and fragments. Coffee plantations with human
residence inside had higher species compared to those not inhabiting. This study
highlights the importance of scale for assessing the biodiversity potential of human
modified habitats and for assessing their conservation potential.
Keywords- Formicidae, Western Ghats, Coffee plantations, Beta diversity, Land use
change
INTRODUCTION
Human-induced changes in the ecosystems in the form of habitat
fragmentation, habitat conversion, and degradation are causing loss of biodiversity
both locally and globally (Newbold et al., 2015). Tropical forests, although harbouring
a large chunk of biodiversity, are at the receiving end due to the human induced
changes and need prioritized conservation (Laurance and Bierregaard, 1997; Malhi et
al., 2014). Studies on the effects of fragmentation and habitat alterations have used a
diverse set of groups ranging from mammals, birds, herpetofauna, and invertebrates
(Almeida et al., 2016; Kapoor, 2008; Kurz et al., 2014; Ramesh and Downs, 2015).
Some of the studies have found that habitat fragmentation and habitat alterations pose
a serious threat not only to species but also to the key ecological processes like
predation, decomposition, pollination, and parasitism (Baur and Erhardt, 1995;
Kareiva, 1987; Klein, 1989; Koh, 2007; Kruess and Tscharntke, 2000; Powell and
Powell, 1987; Riemann et al., 2017). Ants have been used as models to study the
effects of fragmentation and land use changes (Andersen et al., 2002; Armbrecht and
Perfecto, 2003; Brühl et al., 2003; JingWen et al., 2017; Leal et al., 2012; Tawatao et
al., 2014). They are regarded as one of the best groups to study the response to land
use change due to their functional and ecological dominance in the environments they
live in and due to the range of functional roles they carry out as indirect herbivores,
scavengers, and predators (Fittkau and Klinge, 1973; Wilson, 1990; Wilson and
Hölldobler, 2005).
The conversion of land from its natural state has profound effect on various
abiotic and biotic components, which are crucial in many ecosystem processes
(Brambilla et al., 2010; Hadi et al., 2000). Interacting with other global changes,
modifications in land use have a strong impact on the structure of biological
communities factors (Arinaminpathy et al., 2009; Berg et al., 2015; Sala et al., 2000).
The majority of the studies on land use changes have focused on understanding the
effect of forest conversions into plantations within agroforestry systems.
3
These studies consistently report a negative impact on the species richness and
alternation of community structure (Bickel and Suparoek, 2005; Brühl and Eltz, 2010;
Fayle et al., 2010; Liu et al., 2016; Mauda et al., 2018; Nazarreta et al., 2020; Pacheco
et al., 2009; Ratsirarson et al., 2002). Although agroforestry reduces the biodiversity
of an area, traditional agroforestry systems like shade-grown coffee, which retains
much of the native tree cover act as a refuge for biodiversity when compared to the
altered matrix (Donald, 2004; Perfecto et al., 1996). However, intensification of
management practices involving the replacement of native shade trees, with fast-
growing exotic trees can significantly exacerbate the effect of fragmentation on ant
communities (Armbrecht et al., 2005; Armbrecht and Perfecto, 2003; Asfiya et al.,
2015; Perfecto et al., 1997; Philpott, 2005; Roth et al., 1994).
Studies have shown that fragmentation results in the alternation of ant
community structure, disproportionate loss of specialist species, the proliferation of
generalist and tramp species (species that have been spread around the world
inadvertently by human commerce), and an overall reduction in the ant species
richness by up to 50% (Brühl et al., 2003; Bruna et al., 2005; Gibb and Hochuli, 2002;
Leal et al., 2012; Schoerder et al., 2004). Studies have also documented a reduction
in the abundance of species that manage to persist post fragmentation (Leal et al.,
2012; Tawatao et al., 2014). The decrease in the size of fragments has also been found
to reduce the diversity of ants (Schoerder et al., 2004). Primary drivers of change in
species richness of ant communities post fragmentation have been attributed to the
alteration in the habitat characteristics like reduction in tree cover, tree density which
possibly leads to altered microhabitats (for example- less leaf litter can be found, if
the tree density is less thus affecting the leaf litter dwellers), increase in disturbances
and prominent edge effects (Barrera et al., 2015; Carvalho and Vasconcelos, 1999;
Debuse et al., 2007; Tawatao et al., 2014; Vasconcelos et al., 2006; Watt et al., 2002).
In a recent study by González et al. it was found that there is an increased diversity of
ants near the forest edges than the interior in fragmented forests which they attributed
to the influence of species from the matrix (González et al., 2018).
In the Indian context, very few studies have investigated the effect of
fragmentation on ants, although the findings from these are largely consistent with
4
studies from other parts of the world; they have considered largely the taxonomic
diversity (Badrinarayanan, 2001; Gadagkar et al., 1993; Mone et al., 2014; Mujeeb
Rahman et al., 2012; Rajan and Marathe, n.d.; Vineesh et al., 2007). The Western
Ghats in India is recognized as one of the global biodiversity hotspots due to high
levels of diversity, endemism, and loss of primary forest cover (Myers et al., 2000).
The landscapes in the Western ghats of India represent some of the oldest human-
modified tropical forests and thus provide an opportunity to understand the responses
of biodiversity to long-term habitat change (Anand et al., 2010). The understanding of
ant communities in a highly fragmented landscape is lacking and a better
understanding will give an important perspective of the group that is numerically
dominant in these landscapes.
AIM
Objectives: To determine the effect of land use change on ant communities
Hypothesis:
As studies have shown that there is a decrease in species richness with a reduction in
tree diversity and abundance. We hypothesized that a reduction in structural
complexity of the vegetation from contiguous forests to forest fragments to native
shade coffee to exotic shade coffee will lead to a reduction in species richness, change
in species composition.
Research Questions:
1. How does ant species richness vary across different land use types?
2. How does the community composition vary across different land use types?
5
STUDY AREA
The Kodagu district of Karnataka, known widely as Coorg, is a land of mist and
greenery, carved by the diverging roads and never-ending list of hill stations. It is one
of the sparsely populated districts of the Karnataka state and places are well connected
by road (“Kodagu District, Government of Karnataka,” 2021). The district lying
amidst the Western ghats mountain chain is rich in biodiversity and exhibits a high
percentage of endemic flora and fauna. The district has an elevation ranging from 80
m at the Makutta section to ~1750 m on the top of the Tadiandamol peak (the highest
peak in the district). Most of the district lies in the elevation range of 800- 1100 m
creating a landscape that is dotted with hills all along. Major towns namely, Madikeri,
Virajpet, are located on top of these hills and are less populated as there are many
small villages spread out around. The hills, if not populated, are dotted with trees
intended for agroforestry, dominated by coffee and other plantation crops. The valley
floors are used as paddy fields and appear as wastelands during the non-growing
season as they remain waterlogged and other grasses and shrubs grow. These
agroforestry coffee plantations make Kodagu the topmost coffee-producing district in
the country (Coffee Board of India, 2021). The district receives an average of 2000 –
5000 mm rainfall in the central and western parts and central-eastern parts receiving
about 1500- 2000 mm annually and an average temperature of 24°C and increasing up
to 30°C in the hotter months. It received an average rainfall of 2664 mm between the
years 2014-18 (Indian Meteorological Department, 2021). The study area experiences
a dry season of about 4-5 months from December and April characterized by low
moisture levels (Bhagwat, 2002).
The study was carried out during the late winter (mid-January to the first week of
March) and was confined to the Virajpet taluk of the district. Wet-evergreen forest is
the most dominant vegetation type within this district. Although the majority of the
time it was dry, it rained heavily during the mid-January and late-February period for
at least two days. During late February, the coffee plantations were immersed in fog
during the early hours of the day which cleared out by 9 AM.
6
The major land-use types of this area include undisturbed forests, forest fragments,
coffee plantations, and paddy fields.
Figure 1: Map of the study area.
7
Continuous and undisturbed forests
The Kodagu district is covered by evergreen forests along the Western Ghats
mountain chain on the western and north-western parts while the deciduous forest is
mostly found along the eastern parts of the district. Much of the continuous and
undisturbed forests exist either in the form of protected areas (Wildlife sanctuaries or
National parks) or in the form of reserve forests. The forests in the Western Ghats
section are protected as Brahmagiri Wildlife Sanctuary (WLS), Talacauvery WLS,
Pushpagiri WLS, and reserve forests. All the sampling sites including the forest
fragments, coffee plantations, and the contiguous forests fall within the wet-evergreen
forest type.
Figure 2: The different land use types considered in this study, each different from the other. (Taken in: Feb-2021)
8
Forest fragments/ Sacred grove
A large number of forest fragments lie scattered in the form of sacred groves.
Being primarily forest lands, these are owned by the forest department and are thus
legally protected but are managed by the local communities as common property.
Locally called “Devarakad/ Devarakadu”, each fragment has a deity of its own that is
worshipped and maintained by the locals and a small water body for the associated
activities. Each village has at least one devarakadu associated with it and the overall
density of sacred groves in Kodagu is one in every 300 ha (Bhagwat et al., 2005a;
Kushalappa and Bhagwat, 2001). These sacred groves often range from less than a
hectare to a few ten hectares in area (Bhagwat et al., 2005b). There are around 121
different deities each with its unique forms of worship practiced by 18 different local
communities. Bhadrakali, Bhagavathi, Naga, Aiyappa, Eshwara, and Ajappa are some
of the common deities worshipped in Devarakadu and each deity has an associated
folk story, which connects them closely to the villagers. These deities are considered
as protectors and guardians of villages they are huddled in. All the local villagers
gather annually/ biannually in the Devarakadu to celebrate the ‘Devarakadu Habba’
or the sacred grove festival (Kushalappa, 2014). Apart from these festival gatherings
only the priest visits daily for the puja, or few devotees visit during the off season.
A
B
Figure 3: A) A Devarakadu in a matrix of Paddy. B) A fragment that is encroached for resources (here red soil
for construction). (Taken on: 16-Mar-2021)
9
Each of these Devarakadus differs in their deity and also in the way they are
maintained. Some of them have just a deity and a small area roughly of 10 m2 around
it for the devotees and a central large tree in it and a well-used trail often overgrown
with weeds lead to the deity.
Few of them are further upgraded to a small temple for the deity and a gravel
path leading towards it. While those having gotten sponsors have a metalled road
leading to the temple area with a small temple for the deity and few more buildings
for devotees to sit during the festival. These fragments are threatened by humans either
directly due to encroachment of the area from outside or inside for ever-growing coffee
plantations, or for excavating soil that can earn huge profits and indirectly by the
growth of weeds alongside the edges (both path and the periphery). These fragments
deal with huge pressures from human activities daily. There is an increasing trend of
people moving out of the villages and this has partly resulted in erosion of the faith
and conservation values associated with this informal system of protection. As these
forest patches get their protection mainly due to the underlying socio-religious beliefs,
when the beliefs are no longer there, the existence is threatened.
A
B
Figure 4: The path leading to deity. A) A narrow and gravelled road preserving natural environment. B) A wide metalled
road at the cost of the native trees. (Taken on: 16-Mar-2021)
10
Coffee Plantations
Coffee plantations are one of the prominent land use categories occupying up
to 60% of the area in the district (Bhagwat et al., 2005b). Coffea robusta is the
predominant type of coffee grown in Kodagu which is also the top contributor of this
variety in the country (Coffee Board of India, 2021). Much of the coffee is shade-
grown, as coffee is a shade-demanding species, for which most of the native tree cover
is retained to provide shade for the coffee. Management practices like removal of the
grasses and leaf litter during the growing season (April to November) make the coffee
the only layer of understorey vegetation. The shade-giving trees are also pruned or
felled to provide the optimum shade for the coffee below, which varies across the
plantations.
Sometimes a few selected native trees are grown for their benefits like the
Artocarpus heterophyllus (Jackfruit) whose canopy provides shade to coffee and
fruits. While the usage of pesticides in the plantations hasn’t been encountered, some
of the plantations use insecticides like ‘Gammaxene’ and ‘Cypermethrin’ (0.25%
DP) and the latter being used predominantly during the harvesting period. These are
generally dusted in small amounts on the coffee fruits where ants are patrolling
heavily, which kills the ants and the coffee fruits can be harvested.
Species of ants that are predominantly targeted are Oecophylla smaragdina
(Asian weaver ant) and Crematogaster sp. which are arboreal and defend their nests
and nesting trees aggressively and are frequently encountered nesting on the coffee
plants. Sometimes other species which are not that aggressive also get killed when
they move along the coffee plant. Very few plantation owners go to the extreme of
dusting the whole coffee trees and the whole ground below coffee (after the harvest)
giving it a makeover that essentially removes the majority of other invertebrates less
alone ants. Very few coffee plantations exist in the landscape that are not managed
and one can see another layer of vegetation growing below the coffee and the canopy
trees not being pruned.
11
The coffee plantations are further classified into two types based on the nature
of shade-giving trees (Nath et al., 2011).
1. Native-Shaded Coffee Plantations (Native shaded coffee)
In these plantations, the shade is provided predominantly by the trees that
are native and common in the region. Their canopies spreading far and wide
are the predominant type of plantations. Some plantations have a small amount
of young silver oak trees.
1. Silver oak/ Exotic-Shaded Coffee Plantations (Silver shaded coffee)
With the added advantage of harvesting, the non-indigenous silver oak has
been predominantly planted in some plantations replacing the native trees
either completely or partially and predominates the canopy. It makes a uniform
canopy layer due to their similar time of planting.
The study sites were selected such that they represent a uniform area (8-11
hectares), similar elevation (class range of 780- 910 m), evergreen vegetation type,
and have similar matrix around them. All of the coffee plantations and fragments have
paddy fields, coffee plantations, and roads surrounding them, even the forest sites had
coffee plantations and roads near to the edge. All of the study sites were near the town
of Virajpet. The coffee plantations with greater than 50 % of trees as silver oak were
classified as silver oak/ silver shaded coffee plantations.
12
METHODOLOGY
The study was conducted across four predominant land use types in the study
area – the continuous forests (forests), remnant forest fragments (fragments/ sacred
groves), coffee plantations with native (Native shade coffee), and exotic shades (Silver
shade coffee). The sites were selected such that they represented a similar size class
and with similar topographic features. Selected sites were sampled for ants using the
below mentioned techniques.
Structural attributes
For computing the structural attributes of each land use, vegetation data (Tree and
shrub species, height, and girth) was collected during this study and also compiled
from an earlier study (Page et al., 2010). The structural attributes were collected along
a 10 X 60 m transect for trees and 5 X 5 m for the shrubs at the beginning and the end
of each transect. The canopy cover was evaluated at each ant pitfall location along the
transects using the GLAMA application in mobile.
Ant sampling
Leaf litter ants were sampled following the ‘Ants of Leaf Litter’ protocol (Agosti
et al., 2000). Four transects were laid randomly in each site, sometimes using the
existing paths to navigate. Transects were laid such that they spread across the site and
maximizing the distance between the two nearest transects. Each Transect was 60 m
in length and leaf litter ants were sampled at every 20 m. At each sampling point, the
ants were sampled using both pitfall traps and wrinkler extraction (Figure 5).
• Pitfall traps consisted of paper cups filled with detergent (like surf excel) water
kept inside the ground flushed up to the rim. The traps were kept active for a
minimum period of 24 hours and a maximum of 26 hours from deployment.
The catch in the pitfall was preserved in 70% ethanol.
13
• Wrinkler extraction of ants consisted of sampling the leaf litter in 0.5 m2 area.
The leaf litter and the topsoil up to 2 cm were scraped and transferred to a sieve
with a mesh size of 1 inch and sieved. The siftate was collected in a white tray
and the ants were collected using an aspirator. The catch was preserved in 70%
ethanol. This method was modified from the original wrinkle extraction, where
wrinkler apparatus is used and the leaflitter dwelling organisms move
downwards and are collected in alcohol.
• For quantifying the amount of leaflitter, litter was weighed from the 0.5 m2
plots after sifting the litter for invertebrates.
Figure 5: Sampling method of ants at a site.
14
Table 1: Description of the study sites in Kodagu district, Karnataka, India.
Land use
Name
Location
Area (ha)
Elevation
No of transects
Latitude
Longitude
Forest
Heggala RF
12° 7'49.20"
75°45'56.99"
>1000
910
4
Forest
Kokka RF
12° 5'26.34"
75°50'19.10"
>1000
800
4
Fragment
Arji
12°10'53.25"
75°47'49.13"
8.51
900
4
Fragment
Bettoli
12°11'24.55"
75°47'17.98"
9.72
914
4
Fragment
Kadanoor
12°13'6.23"
75°46'51.70"
9.07
908
4
Native shaded coffee plantation
Bittangala
12° 9'52.66"
75°50'44.09"
8
852
4
Native shaded coffee plantation
Chickpet
12°12'27.94"
75°47'28.82"
10
908
4
Native shaded coffee plantation
Mythadi
12°15'20.51"
75°47'12.20"
10
898
4
Native shaded coffee plantation
Panjarpet
12°10'52.80"
75°48'55.07"
8
903
4
Silver shaded coffee plantation
Bhetri
12°16'10.72"
75°45'24.91"
8.2
905
4
Silver shaded coffee plantation
Kotekoppal
12°13'38.55"
75°48'2.47"
8
893
4
15
Identification of ants
The collected ants were identified till the genus level using the key provided
by Fayle et al. (Fayle et al., 2014). Although the key was developed primarily for ants
of Malaysia, however, most of those tropical genera are represented in the Western
Ghats it was used in the study. Multiple morphospecies in a single genus were
considered as different species under the genus and given a species number in the
order of identification. Only the worker caste was identified and considered in this
study. In case a species had major and minor workers; the minor worker was
considered for the identification. The descriptive images for species collected were
drawn for reference (Appendix II).
Statistical methods
Statistical analysis was performed in R and Excel, and visualized using ggplot2
(R Core Team, 2020; Wickham, 2011). Data collected by both methods (pitfall and
leaflitter extraction) was pooled into a single sample for further analysis. Species
richness was rarefied to the least number of samples across all the sites across land use
types. Species richness estimates for each land use were derived using classic Chao I
estimates for the land use types using EstimateS,v.9.1.0 (Colwell, 2013). The rank
abundance of species across sites was computed in R (BiodiversityR:: rankabuncomp)
and plotted using ggplot2 (Kindt and Kindt, 2015). Mantel test was performed on the
community data with the environmental variables to understand the variation in
communities across sites with the variation in environment (Vegan:: Mantel)
(Oksanen et al., 2013).
Generalized Mixed Effect Model’s (GLMM’s) were performed (glmmTMB::
glmmTMB) to understand the factors explaining the observed species richness
(Magnusson et al., 2017). The predictor variables were scaled before running the
model with a Poisson distribution. Non-metric multidimensional scaling (NMDS) was
performed using bray-curtis dissimilarity with three dimensions at 1000 permutations
16
(Vegan::metaMDS) on transects grouped at land use type to visualize the grouping of
transects across land use. The compositional similarity of communities across land use
was checked using an analysis of similarity test (ANOSIM, Vegan::anosim). Berta
diversity (Whittaker’s Beta- β = γ/α) was calculated at transects and site scale. Beta
diversity was further decomposed into contributions by differences in richness and
turnover [Jaccard index from the Baselga family (adespatial::beta.div.comp)]
(Baselga, 2010; Dray et al., 2018). Local contribution of sites to the overall beta
diversity (adespatial::LCBD.comp) was calculated to identify the sites that are driving
the change in beta diversity across the landscape.
17
RESULTS
Overview
Eleven sites were sampled in total spanning the landscape around Virajpet
comprising 2 sites for the forests, 3 sites for fragments, 4 shades for native shaded
coffee, and 2 sites for the silver shaded coffee. A total of 70 species of ants
representing 31 genera spanning 7 sub-families were collected during the sampling
(Appendix I– Table 1). Mymricinae was the most widely represented subfamily with
29 species from 11 genera. The most species-rich genus was Leptogenys (Ponerinae)
with 7 species and the most abundant species is Nylanderia sp. 1 (Formicinae).
Nylanderia sp. 1 was the most frequently occurring species found in 24.12% of the
340 individual traps (trap level). Leptogenys sp. 1 and Pheidole sp. 2 were the most
frequently occurring species in the transects, and were recorded in 77.27% of the 44
transects sampled.
Structural complexity
We examined multiple attributes of structural heterogeneity to check
whether it declined from forests to fragments to coffee plantations, as initially
hypothesized. We found that the land use types varied with each measure of the habitat
characters assessed. The canopy cover (Figure 6A) and litter weight (Figure 6B)
decreased from forests to silver shade coffee plantations, however between the forests
and fragments canopy cover was comparable. The mean tree basal area was highest in
fragments followed by native shade coffee plantation and then forests and silver shade
coffee. The average height of vegetation was highest in forests followed by coffee
plantations (both types are similar) and least in fragments (Figure 6D).
18
Figure 6: Structural complexity of land use with respect to various habitat characters. A) Canopy closure & B) Litter
weight decreased from Forest to coffee plantations, while C) Tree basal area was highest in fragments and least in
coffee plantations D) Height of vegetation decreased from forests to coffee plantations and was least in fragments.
Figure 7: Diversity in Girth and height classes of vegetation didn’t show any trend.
19
The average species richness observed at the sample and transect levels
complies with the hypothesis that forests are the high species-rich land use type and
the silver shade coffee with the least and fragments and native shade coffee in between
(Figure 8 A&B). The observed species richness was highest in the native shaded coffee
followed by silver shade coffee, while the fragments had lower than coffee plantations,
the forests represented the least number of species, when it’s pooled at land use level
(Figure 8D). Rarefied species richness (to account for the differences in sampling in
each land use) estimated at the least samples across land use types (n=32) indicated
the native shaded coffee was the most species-rich followed by silver shaded coffee.
Rarefied species richness was comparable for forests and fragments and lower than
coffee plantations. Estimated species richness also showed a similar trend of consistent
decline in species richness from coffee plantations to fragments and forests (Figure 8).
20
Figure 8: Observed Species richness (Mean & SE) and the line is just to denote the trend. (A- D) The trend observed at
sample level, transects vanishes at sites and reverses at land use, D) Rarefied and Estimated species (Chao I) richness
across land use, (E- F) The contribution of beta diversity increased from forests and fragments to coffee plantations at
both transect and sites.
21
Factors explaining the species richness
Generalized Linear Mixed Effect Models (GLMM’s) were considered to
incorporate the effect the randomness in transect might have on the species richness
while accounting for habitat variables. The intercept-only model and the global model
were not different from each other either (Δ AIC=2.8). The results thus suggested that
there was no significant contribution by any of the measured habitat variables in
explaining the observed species richness.
Table 2: Intercept only model. The estimates on intercept are significant as they are not overlapping zero.
Table 3: Although the best model explained little variation (marginal R2 = 0.065), it was within 2 delta AIC units
from the intercept only model (Δ AIC=1.5).
Family: Poisson (log)
Formula: species ~ Stem density + human habitation + (1 | transect)
AIC
BIC
logLik
deviance
df.resid
748.2
759.6
-369.5
738.9
172
Random effects:
Groups
Name
Variance
Std.Dev.
transect
(Intercept)
0.03474
0.1864
Conditional model:
Estimate
Std.
Error
zvalue
Pr(>|z|)
(Intercept)
1.27150
0.07012
18.134
<2e-16
***
Stem density
0.12604
0.05398
2.335
0.0195
*
Human habitation
0.24561
0.10702
2.295
0.0217
*
Family: Poisson (log)
Formula: species ~ 1 + (1 | transect)
AIC
BIC
logLik
deviance
df.resid
749.7
756.1
-372.9
745.7
174
Random effects:
Groups
Name
Variance
Std.Dev.
transect
(Intercept)
0.05135
0.2266
Conditional model:
Estimate
Std.
Error
z value
Pr(>|z|)
(Intercept)
1.3817
0.0517
26.73
<2e-16
***
Signif. codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’ 0.1 ‘ ’ 1
22
Effect of residence and insecticide
Built-up or residential buildings were categorized as human habitations and
were observed across fragments and coffee plantations. The species richness per
transect was not significantly different from the native shade coffee plantations (only
native shade had human habitations in them) with and without human habitation (t-
test- P = 0.1759, t = 1.5232, df = 6.3489). We did not find any significant difference
in species richness between plantations which used insecticides and plantations which
did not (Native shade coffee -> t-test- P = 0.740, t = -0.33809, df = 13.898; Silver
shade coffee -> t-test- P = 0.67, t = -0.44721, df = 5.7692). There was no significant
difference in species richness between fragments with human habitations and
fragments without human habitation (paired t-test- P = 0.63, t = -0.49835, df = 9.5036).
Species Composition
While Nylanderia sp. 1 was one of the abundant species across forests,
fragments, and native shaded coffee, Pheidole sp. 1 and Technomyrmex sp. 1 were
abundant across forests, native shaded coffee, and silver shaded coffee. The
dominance order of species changed with the change in land use type (Figure 11). The
number of rare species also increased in coffee plantations compared to forests and
fragments. The Mantel test showed a weak but significant correlation between
environmental dissimilarity matrix and compositional dissimilarity matrix (Mantel
statistic based on Spearman's rank correlation, r = 0.1786; p = 0.0017). This suggests
that change in compositional similarity is associated with change in habitat variables
such as canopy, stem density, leaflitter, pH, tree and shrub species richness.
Only18.6% of the total species were shared across the four land use types, while
forests shared 22.9% with fragments and 27.1% with silver shade coffee plantations.
Native shade coffee plantations shared 32.9% of species with each other, whereas both
23
the native and silver shade coffee plantations shared 37.1% of the species. Coffee
plantations shared 56% of species with forest and fragments together.
The community composition visualized from an NMDS plot suggests that
there is an overlap of communities of coffee plantations with each other and little with
fragments, while forest communities are mostly unique with a little overlap with only
fragments. ANOSIM showed that the composition of ants in each land use together
doesn’t differ much (R: 0.08724). The forested and non-forested land use differed
slightly (R: 0.123).
Figure 9: NMDS plot of various land use types. The vectors represented here shows the direction of their increase in the plot
and thus the contribution to the sites (vectors represented are those with p ≤ 0.02).
24
25
Figure 10: Rank abundance curves across land use types (The top five most abundant species are labelled). Although some species are dominant across other land use their dominance order changed. There
is an increase of rare species in coffee plantations.
26
Beta diversity
Beta diversity of the landscape (across all land uses) is found to be 0.448 when
decomposed further into contribution by the difference in richness and due to species
replacement. Results show that the contribution to overall beta diversity by replacement
(52.6%) was slightly higher than by richness (47.4%) (Table 4Table 5). Native shade coffee
had the highest beta diversity followed by forests and silver shade coffee, whereas fragments
were the ones with the least beta diversity (Table 4). The contribution by transects from the
forest was higher followed by the one in native shade coffee. When beta diversity was
calculated for individual land use types, it showed that there was an increase in contribution
to beta diversity due to the replacement of species in coffee plantations when compared to
forests and fragments (Table 4 & Figure 12).
Table 4: Table of results of Beta diversity of individual land use and for the landscape
Land use
Beta diversity
Replacement
Nestedness/Richness
Forest
0.449
0.357
0.092
Fragment
0.406
0.359
0.047
Native shade coffee
0.456
0.407
0.049
Silver shade coffee
0.424
0.401
0.023
Entire Landscape
0.448
0.235
0.212
27
Figure 11: The contribution of replacement and nestedness to the total beta diversity for each land use. There is
an increase in contribution by replacement from forest to coffee plantations.
28
DISCUSSION
This is one of the few studies to investigate the effect of land use change on
predominantly leaf litter ant communities in the Western Ghats biodiversity hotspot
(Badrinarayanan, 2001; Gadagkar et al., 1993; Rajan and Marathe, n.d.; Vineesh et al.,
2007). The land use types examined in the study represent a gradient in structural
heterogeneity of vegetation which largely declines from forests to fragments to native
shade coffee plantations to and silver oak dominated coffee plantations.
Structural complexity
Our study was based on the premise that structural heterogeneity quantified
through vegetation attributes would decrease from forests to fragments to coffee
plantations. Our results were partially consistent with this prediction (Figure 6).
Canopy closure and leaf litter weight declined from forests to coffee plantations. A
similar trend was observed in a study from a nearby landscape where litter biomass
and litter moisture decreased from forests to coffee plantations (Badrinarayanan,
2001). However, mean basal area of trees increased from forests to fragments and
native shade coffee plantations and was least in silver shade coffee plantations.
Although the vegetation girth and height classes represented a decrease from forests
to coffee plantations, the shannon’s diversity didn’t show any trend, but the fragments
were least diverse because of higher number of shrubs. The management strategies
followed in coffee plantations leads to opening up of canopies for ensuring adequate
light for coffee below. This results in reduction of foliage thus affecting the amount of
litter seen. The leaf litter in the coffee plantations is also subjected to continuous
removal owing to the management strategies thus contributing to the reduction in litter
in plantations as well. Majority of the litter in coffee plantations was contributed by
coffee leaves (personal observation). Certain life forms such as lianas and understorey
trees and shrubs were entirely missing from the coffee plantations.
28
Species richness
Observed species richness at the level of a transect showed a consistent
decline from forests to coffee plantations (Figure 8). The trend is consistent with a
number of other studies that have examined the effect of conversion of forests to other
land use types (Armbrecht et al., 2005; Bickel and Suparoek, 2005; Mauda et al., 2018;
Nazarreta et al., 2020; Perfecto and Snelling, 1995; Philpott et al., 2008). However,
this trend disappeared at the scale of individual sites and reversed at the scale of land
use type when species richness of different sites within a land use type was pooled
together. At this scale, species richness showed a positive trend from forests to coffee
plantations (Figure 8D).
This result was contrary to our expectation that forests with intact structural
heterogeneity would show the highest species richness among the four land-use types.
The results suggest that coffee plantations together support the highest species
richness while the two forests sites together exhibit lowest species richness (Figure 13).
This result was not an artefact of sampling bias as evident from sample-based
rarefaction curves (Figure 12) which shows that the two categories of coffee plantations
Figure 12: Sample based accumulation curves for each land use. None of the land use reached saturation.
29
showed higher rarefied species richness as compared to the forest or the fragments.
This is in contrast with the trend observed in other studies that focused on the effect
of land use conversion at comparable scales. These studies found a consistent decrease
in species richness from forests to the plantations (Fayle et al., 2010; Liu et al., 2016;
Pacheco et al., 2009; Ratsirarson et al., 2002). While the study by Badrinarayanan has
found similar species richness between forests and coffee plantations
(Badrinarayanan, 2001). A similar shift in the pattern of species richness at different
scales (landscape and transect level) was also reported by Betselmeyer & Weins from
Argentina (Bestelmeyer and Wiens, 1996). Thus, the patterns in species richness
observed in our study showed a strong scale dependence.
The increase in species richness with the decrease in habitat complexity has
been observed in two studies from Australia, in Sydney sandstone ridge-top
woodlands (Lassau and Hochuli, 2004) and in urban green spaces (Ossola et al.,
2015). They attributed this pattern to enhanced negotiability on the ground in less
complex areas and greater light availability for fulfilling the energy requirements of
ants as they are thermophilic. In contrast to this, Nooten et al., (2019) found that among
the urban golf courses, those with highly complex habitats were the ones with highest
species richness of ants. Comparison of the present study with other studies should be
carried out with caution as the measure of habitat complexity was not the same across.
That said, none of the habitat variables measured during the study viz., canopy, stem
density, and amount of leaf litter explained the variation in species richness, although
all these varied with land use (Figure 6).
Beta diversity
This reversal and strong scale dependence of species richness observed in
our study can be attributed largely to higher beta diversity or species turnover in coffee
plantations than forest and fragments (Figure 8-E & F). Average beta diversity showed
an increasing trend from forests to plantations with the exception of fragments which
exhibited lowest beta diversity. This suggests that, on an average any two transects in
silver shade coffee plantations are more dissimilar than those in fragments and forests.
30
A similar pattern was found when beta diversity was examined across sites. In other
words, we found maximum turnover in species composition across sites for coffee
plantations as compared to forests and fragments. High beta diversity in coffee
plantations is in contrast with studies in agricultural landscapes, where intensification
was found to lead to a decrease in beta diversity (Escobar-Ramírez et al., 2020).
The beta diversity when decomposed into contribution by species
replacement and nestedness, revealed that there was a steady increase in species
replacement from forests to silver shade coffee plantations with a simultaneous
decrease in nestedness (Figure 11). A similar trend was observed from a study from
South America, where the decrease in similarity of pine plantations with forests was
studied (Santoandré et al., 2019). It led to an increase in the contribution by
replacement towards beta diversity in plantations with most dissimilarity, attributed to
possible environmental filtering. The increase in replacement of species across
transects, leads to higher species pool and thus the higher species richness in silver
shade coffee when compared to forests.
Insecticides and Human influence
Insecticide’s usage has long been known to negatively impact targeted and
non-targeted groups when applied to crops, while the human influence/ disturbances
negatively impacts native fauna as it brings up tramp species (Perfecto, 1990; Rizali
et al., 2010). The reduction in habitat complexity coupled with insecticide usage and
human presence could be driving the high replacement/ turnover across transects in
coffee plantations. The usage of insecticides in the coffee plantations didn’t have any
impact on species richness as the plantations with and without insecticide use, have
comparable species richness. This finding is inconsistent with an earlier study from
the same landscape which found a reduction in species richness due to insecticides
(Mone et al., 2014). The trend observed in this study could be because of the way the
insecticides are applied in the coffee plantations sampled. It is applied in powdered
form and generally dusted on the branches specifically targeting Oecophylla and
Crematogaster sp. to prevent coffee pickers from getting attacked during the picking
31
of coffee. When applied some of it falls on the litter below the coffee plant and might
lead to non-targeted killing/ reduction in abundance of the ants in those areas.
Reduction of abundance or richness of species due to the use of insecticides coupled
with alternation of microhabitat conditions due to canopy opening may facilitate
colonisation by species adapted to drier and more open habitats.
The observed patterns in the species richness might also be influenced by the
type and quality of the matrix surrounding the coffee plantations and fragments
(Perfecto and Vandermeer, 2002). Human presence in terms of residence are known
to bring in tramp species and invasives that might displace the native species around
their area of influence and contribute to higher turnover across transects (Rizali et al.,
2010). Our results also show that coffee plantations with human habitation had a
significantly higher species richness than those without any human habitation.
Species Composition
Species like Meranoplus sp. which have morphological adaptations to living in drier
conditions were found only in coffee plantations (observed in silver shade coffee also
but not collected in pitfalls). It indicates a change in the microhabitat characters in
coffee plantations which favour these species seen in a study in a nearby landscape
where the forests are found to be generally cooler, moist and with little variation in
temperatures, while the coffee plantations are drier and have variable climate
(Badrinarayanan, 2001). Species like Strumigenys sp. which are leaf litter dwelling
species were recorded primarily from forests and fragments where the amount,
thickness, and diversity of leaf litter was high. Genera such as Discothyrea,
Lophomyrmex, Myrmoteras were recorded only in the coffee plantations while the
invasive ant species Anoplolepis gracilepis was encountered only in the native shade
coffee plantations.
Results from rank abundance curve (Figure 10) show that the species
dominance changes with a change in land use, for instance, Oecophylla smaragdina
which is one of the frequently encountered species in coffee plantations is relatively
less common in fragments and encountered just once in forests (one individual).
32
Number of unique species is higher in coffee plantations, a trend observed in
a study by Badrinarayanan (Badrinarayanan, 2001). It can be an artifact of insufficient
sampling effort in this study (Figure 12), as some of the species unique to some land
uses might have been missed due to their low abundance in other land use types.
Coffee plantations, while harbouring some of the species from forests and fragments,
also provide refuge to species unique to them. 34% of species were found only in the
coffee plantations. This shows that coffee plantations play an important role in
maintaining and contributing to the overall diversity of the landscape.
Limitations
• This study was carried out during the late winter, which is not the period of
high activity for ants, as compared to the monsoon season.
• Only leaf litter ant communities were sampled in this study. Sampling the
canopy dwellers is also required to further enhance the understanding of the
ant communities inside these land use types and thus the opportunity to
understand the direct effect of reduction in habitat complexity. However, the
Figure 13: A Venn diagram of species shared between each land use (Total species = 70).
33
number of exclusively arboreal species are very few and hence sampling these
is unlikely to change the results of this study.
• It is likely that we have missed a number of rare species as inferred from the
species accumulation curve.
CONCLUSION
The findings from our study suggest that responses in ant species richness and
composition are not just dependant on structural complexity within a patch but more
by the habitat diversity and surrounding heterogeneity collectively represented by a
land use type. A number of studies have investigated the role of coffee based agro-
forestry in biodiversity conservation. These studies largely advocate the importance
of maintaining native tree cover as shade for enhancing the biodiversity value of this
land use type as compared to mono-dominant silver-oak shade coffee. Our results
suggest that coffee plantations regardless of nature of tree cover are effective in
supporting high diversity of ants. However, our results should be taken with caution
as this was a short-term study. It was not a complete and comprehensive sampling of
ant assemblages in these land use types. It is also important to consider that although
the overall richness of coffee plantations was higher, the species composition of ants
was observed to be different in different land use types highlighting the importance of
each of these individually and also collectively in maintaining the regional species
pool of the Kodagu landscape.
***********
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Appendix 1
Species by site matrix (pooled at sites)
Table 5: Species by site matrix.
Sub family
Species
Acronym
used
Forest
Fragment
Native shade coffee
Silver shade coffee
fr1
fr2
fg1
fg2
fg3
ns1
ns2
ns3
ns4
si1
si2
Formicinae
Acropyga sp1
Aeni_sp1
0
0
0
1
0
0
0
0
0
0
1
Aenictinae
Aenictus sp1
Aeni_sp2
0
0
0
0
0
0
0
0
0
0
2
Aenictinae
Aenictus sp2
Acro_sp1
0
0
0
0
3
62
0
0
0
0
0
Formicinae
Anoplolepis
gracilepis
Anop_grac
0
0
0
0
0
34
70
0
0
0
0
Formicinae
Camponotous
sp1
Camp_sp1
3
68
1
0
1
0
0
4
0
0
0
Formicinae
Camponotous
sp2
Camp_sp2
0
0
1
0
0
0
0
0
0
0
0
Formicinae
Camponotous
sp3
Camp_sp3
3
1
0
0
0
0
0
10
0
1
1
Formicinae
Camponotous
sp4
Camp_sp4
0
0
0
0
0
0
0
0
0
5
0
Formicinae
Camponotous
sp5
Camp_sp5
1
1
0
0
0
0
0
0
0
0
0
Formicinae
Camponotous
spx
Camp_spx
0
0
1
0
0
0
0
0
0
0
0
Myrmicinae
Cardiocondyla
sp1
Card_sp1
2
0
0
0
0
1
8
4
12
1
0
Myrmicinae
Carebara sp1
Care_sp1
0
0
0
0
0
38
6
12
6
1
3
Myrmicinae
Carebara sp2
Care_sp2
23
16
0
0
3
1
0
5
4
0
0
Myrmicinae
Carebara sp3
Care_sp3
0
2
0
0
0
0
0
0
0
0
0
Dorylinae
Cerapachys sp1
Cera_sp1
0
1
0
0
0
0
0
0
0
0
0
Myrmicinae
Crematogaster
sp1
Crem_sp1
209
7
3
1
5
1
5
2
0
0
0
Myrmicinae
Crematogaster
sp2
Crem_sp2
1
1
1
0
0
0
0
0
1
0
1
Myrmicinae
Crematogaster
sp3
Crem_sp3
21
0
0
0
0
0
0
0
0
0
0
Ponerinae
Diacamma sp1
Diac_sp1
0
0
0
0
0
0
0
0
0
1
0
Proceratiinae
Discothyrea sp1
Disc_sp1
0
0
0
0
0
0
0
0
1
0
0
Dorylinae
Dorylus sp1
Dory_sp1
0
0
0
1
0
0
0
0
0
0
0
Ponerinae
Hypoponera sp1
Hypo_sp1
1
5
7
10
0
0
0
2
0
4
1
Formicinae
Lepisiota sp1
Lepi_sp1
0
0
0
0
0
0
0
2
0
0
0
Ponerinae
Leptogenys sp1
Lept_sp1
12
24
9
6
9
3
14
25
5
8
35
Ponerinae
Leptogenys sp2
Lept_sp2
0
0
2
3
1
0
1
3
0
0
0
Ponerinae
Leptogenys sp3
Lept_sp3
0
0
0
0
1
0
0
0
0
0
0
Ponerinae
Leptogenys sp4
Lept_sp4
0
0
0
0
0
0
0
20
0
0
1
Ponerinae
Leptogenys sp5
Lept_sp5
0
14
0
0
0
0
0
5
0
0
5
Ponerinae
Leptogenys sp6
Lept_sp6
0
0
0
0
0
0
0
0
0
114
0
Ponerinae
Leptogenys sp7
Lept_sp7
0
1
0
0
0
0
0
0
0
1
0
Myrmicinae
Lophomyrmex
sp1
Loph_sp1
0
0
0
0
0
0
0
0
0
0
1
Myrmicinae
Meranoplus sp1
Mera_sp1
0
0
0
0
0
0
1
1
10
0
0
Myrmicinae
Meranoplus sp2
Mera_sp2
0
0
0
0
0
0
0
0
1
0
0
Myrmicinae
Monomorium
sp1
Mono_sp1
0
0
0
0
0
2
1
1
0
0
0
Myrmicinae
Monomorium
sp2
Mono_sp2
0
0
0
0
0
0
4
0
3
4
0
Myrmicinae
Monomorium
sp3
Mono_sp3
0
0
0
0
1
0
1
0
0
0
0
Myrmicinae
Monomorium
sp4
Mono_sp4
0
0
2
20
3
0
0
0
0
0
0
Myrmicinae
Myrmecina sp1
Myca_sp1
0
0
0
0
0
0
0
2
0
0
0
Myrmicinae
Myrmicaria sp1
Myci_sp1
0
0
0
0
0
0
0
1
0
0
1
Formicinae
Myrmoteras sp1
Myrm_sp1
0
0
0
0
0
1
0
0
0
0
0
Formicinae
Nylanderia sp1
Nyla_sp1
22
67
158
23
143
0
0
1611
0
39
1
Formicinae
Nylanderia sp2
Nyla_sp2
0
0
0
0
0
0
2
0
0
0
0
Ponerinae
Odontoponera
sp1
Odon_sp1
6
7
3
6
0
3
1
6
1
1
1
Formicinae
Oecophylla
smaragdina
Oeco_smar
1
0
3
6
11
18
16
30
22
18
13
Ponerinae
Pachycondyla
sp1
Pach_sp1
2
3
2
2
6
1
0
2
1
1
0
Ponerinae
Pachycondyla
sp2
Pach_sp2
0
0
12
50
7
5
0
0
85
3
0
Ponerinae
Pachycondyla
sp3
Pach_sp3
0
0
1
0
1
0
0
1
0
0
0
Ponerinae
Pachycondyla
sp4
Pach_sp4
0
0
0
0
0
0
0
0
87
0
0
Myrmicinae
Pheidole sp1
Phei_sp1
1
174
24
20
0
18
40
133
19
26
169
Myrmicinae
Pheidole sp2
Phei_sp2
18
27
18
27
48
21
12
7
33
21
20
Myrmicinae
Pheidole sp3
Phei_sp3
20
19
15
2
64
0
18
0
0
47
6
Myrmicinae
Pheidole sp4
Phei_sp4
2
6
0
15
0
0
0
4
4
4
0
Myrmicinae
Pheidole sp5
Phei_sp5
0
0
0
2
0
0
0
1
5
0
0
Formicinae
Polyrhachis sp1
Poly_sp1
0
0
0
0
0
0
1
0
0
0
0
Formicinae
Polyrhachis sp2
Poly_sp2
0
0
0
0
1
0
0
0
0
0
1
Formicinae
Polyrhachis sp3
Poly_sp3
0
0
0
0
0
0
0
0
1
1
0
Myrmicinae
Strumigenys sp1
Stru_sp1
0
0
0
2
1
0
0
0
0
1
0
Myrmicinae
Strumigenys sp2
Stru_sp2
0
0
0
66
0
0
0
0
0
0
0
Myrmicinae
Strumigenys sp3
Stru_sp3
1
1
0
0
0
0
0
0
0
0
0
Dolichoderinae
Tapinoma sp1
Tapi_sp1
0
0
0
0
0
0
2
0
0
0
0
Dolichoderinae
Tapinoma sp2
Tapi_sp2
0
0
14
0
0
0
0
0
0
0
0
Dolichoderinae
Technomyrmex
sp1
Tech_sp1
3
92
30
15
54
96
5
2
4
80
1
Dolichoderinae
Technomyrmex
sp2
Tech_sp2
0
0
8
0
0
0
0
0
1
1
0
Pseudomyrmecinae
Tetraponera
rufonigra
Tepo_rufo
0
0
0
0
0
0
0
0
1
0
0
Pseudomyrmecinae
Tetraponera sp1
Tepo_sp1
0
1
0
0
0
0
0
0
1
0
0
Myrmicinae
Tetramorium sp1
Tetr_sp1
13
20
8
4
23
5
5
18
3
2
6
Myrmicinae
Tetramorium sp2
Tetr_sp2
0
0
0
0
0
0
0
1
3
0
1
Myrmicinae
Tetramorium sp3
Tetr_sp3
0
2
0
0
0
0
0
0
2
3
2
Myrmicinae
Tetramorium sp4
Tetr_sp4
0
4
0
0
0
0
0
0
1
0
1
Myrmicinae
Tetramorium sp5
Tetr_sp5
0
1
0
0
0
0
0
0
0
0
0
fr1
fr2
fg1
fg2
fg3
ns1
ns2
ns3
si1
si2
si3
Species richness
21
26
22
21
20
17
20
28
27
25
23
Appendix- II-
Drawings of the species collected during the study, few common and stand-alone species weren’t drawn.
