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Int. J. Agri. Agri. R.
Rowena P. Varela Page 140
RESEARCH PAPER OPEN ACCESS
Aquatic Insect Similarity Connecting Natural Wetland Habitat
and Ricefield for Ecological Rice Production in Agusan Marsh
in Mindanao, Philippines
Rowena P. Varela*
Department of Agricultural Sciences, College of Agricultural Sciences and Natural Resources,
Caraga State University, Ampayon, Butuan City Philippines
Article published on July 29, 2016
Key words: Diversity, Species richness, Natural wetland, Lowland rice field, Species similarity.
Abstract
This study describes the relationship of the natural wetland habitats in Agusan Marsh to nearby rice fields and its
implication to ecological rice production. Aquatic insects play multiple roles in the ecosystem such as predators,
prey to other animals and decomposers which help in maintaining ecological balance. Results revealed that the
diversity of odonates was highest in the sedge-dominated swamp among natural habitats which corresponds to
the adjoining ricefields. The pattern of clustering of odonates show 3 groups; the rice-sago and rice-sedges sub-
cluster, the rice-bangkal, rice-Terminalia, rice-fern, bangkal, sago and sedges sub-cluster, and the Terminalia
forest as the outlier. The diversity of semi-aquatic and aquatic bugs was highest in the Bangkal forest while the
lowest was in the fern-dominated swamp. The pattern of clustering shows 2 sub-clusters and the outlier Bangkal
forest. On aquatic beetles, highest diversity was in the Terminalia forest. The sub-cluster consists of Terminalia
habitat and rice-fern, while the other sub-cluster includes rice-Bangkal and rice-Terminalia. The resulting
patterns of similarity in diversity and distribution of species in natural habitats and nearby ricefields indicate that
ricefields are important temporary habitats for some aquatic insect species and serve as stepping stones for the
movement of the insects.
* Corresponding Author: P. Varela  rpvarela@carsu.edu.ph
International Journal of Agronomy and Agricultural Research (IJAAR)
ISSN: 2223-7054 (Print) 2225-3610 (Online)
http://www.innspub.net
Vol. 9, No. 1, p. 140-147, 2016
Int. J. Agri. Agri. R.
Rowena P. Varela Page 141
Introduction
Lowland ricefields are originally natural wetlands,
thus the natural wetland and adjacent ricefields are
expected to be ecologically related in many ways.
However, ecological studies in ricefields for
biodiversity conservation have been conducted
recently. According to Roger (1996), the ecology of
wetland ricefield ecosystems is less studied since it is
regarded as artificial and temporary. Thus only
limited information is available on the sharing of
biodiversity between ricefields and neighboring
ecosystems. There have been reports on ecology of
ricefields citing aquatic insects and their roles in
nutrient cycling and other ecological processes
(Lawler, 2001; Bambaradeniya, et al., 2004). The
high faunal diversity in ricefields has been attributed
to the rapid re-colonization by fauna from contiguous
water bodies after the disturbances caused by cultural
practices or drying during the fallow period. In
contrast, frequent disturbances and the use of
agrochemicals in this ecosystem disrupt the natural
community structure, energy flow, nutrient cycling,
and population dynamics of component species.
Additionally, the considerable decrease in the aquatic
biodiversity in ricefields has been ascribed to the loss
of permanent water reservoirs near ricefield.
The colonization of aquatic insects in ricefields from
adjoining water bodies is a clear indication of the
relationship between natural wetlands and ricefields.
However, studies conducted on the relationship of
flora and fauna of ricefields to adjacent habitats gave
emphasis only to the terrestrial and canopy-dwelling
species (Marcos, et al., 2001). Some studies on agro
biodiversity in relation to other habitats have
recommended the need to consider the landscape
approach (Liu, et al., 2013; Lan, et al., 2001).
Movement of aquatic insects between and within
habitats is an important aspect to consider in agro
biodiversity, however, studies on this remain
inadequate. Many aquatic insects are strong flyers.
Movement through flight may be one of the
mechanisms relating the diversity of the natural
wetland habitats and ricefields.
Due to their flight capability, these insects readily
move from one aquatic habitat to another, especially
when conditions are not favorable for survival.
However, most aquatic insects move through passive
dispersal. In passive dispersal of animals, corridors
are important. Corridors are units of the landscape
that interconnect one patch with another (Foreman
and Godron, 1986; Foreman, 1995). One interesting
study by Fairchild, et al. (2003) focused on the
influence of microhabitat and landscape on aquatic
beetle assemblages in temporary and permanent
ponds. Indeed, the aquatic beetle assemblages were
strongly correlated with the characteristics of
landscape and microhabitats. In ricefields, for
instance, where habitats change with the application
of cultural practices, aquatic insects may disperse in
search of suitable habitats. Thus this study was done
to determine the relationship of the natural wetland
habitats in Agusan Marsh to the continuing expansion
of ricefields in the vicinity. This study likewise intends
to examine if ricefields have the potential role as
corridors for biodiversity movement.
Materials and method
Study Site
The study was conducted in the Agusan Marsh, in the
northeastern part of the island of Mindanao in
Southern Philippines in July 2010 to June 2011 (Fig.
1). The marsh has an area of 111,540 hectares which is
categorized into 7 habitat types that include
freshwater swamp forests, secondary scrubs,
herbaceous swamps, lakes, pools and rivers, and the
human-dominated habitats such as rice paddies and
other agricultural lands (AMWS Master Plan, 2001).
Fig. 1. Land cover map of Agusan Marsh in
Mindanao, Philippines showing the study sites
(enclosed in yellow box).
Int. J. Agri. Agri. R.
Rowena P. Varela Page 142
Insect sampling and processing of specimens
Aquatic insects with emphasis on Hemiptera,
Odonata, and Coleoptera were sampled from the
major aquatic habitats of Agusan Marsh and adjacent
ricefields. The sampled area measured approximately
500 sq. meters in each study site. Insects that actively
swim were collected from the sites using a dip net,
while those that cling on aquatic vegetation and other
debris within the 1 sq m quadrat were collected by
washing them off in a container with water.
For insects dwelling on the water surface (e.g. water
striders, pond skaters), the net was swept over the
water swiftly. Sampling was done for two rice
cropping seasons.
Aquatic insects were identified using taxonomic keys.
Aquatic and semi-aquatic bugs were identified using
Chen et al. (2005) and Zettel and Gapud (1999).
Aquatic beetles were determined using Balke, et al.
(2002), and the hydrophilids using Hansen (1999).
Community structure analysis
Trends in community structure of aquatic insects
were analyzed using the corresponding measu-
rements. Species richness involved actual counts of
species collected for the different insect groups in the
natural habitats and adjoining ricefields. Species
diversity indices were measured using the Shannon-
Wiener function which accounts for the number of
species and the number of individuals in each species,
and is expressed as:
where H’ = index of species diversity ; log2 =
3.321928
s = number of species
pi = proportion of total sample belonging to
ith species
In determining equitability of distribution of
individuals in each species, the Jaccard’s index of
evenness was computed as follows:
J’ = H’
H’MAX
where J’ = evenness measure (range 0 – 1)
H’ = Shannon-Wiener function
H’MAX = maximum value of H’ = log S (no.
of species in sample)
The similarities in species compositions among the
natural habitats and ricefelds were analyzed using the
Pearson’s coefficient of similarity. To accomplish
this, the single linkage cluster analysis in the
Multivariate Statistical Package (MVSP) was used.
Results and discussion
Diversity of dragonflies and damselflies between
natural habitat and rice fields.
The diversity of dragonflies and damselflies (Order
Odonata) was highest in the sedge-dominated swamp
among the natural habitats (Table 1).
The lowest was in the Bangkal forest at 2.077. In
adjacent ricefields, the diversity was similarly highest
in site near the sedge-dominated swamp. The lowest
diversity index was in ricefield adjoining the fern-
dominated swamp.
The result reflects the physical characteristics of the
habitats. In the sedge-dominated swamp, water is
generally present even during the dry months. A
similar characteristic is also noted in the Terminalia
forest, where sedges also grow in between the
Terminalia trees.
In the rest of the natural habitats, water becomes
scarce during the drier months thus some insect
species move to where water is available. In ricefield
near the sedge-dominated swamp, the characteristic
is very similar to that of the natural habitat because
these ricefields are originally sedge-dominated
swamps. Elphick (2001) studied the potential for
human-disturbed habitats to act as surrogates for the
natural habitats they replace.
He noted that flooded fields seemingly provide
comparable foraging habitat to seminatural wetlands
for waterbirds. Thus flooded ricefields, if managed
properly, can provide valuable waterbird habitat.
Int. J. Agri. Agri. R.
Rowena P. Varela Page 143
Table 1. Diversity of dragonflies and damselflies in natural habitats and adjacent ricefields of Agusan Marsh,
Mindanao, Philippines.
Sampling Site Shannon Diversity Evenness Species Richness
Fern-dominated swamp 2.507 0.893 7
Sedge-dominated swamp 2.906 0.875 10
Sago forest 2.206 0.853 6
Terminalia forest 2.524 0.899 7
Bangkal forest 2.077 0.895 5
Rice-Fern 1.648 0.824 4
Rice-Sedges 2.768 0.873 9
Rice-Sago 2.190 0.847 6
Rice-Terminalia 1.963 0.759 6
Rice-Bangkal 2.026 0.873 5
Species composition of aquatic insects in the different
natural habitats of the Agusan Marsh was expected to
have closer similarities with their adjacent rice fields.
However, the pattern of clustering of the odonates
show 3 groups: the rice-sago and rice-sedges sub-
cluster, then the rice-bangkal, rice-Terminalia, rice-
fern, bangkal, sago and sedges sub-cluster, and the
Terminalia forest as the outlier (Fig. 2).
A large sub-cluster consisting of the rice-bangkal,
rice-Terminalia, rice-fern, bangkal, sago and sedges
has a similarity index of about 91%.
The rice-sago and rice-sedges have 94% similarity
and joined to the other sub-cluster by a similarity
index of about 90%. The outlier habitat, the
Terminalia forest, joins the 2 sub-clusters at 83%
level of similarity. This suggests that the Terminalia
forest was more dissimilar in the odonate species
composition. This can be attributed to the
characteristics of the habitat where open spaces are
smaller resulting to less sunlight reaching the ground.
Fig. 2. Similarity of Odonatan species in the different
natural habitats and adjoining ricefields in Agusan
Marsh.
Similarity of semi-aquatic and aquatic bug diversity
between natural habitat and rice fields.
The diversity of semi-aquatic and aquatic bugs (Order
Hemiptera) in natural habitats and their adjacent
ricefields was highest in the Bangkal forest (Table 2).
The lowest was in the fern-dominated swamp, which
is very similar to the diversity indices of ricefield
habitats.
The result can be attributed to the behavior of the
insects and natural habitat characteristics. In Bangkal
forest, the water is a combination of stagnant and
moving which attracted different species of aquatic
and semi-aquatic bugs. A closer characteristic was
noted in the Sago forest, thus species that dwells on
both stagnant and slowly moving water were also
found.
The diversity indices are generally low in all habitats
due to the uneven distribution of the population. In
the sedge-dominated swamp, sago and Bangkal
forests, the indices of evenness are relatively higher
resulting from the higher species richness and
populations that are more evenly distributed. Species
composition of aquatic and semi-aquatic bugs in the
different natural habitats of the Agusan Marsh was
expected to have closer similarities. Nonetheless, the
clustering pattern shows 2 sub-clusters and the
outlier Bangkal forest (Fig. 3).
The bigger sub-cluster includes 6 habitats: rice-sago
and rice-Bangkal, the sago, rice-sedges, Terminalia
and sedges. The other sub-cluster is composed of rice-
fern, the rice- Terminalia forest and fern.
Int. J. Agri. Agri. R.
Rowena P. Varela Page 144
Table 2. Diversity of semi-aquatic and aquatic bugs in natural habitats and adjacent ricefields of Agusan Marsh,
Mindanao, Philippines.
Sampling Site Shannon Diversity Evenness Species Richness
Fern-dominated swamp 1.286 0.458 7
Sedge-dominated swamp 2.277 0.658 11
Sago forest 2.340 0.705 10
Terminalia forest 1.769 0.533 10
Bangkal forest 2.645 0.882 8
Rice-Fern 1.101 0.392 7
Rice-Sedges 1.504 0.582 6
Rice-Sago 1.505 0.582 6
Rice-Terminalia 1.164 0.450 6
Rice-Bangkal 1.274 0.493 6
The same is true for the other sub-cluster, consisting
of rice-fern, the rice-Terminalia forest and fern,
having a similarity index of almost 100%. The outlier
habitat, the Bangkal forest, joins the 2 sub-clusters at
70% level of similarity. This suggests that the species
composition of Bangkal forest was more different
than the rest of the habitats.
Fig. 3. Similarity of Hemipteran species in the
different natural habitats and adjoining ricefields in
Agusan Marsh.
The rice fields have generally lower diversity indices
than the natural habitats resulting from the fewer
number of species. In the study of Ma et al. (2004),
they found that natural wetlands are better habitats
for water birds than artificial wetlands on Chongming
Island, while the artificial ones are suitable habitats
for water birds in winter.
The birds use artificial wetlands only when natural
wetlands are unavailable or of poor quality. However,
they cautioned that overemphasis on suitability of
artificial wetlands for
water birds might encourage land managers to
convert natural wetlands into the artificial ones which
may result in considerable loss of bird diversity.
Similarity of aquatic beetle diversity between
natural habitat and ricefields.
The diversity of aquatic beetles in natural habitats
and adjacent ricefields was highest in the Terminalia
forest (Table 3). The result can be attributed to the
presence of some unique species found only in
Terminalia forest.
All other natural habitats have diversity indices closer
to that of the Terminalia forest. The lowest was in the
Bangkal forest, which has generally similar diversity
indices with ricefield habitats. This suggests that the
various species of aquatic beetles differed in habitat
requirements thus there are species that are not
found in certain habitats. In Bangkal forest, the
diversity index of aquatic beetles is closer to the
diversity of ricefield habitats.
This can be attributed to the habitat requirements of
certain beetle species, such as the water scavenger
beetles that thrive in microhabitats where decaying
materials abound. The indices of evenness are
generally higher in all habitats due to the relatively
even distribution of various populations,except for
the unique species. In Terminalia forests and its
adjoining ricefield, the number of species is higher,
leading to a relatively higher diversity index for both
habitats.
Int. J. Agri. Agri. R.
Rowena P. Varela Page 145
Table 3. Diversity of aquatic beetles in natural habitats and adjacent ricefields of Agusan Marsh, Mindanao,
Philippines.
Sampling Site Shannon Diversity Evenness Species Richness
Fern-dominated swamp 2.394 0.798 8
Sedge-dominated swamp 2.411 0.761 9
Sago forest 2.343 0.781 8
Terminalia forest 2.578 0.776 10
Bangkal forest 2.174 0.774 7
Rice-Fern 2.075 0.739 7
Rice-Sedges 2.118 0.754 7
Rice-Sago 1.930 0.688 7
Rice-Terminalia 2.173 0.685 9
Rice-Bangkal 1.831 0.708 6
The water beetles, which constitute about half of the
total species, have a very visible pattern of species
similarities (Fig. 4). Two clusters are apparent, with 2
sub-clusters splitting at 96%. The first cluster
consists of Terminalia habitat and rice-fern, while the
second cluster includes rice-Bangkal and rice-
Terminalia, joining with rice-sago, Bangkal, sago,
sedges and rice-sedges. The first cluster consisting of
Terminalia habitat and rice-fern is separated by a
considerable distance from the other sub-cluster. It is
apparent that the species similarities within the two
sub-clusters are not due to proximity but possibly on
the similarity in physical characteristics or vegetation
structure in these habitats.
Fig. 4. Similarity of Coleopteran species in the
different natural habitats and adjoining ricefields in
Agusan Marsh.
Land-use is an important factor that influences faunal
distribution and diversity. According to Bennett et al.
(2007), the extent of habitat is usually an important
influence on single species existence or the richness of
assemblages defined by habitat type. Similarly, the
composition of the land-use mosaic, based on the
proportions of elements, strongly influences the
species composition of faunal assemblages and that
heterogeneity of elements is often positively
correlated with the richness of taxonomic assem-
blages. Wilby et al. (2006) reported that the
arthropod community associated with rice revealed
characteristic seasonal differences in guild structure
due to patterns in community assembly through the
cropping season. Moreover, the arthropod diversity
across all land-use types generally declined with the
land use gradient towards extensive rice monoculture.
They added that with particular reference to diversity
and community structure in rice, landscape diversity
influenced the processes of community assembly in
rice largely through effects on abundance rather than
diversity and that the predator guild was least
affected. This finding is a necessary input for
planning the ecological production of rice where
diversity of arthropods is viewed to contribute in
bringing equilibrium resulting from the interactions
in the ecosystem.
Conclusion
The resulting patterns of similarity in diversity and
distribution of aquatic insect species in natural
habitats and nearby ricefields indicate that indeed
ricefields are important temporary habitats for
certain species of aquatic insects. Although in some
cases no close similarities in species composition is
apparent between natural habitats and rice fields, the
rice fields can still be regarded as corridors or
stepping stones for the movement of aquatic insects
from one natural habitat to the next. It is evident that
the major reasons in the similarity of species
composition between natural habitats and the
associated rice fields are those pertaining to resource
availability.
Int. J. Agri. Agri. R.
Rowena P. Varela Page 146
In addition, the similarity relationships clearly
demonstrate the active movement of aquatic insects,
regardless of habitat proximity. The similarity of
species between the natural wetland habitats and
ricefields indicates that the aquatic insect
communities are important factors for check and
balance in ricefields thereby controlling increase in
insect pest populations. This condition is important in
understanding ecological rice pest management.
Acknowledgment
The author acknowledges the research grant from the
Commission on Higher Education (CHED) for the
financial support.
Literature cited
AMWS. 2001. Agusan Marsh Wildlife Sanctuary
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Balke M, Jach, MA and Hendrich, L. 2002.
Water beetles of Malaysia (Coleoptera). Report on the
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Bambaradeniya CN, Amerasinghe FP. 2004.
Biodiversity associated with the rice field agroecosystem
in Asian countries: a brief review IWMI;
Barrion AT and Litsinger, JA. 1994. Biology and
Management of Rice Insects. In: EA Heinreich (ed.).
Wiley and Sons Ltd., Delhi, India. pp 50-345.
Bennett AF, Radford JQ, Haslem A. 2006.
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Chen PP, Nieser N, Zettel H. 2005. Aquatic and
semi-aquatic bugs (Heteroptera: Nepomorpha &
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Fairchild GW, Cruz J, Faulds AM, Short AEZ
Matta JF. 2003. Microhabitat and landscape influences
on aquatic beetle assemblages in a cluster of temporary
and permanent ponds. Journal of the North American
Benthological Society 22(2), pp. 224-240.
Foreman RTT, Godron M. 1986. Landscape
Ecology. John Wiley and Sons, New York 620 p.
Forman RT. 1995. Some general principles of
landscape and regional ecology. Landscape ecology 1,
10(3), 133-42.
Gapud, VP, Zettel H. 1999. "The Philippine Water
Bug Inventory Project (PWBIP) and a bibliography
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Aquatic Insect Similarity Connecting Natural Wetland Habitat and Ricefield for Ecological Rice Production in Agusan Marsh in Mindanao, Philippines

  • 1. Int. J. Agri. Agri. R. Rowena P. Varela Page 140 RESEARCH PAPER OPEN ACCESS Aquatic Insect Similarity Connecting Natural Wetland Habitat and Ricefield for Ecological Rice Production in Agusan Marsh in Mindanao, Philippines Rowena P. Varela* Department of Agricultural Sciences, College of Agricultural Sciences and Natural Resources, Caraga State University, Ampayon, Butuan City Philippines Article published on July 29, 2016 Key words: Diversity, Species richness, Natural wetland, Lowland rice field, Species similarity. Abstract This study describes the relationship of the natural wetland habitats in Agusan Marsh to nearby rice fields and its implication to ecological rice production. Aquatic insects play multiple roles in the ecosystem such as predators, prey to other animals and decomposers which help in maintaining ecological balance. Results revealed that the diversity of odonates was highest in the sedge-dominated swamp among natural habitats which corresponds to the adjoining ricefields. The pattern of clustering of odonates show 3 groups; the rice-sago and rice-sedges sub- cluster, the rice-bangkal, rice-Terminalia, rice-fern, bangkal, sago and sedges sub-cluster, and the Terminalia forest as the outlier. The diversity of semi-aquatic and aquatic bugs was highest in the Bangkal forest while the lowest was in the fern-dominated swamp. The pattern of clustering shows 2 sub-clusters and the outlier Bangkal forest. On aquatic beetles, highest diversity was in the Terminalia forest. The sub-cluster consists of Terminalia habitat and rice-fern, while the other sub-cluster includes rice-Bangkal and rice-Terminalia. The resulting patterns of similarity in diversity and distribution of species in natural habitats and nearby ricefields indicate that ricefields are important temporary habitats for some aquatic insect species and serve as stepping stones for the movement of the insects. * Corresponding Author: P. Varela  rpvarela@carsu.edu.ph International Journal of Agronomy and Agricultural Research (IJAAR) ISSN: 2223-7054 (Print) 2225-3610 (Online) http://www.innspub.net Vol. 9, No. 1, p. 140-147, 2016
  • 2. Int. J. Agri. Agri. R. Rowena P. Varela Page 141 Introduction Lowland ricefields are originally natural wetlands, thus the natural wetland and adjacent ricefields are expected to be ecologically related in many ways. However, ecological studies in ricefields for biodiversity conservation have been conducted recently. According to Roger (1996), the ecology of wetland ricefield ecosystems is less studied since it is regarded as artificial and temporary. Thus only limited information is available on the sharing of biodiversity between ricefields and neighboring ecosystems. There have been reports on ecology of ricefields citing aquatic insects and their roles in nutrient cycling and other ecological processes (Lawler, 2001; Bambaradeniya, et al., 2004). The high faunal diversity in ricefields has been attributed to the rapid re-colonization by fauna from contiguous water bodies after the disturbances caused by cultural practices or drying during the fallow period. In contrast, frequent disturbances and the use of agrochemicals in this ecosystem disrupt the natural community structure, energy flow, nutrient cycling, and population dynamics of component species. Additionally, the considerable decrease in the aquatic biodiversity in ricefields has been ascribed to the loss of permanent water reservoirs near ricefield. The colonization of aquatic insects in ricefields from adjoining water bodies is a clear indication of the relationship between natural wetlands and ricefields. However, studies conducted on the relationship of flora and fauna of ricefields to adjacent habitats gave emphasis only to the terrestrial and canopy-dwelling species (Marcos, et al., 2001). Some studies on agro biodiversity in relation to other habitats have recommended the need to consider the landscape approach (Liu, et al., 2013; Lan, et al., 2001). Movement of aquatic insects between and within habitats is an important aspect to consider in agro biodiversity, however, studies on this remain inadequate. Many aquatic insects are strong flyers. Movement through flight may be one of the mechanisms relating the diversity of the natural wetland habitats and ricefields. Due to their flight capability, these insects readily move from one aquatic habitat to another, especially when conditions are not favorable for survival. However, most aquatic insects move through passive dispersal. In passive dispersal of animals, corridors are important. Corridors are units of the landscape that interconnect one patch with another (Foreman and Godron, 1986; Foreman, 1995). One interesting study by Fairchild, et al. (2003) focused on the influence of microhabitat and landscape on aquatic beetle assemblages in temporary and permanent ponds. Indeed, the aquatic beetle assemblages were strongly correlated with the characteristics of landscape and microhabitats. In ricefields, for instance, where habitats change with the application of cultural practices, aquatic insects may disperse in search of suitable habitats. Thus this study was done to determine the relationship of the natural wetland habitats in Agusan Marsh to the continuing expansion of ricefields in the vicinity. This study likewise intends to examine if ricefields have the potential role as corridors for biodiversity movement. Materials and method Study Site The study was conducted in the Agusan Marsh, in the northeastern part of the island of Mindanao in Southern Philippines in July 2010 to June 2011 (Fig. 1). The marsh has an area of 111,540 hectares which is categorized into 7 habitat types that include freshwater swamp forests, secondary scrubs, herbaceous swamps, lakes, pools and rivers, and the human-dominated habitats such as rice paddies and other agricultural lands (AMWS Master Plan, 2001). Fig. 1. Land cover map of Agusan Marsh in Mindanao, Philippines showing the study sites (enclosed in yellow box).
  • 3. Int. J. Agri. Agri. R. Rowena P. Varela Page 142 Insect sampling and processing of specimens Aquatic insects with emphasis on Hemiptera, Odonata, and Coleoptera were sampled from the major aquatic habitats of Agusan Marsh and adjacent ricefields. The sampled area measured approximately 500 sq. meters in each study site. Insects that actively swim were collected from the sites using a dip net, while those that cling on aquatic vegetation and other debris within the 1 sq m quadrat were collected by washing them off in a container with water. For insects dwelling on the water surface (e.g. water striders, pond skaters), the net was swept over the water swiftly. Sampling was done for two rice cropping seasons. Aquatic insects were identified using taxonomic keys. Aquatic and semi-aquatic bugs were identified using Chen et al. (2005) and Zettel and Gapud (1999). Aquatic beetles were determined using Balke, et al. (2002), and the hydrophilids using Hansen (1999). Community structure analysis Trends in community structure of aquatic insects were analyzed using the corresponding measu- rements. Species richness involved actual counts of species collected for the different insect groups in the natural habitats and adjoining ricefields. Species diversity indices were measured using the Shannon- Wiener function which accounts for the number of species and the number of individuals in each species, and is expressed as: where H’ = index of species diversity ; log2 = 3.321928 s = number of species pi = proportion of total sample belonging to ith species In determining equitability of distribution of individuals in each species, the Jaccard’s index of evenness was computed as follows: J’ = H’ H’MAX where J’ = evenness measure (range 0 – 1) H’ = Shannon-Wiener function H’MAX = maximum value of H’ = log S (no. of species in sample) The similarities in species compositions among the natural habitats and ricefelds were analyzed using the Pearson’s coefficient of similarity. To accomplish this, the single linkage cluster analysis in the Multivariate Statistical Package (MVSP) was used. Results and discussion Diversity of dragonflies and damselflies between natural habitat and rice fields. The diversity of dragonflies and damselflies (Order Odonata) was highest in the sedge-dominated swamp among the natural habitats (Table 1). The lowest was in the Bangkal forest at 2.077. In adjacent ricefields, the diversity was similarly highest in site near the sedge-dominated swamp. The lowest diversity index was in ricefield adjoining the fern- dominated swamp. The result reflects the physical characteristics of the habitats. In the sedge-dominated swamp, water is generally present even during the dry months. A similar characteristic is also noted in the Terminalia forest, where sedges also grow in between the Terminalia trees. In the rest of the natural habitats, water becomes scarce during the drier months thus some insect species move to where water is available. In ricefield near the sedge-dominated swamp, the characteristic is very similar to that of the natural habitat because these ricefields are originally sedge-dominated swamps. Elphick (2001) studied the potential for human-disturbed habitats to act as surrogates for the natural habitats they replace. He noted that flooded fields seemingly provide comparable foraging habitat to seminatural wetlands for waterbirds. Thus flooded ricefields, if managed properly, can provide valuable waterbird habitat.
  • 4. Int. J. Agri. Agri. R. Rowena P. Varela Page 143 Table 1. Diversity of dragonflies and damselflies in natural habitats and adjacent ricefields of Agusan Marsh, Mindanao, Philippines. Sampling Site Shannon Diversity Evenness Species Richness Fern-dominated swamp 2.507 0.893 7 Sedge-dominated swamp 2.906 0.875 10 Sago forest 2.206 0.853 6 Terminalia forest 2.524 0.899 7 Bangkal forest 2.077 0.895 5 Rice-Fern 1.648 0.824 4 Rice-Sedges 2.768 0.873 9 Rice-Sago 2.190 0.847 6 Rice-Terminalia 1.963 0.759 6 Rice-Bangkal 2.026 0.873 5 Species composition of aquatic insects in the different natural habitats of the Agusan Marsh was expected to have closer similarities with their adjacent rice fields. However, the pattern of clustering of the odonates show 3 groups: the rice-sago and rice-sedges sub- cluster, then the rice-bangkal, rice-Terminalia, rice- fern, bangkal, sago and sedges sub-cluster, and the Terminalia forest as the outlier (Fig. 2). A large sub-cluster consisting of the rice-bangkal, rice-Terminalia, rice-fern, bangkal, sago and sedges has a similarity index of about 91%. The rice-sago and rice-sedges have 94% similarity and joined to the other sub-cluster by a similarity index of about 90%. The outlier habitat, the Terminalia forest, joins the 2 sub-clusters at 83% level of similarity. This suggests that the Terminalia forest was more dissimilar in the odonate species composition. This can be attributed to the characteristics of the habitat where open spaces are smaller resulting to less sunlight reaching the ground. Fig. 2. Similarity of Odonatan species in the different natural habitats and adjoining ricefields in Agusan Marsh. Similarity of semi-aquatic and aquatic bug diversity between natural habitat and rice fields. The diversity of semi-aquatic and aquatic bugs (Order Hemiptera) in natural habitats and their adjacent ricefields was highest in the Bangkal forest (Table 2). The lowest was in the fern-dominated swamp, which is very similar to the diversity indices of ricefield habitats. The result can be attributed to the behavior of the insects and natural habitat characteristics. In Bangkal forest, the water is a combination of stagnant and moving which attracted different species of aquatic and semi-aquatic bugs. A closer characteristic was noted in the Sago forest, thus species that dwells on both stagnant and slowly moving water were also found. The diversity indices are generally low in all habitats due to the uneven distribution of the population. In the sedge-dominated swamp, sago and Bangkal forests, the indices of evenness are relatively higher resulting from the higher species richness and populations that are more evenly distributed. Species composition of aquatic and semi-aquatic bugs in the different natural habitats of the Agusan Marsh was expected to have closer similarities. Nonetheless, the clustering pattern shows 2 sub-clusters and the outlier Bangkal forest (Fig. 3). The bigger sub-cluster includes 6 habitats: rice-sago and rice-Bangkal, the sago, rice-sedges, Terminalia and sedges. The other sub-cluster is composed of rice- fern, the rice- Terminalia forest and fern.
  • 5. Int. J. Agri. Agri. R. Rowena P. Varela Page 144 Table 2. Diversity of semi-aquatic and aquatic bugs in natural habitats and adjacent ricefields of Agusan Marsh, Mindanao, Philippines. Sampling Site Shannon Diversity Evenness Species Richness Fern-dominated swamp 1.286 0.458 7 Sedge-dominated swamp 2.277 0.658 11 Sago forest 2.340 0.705 10 Terminalia forest 1.769 0.533 10 Bangkal forest 2.645 0.882 8 Rice-Fern 1.101 0.392 7 Rice-Sedges 1.504 0.582 6 Rice-Sago 1.505 0.582 6 Rice-Terminalia 1.164 0.450 6 Rice-Bangkal 1.274 0.493 6 The same is true for the other sub-cluster, consisting of rice-fern, the rice-Terminalia forest and fern, having a similarity index of almost 100%. The outlier habitat, the Bangkal forest, joins the 2 sub-clusters at 70% level of similarity. This suggests that the species composition of Bangkal forest was more different than the rest of the habitats. Fig. 3. Similarity of Hemipteran species in the different natural habitats and adjoining ricefields in Agusan Marsh. The rice fields have generally lower diversity indices than the natural habitats resulting from the fewer number of species. In the study of Ma et al. (2004), they found that natural wetlands are better habitats for water birds than artificial wetlands on Chongming Island, while the artificial ones are suitable habitats for water birds in winter. The birds use artificial wetlands only when natural wetlands are unavailable or of poor quality. However, they cautioned that overemphasis on suitability of artificial wetlands for water birds might encourage land managers to convert natural wetlands into the artificial ones which may result in considerable loss of bird diversity. Similarity of aquatic beetle diversity between natural habitat and ricefields. The diversity of aquatic beetles in natural habitats and adjacent ricefields was highest in the Terminalia forest (Table 3). The result can be attributed to the presence of some unique species found only in Terminalia forest. All other natural habitats have diversity indices closer to that of the Terminalia forest. The lowest was in the Bangkal forest, which has generally similar diversity indices with ricefield habitats. This suggests that the various species of aquatic beetles differed in habitat requirements thus there are species that are not found in certain habitats. In Bangkal forest, the diversity index of aquatic beetles is closer to the diversity of ricefield habitats. This can be attributed to the habitat requirements of certain beetle species, such as the water scavenger beetles that thrive in microhabitats where decaying materials abound. The indices of evenness are generally higher in all habitats due to the relatively even distribution of various populations,except for the unique species. In Terminalia forests and its adjoining ricefield, the number of species is higher, leading to a relatively higher diversity index for both habitats.
  • 6. Int. J. Agri. Agri. R. Rowena P. Varela Page 145 Table 3. Diversity of aquatic beetles in natural habitats and adjacent ricefields of Agusan Marsh, Mindanao, Philippines. Sampling Site Shannon Diversity Evenness Species Richness Fern-dominated swamp 2.394 0.798 8 Sedge-dominated swamp 2.411 0.761 9 Sago forest 2.343 0.781 8 Terminalia forest 2.578 0.776 10 Bangkal forest 2.174 0.774 7 Rice-Fern 2.075 0.739 7 Rice-Sedges 2.118 0.754 7 Rice-Sago 1.930 0.688 7 Rice-Terminalia 2.173 0.685 9 Rice-Bangkal 1.831 0.708 6 The water beetles, which constitute about half of the total species, have a very visible pattern of species similarities (Fig. 4). Two clusters are apparent, with 2 sub-clusters splitting at 96%. The first cluster consists of Terminalia habitat and rice-fern, while the second cluster includes rice-Bangkal and rice- Terminalia, joining with rice-sago, Bangkal, sago, sedges and rice-sedges. The first cluster consisting of Terminalia habitat and rice-fern is separated by a considerable distance from the other sub-cluster. It is apparent that the species similarities within the two sub-clusters are not due to proximity but possibly on the similarity in physical characteristics or vegetation structure in these habitats. Fig. 4. Similarity of Coleopteran species in the different natural habitats and adjoining ricefields in Agusan Marsh. Land-use is an important factor that influences faunal distribution and diversity. According to Bennett et al. (2007), the extent of habitat is usually an important influence on single species existence or the richness of assemblages defined by habitat type. Similarly, the composition of the land-use mosaic, based on the proportions of elements, strongly influences the species composition of faunal assemblages and that heterogeneity of elements is often positively correlated with the richness of taxonomic assem- blages. Wilby et al. (2006) reported that the arthropod community associated with rice revealed characteristic seasonal differences in guild structure due to patterns in community assembly through the cropping season. Moreover, the arthropod diversity across all land-use types generally declined with the land use gradient towards extensive rice monoculture. They added that with particular reference to diversity and community structure in rice, landscape diversity influenced the processes of community assembly in rice largely through effects on abundance rather than diversity and that the predator guild was least affected. This finding is a necessary input for planning the ecological production of rice where diversity of arthropods is viewed to contribute in bringing equilibrium resulting from the interactions in the ecosystem. Conclusion The resulting patterns of similarity in diversity and distribution of aquatic insect species in natural habitats and nearby ricefields indicate that indeed ricefields are important temporary habitats for certain species of aquatic insects. Although in some cases no close similarities in species composition is apparent between natural habitats and rice fields, the rice fields can still be regarded as corridors or stepping stones for the movement of aquatic insects from one natural habitat to the next. It is evident that the major reasons in the similarity of species composition between natural habitats and the associated rice fields are those pertaining to resource availability.
  • 7. Int. J. Agri. Agri. R. Rowena P. Varela Page 146 In addition, the similarity relationships clearly demonstrate the active movement of aquatic insects, regardless of habitat proximity. The similarity of species between the natural wetland habitats and ricefields indicates that the aquatic insect communities are important factors for check and balance in ricefields thereby controlling increase in insect pest populations. This condition is important in understanding ecological rice pest management. Acknowledgment The author acknowledges the research grant from the Commission on Higher Education (CHED) for the financial support. Literature cited AMWS. 2001. Agusan Marsh Wildlife Sanctuary Management Plan. PAMB and DENR Caraga Region, Mindanao 114 p. Balke M, Jach, MA and Hendrich, L. 2002. Water beetles of Malaysia (Coleoptera). Report on the Survey on Aquatic Insects in Malaysia. Bambaradeniya CN, Amerasinghe FP. 2004. Biodiversity associated with the rice field agroecosystem in Asian countries: a brief review IWMI; Barrion AT and Litsinger, JA. 1994. Biology and Management of Rice Insects. In: EA Heinreich (ed.). Wiley and Sons Ltd., Delhi, India. pp 50-345. Bennett AF, Radford JQ, Haslem A. 2006. Properties of land mosaics: implications for nature conservation in agricultural environments. Biological Conservation 30, 133(2):250-64. Chen PP, Nieser N, Zettel H. 2005. Aquatic and semi-aquatic bugs (Heteroptera: Nepomorpha & Gerromorpha) of Malesia Brill. Elphick CS. 2000. Functional equivalency between rice fields and seminatural wetland habitats. Conservation Biology 1, 14(1):181-91. Fairchild GW, Cruz J, Faulds AM, Short AEZ Matta JF. 2003. Microhabitat and landscape influences on aquatic beetle assemblages in a cluster of temporary and permanent ponds. Journal of the North American Benthological Society 22(2), pp. 224-240. Foreman RTT, Godron M. 1986. Landscape Ecology. John Wiley and Sons, New York 620 p. Forman RT. 1995. Some general principles of landscape and regional ecology. Landscape ecology 1, 10(3), 133-42. Gapud, VP, Zettel H. 1999. "The Philippine Water Bug Inventory Project (PWBIP) and a bibliography for Philippine Nepomorpha, Gerromorpha, and Leptopodomorpha (Insecta: Heteroptera)." Annalen des Naturhistorischen Museums in Wien. Serie B für Botanik und Zoologie 35-40. Hansen M. 1999. World catalogue of insects. Volume 2: Hydrophiloidea (s. str.) (Coleoptera). Apollo Books. Lan LP, Huyen ND, Quang NH, Minh NV. 2001. Habitat diversity: an approach to the preservation of natural enemies of tropical irrigated rice insect pests. Exploiting Bidiversity for Sustainable Pest management (Eds., Mew, TW, Borromeo, E. and Hardy B.). 51-63. Lawler SP. 2001. Rice fields as temporary wetlands: a review. Israel Journal of Zoology 1, 47(4), 513-28. Liu Y, Duan M, Yu Z. 2013. Agricultural landscapes and biodiversity in China. Agriculture, ecosystems & environment 15, 166:46-54. Ma Z, Li B, Zhao B, Jing K, Tang S, Chen J. 2004. Are artificial wetlands good alternatives to natural wetlands for waterbirds?–A case study on Chongming Island, China. Biodiversity & Conse- rvation 1, 13(2), 333-50. Marcos TF, Flor LB, Velilla AR, Schoenly KG, Manalo JO, Ofilas OM, Teng PS, Ulep JR, Tinguil MB, Mew TW, Estoy AB. 2001. Relationships between pests and natural enemies in rainfed rice and associated crop and wild habitats in Ilocos Norte, Philippines. Exploiting biodiversity for sustainable pest management: Proceedings, Mew, TW, E. Borromeo, and B. Hardy (Eds.). Los Baños, Laguna (Philippines): IRRI 23-42.
  • 8. Int. J. Agri. Agri. R. Rowena P. Varela Page 147 Roger, PA. 1996. Biology and management of the floodwater ecosystem in rice fields. Int. Rice Res Inst. Settle WH, Ariawan H, Astuti ET, Cahyana W, Hakim AL, Hindayana D, Lestari AS. Pajarningsih S. 1996. Managing tropical rice pests through conservation of general natural enemies and alternative prey. Ecology 77, 1975-1988. Shepard BM, Barrion AT. Litsinger JA. 2000. Helpful Insects, Spiders and Pathogens: Friends of the Rice Farmers. IRRI, Los Baños, Laguna, Philippines 127 p. Sutherland WJ. 1995. Introduction and principle of ecological management. p. 1-21. In: Sutherland WJ and DA Hill (eds.). Managing Habitats for Conservation. Cambridge University Press, Cambridge England. Wilby A, Heong KL, Huyen NP, Quang NH, Minh NV, Thomas MB. 2006. Arthropod diversity and community structure in relation to land use in the Mekong Delta, Vietnam. Ecosystems 1, 9(4), 538-49.