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Chapter 23
INSECTA : HEMIPTERA (AQUATIC BUGS)
1
ICAR-NBAIR, H.A. Farm post, Bellary road, Bangalore-560024
2
Survey of India, Southern Regional Centre, Santhome High Road, Chennai-600028
SRIMOYEE BASU1 AND K.A. SUBRAMANIAN2
ABSTRACT
India is represented by 325 species under 84 genera and 18 families of freshwater
hemipterans. The Western Ghats and Central India are well explored, whereas,
the Eastern Himalayan States and Western Himalayan regions of India are still
unexplored. Systematic identication of these aquatic bugs through morphological
studies has progressed rapidly. But, molecular identication involving DNA
barcoding is still lacking in India. An intensive eld survey is urgently required
to update the baseline databases of freshwater hemipterans from India and to
understand their regional diversity and distribution pattern in inland freshwater
ecosystems.
Keywords: Hemiptera, Freshwater, India, Diversity, Distribution, Systematics
INTRODUCTION
Freshwater Hemipterans, commonly known as ‘aquatic bugs’, are secondarily
adapted to aquatic ecosystems. However, their nymphal and adult stages are
fully aquatic. The aquatic bugs, belonging to the suborder Heteroptera (true bugs)
are encompassed under three Infraorders such as, Nepomorpha, Gerromorpha
and Leptopodomorpha based on their niche and habitat preferences. They are an
integral component of freshwater ecosystems and play a major role in the food web
of the ecosystem as being predators, scavengers or collectors. The evolutionary
history shows that all families were present in the Mesozoic and probably date back
into the Triassic (Popov, 1971). The Nepomorphan has an extensive fossil record,
while the Gerromorphan have less fossil record in comparison to Nepomorpha.
Gerromorpha is a sister group to the Panheteroptera including Nepomorpha and
Leptopodomorpha (Schuh and Slater, 1995) and all members of the aquatic bugs
are predicted to be at least of the same age.
Nepomorphans are the true aquatic bugs, living underwater. Gerromorphans
are semi-aquatic bugs, living on the surface water, whereas Leptopodomorphans
are the shore dwellers, inhabiting the littoral zones or intertidal zones. They are
hemimetabolous insects and both nymphs and adults of these bugs play a major
role in sustainable management of aquatic habitat. These aquatic bugs are found
in both lotic and lentic freshwater habitats ranging from mountainous lake,
forested pools, ponds, hill streams, rivers, waterfalls, water splashed rocks, hot
springs, cascades etc. Because of their habitat specicity and sensitivity, they are
BASU and SUBRAMANIAN
358
indicators of the changes in the freshwater ecosystem’s health and ideal candidates
for biomonitoring. Their size vary from 1 mm to about 100mm. They possess a
specialized beak-like structure, rostrum, which helps in piercing and sucking.
Besides, the poor dispersal capabilities of these insects also serve as
zoogeographical indicators (Jordon, 1951; Hungerford and Matsuda, 1958a, b;
Thirumalai, 1999) and they are important indicators of long-term environmental
changes. Knowledge on the diversity and distribution of this group helps in
understanding the functional aspects of community structure and provides the
baseline data required for developing a strategy for the effective conservation
and management of the freshwater ecosystem. They have tackled the constraints
of living in aquatic environments by possessing several morphological and
physiological adaptations. Some of them (Corixidae, Nepidae, Geridae etc.) are
efcient bio-control agents of harmful vector like mosquito larvae. Nepomorpha,
Gerromorpha and Leptopodomorpha colonize new habitats as they are migratory
in nature and many species are attracted to light (Fernando, 1961). However, they
are encountered mostly in apterous forms. But, population living in temporary
pools consists of macropterous forms.
Fossil History
The evolutionary history of aquatic Hemiptera dates back into the Mesozoic
and Triassicera. However, the oldest fossil of Gerromorphan bug is Karanabis
kiritschenkoi Bekker-Migdisova, 1962 from the Upper Jurassic which belongs
to Mesoveliidae (Popov and Bechly, 2007). However, the rich fossil record of
Nepomorpha in the Mesozoic creates a mystery on the habitat preferences of
Gerromorpha due to the scarcity of Mesozoic Gerromorphan fossil (Grimaldi and
Engel, 2005).
Historical Resume
The Heteroptera or true bugs are the largest and most diverse groups of insects.
The most comprehensive work on this group, until now, is ‘The Biology of
Heteroptera’ by Miller (1956, 1971). The taxonomic studies on aquatic Hemiptera
was undertaken by Linnaeus (1758) in his 10th edition of Systema Naturae, which
included 12 species from Neotropical, Afrotropical and Palaearctic regions. The
earliest volume dealing with this group was Fabricius’s Systema Rhyngotorum
(1775). This was followed by Amyot and Serville (1843) in Histoire naturelle des
insectes hemipteres. Stal (1868-1869) and (1870-1875) described 66 species of
aquatic Hemiptera. Schuh and Slater (1995) discussed the systematic of these
bugs. Reuter (1912) discussed and analyzed about their synonymy.
The Gerromorpha or semi-aquatic bugs are dened by the members having the
ability to traverse the water surface lm accompanied by four long and segmented
antennae. Earlier, they were treated under the group Amphibicorisae (Dufour,
1833; Leston et al., 1954). But, recent treatment by Schuh and Slater (1995);
Andersen (1982); Polhemus and Polhemus (2007, 2008); Henry (2009) showed that
it consists of eight families placed among four super families. Among the 8 families,
Hermatobatidae, Macroveliidae and Paraphrynoveliidae are not reported from
Insecta : Hemiptera (Aquatic bugs)
359
GERROMORPHA
Gerridae
Mesoveliidae
Hebridae
Hydrometridae
Veliidae
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NEPOMORPHA
Aphelocheiridae
Helotrephidae
Nepidae
Belostomatidae
Notonectidae
Micronectidae
Corixidae
Naucoridae
Pleidae
Insecta : Hemiptera (Aquatic bugs)
361
NEPOMORPHA
LEPTOPODOMORPHA
Ochteridae
Saldidae
Gelastocoridae
Leptopodidae
India. Most of the information on morphology, classication, phylogeny and bio-
geography is derived from the monographic work of Andersen (1982); Spence and
Andersen (1994). Faunistic knowledge on Gerromorpha is limited to the taxonomic
studies by Distant (1903, 1910a and b), Annandale (1919), Bergroth (1915), Paiva
(1919a and b), Dover (1928), Haz and Mathai (1938), Haz and Riberio (1939),
Haz and Pradhan (1947), Pradhan (1950a, b and 1975), Gupta (1981), Roy et al.
(1988), Ghosh et al. (1989), Polhemus and Starmuhlner (1990), Bal and Basu (1994,
1997, 2000), Chen and Zettel (1999), Thirumalai (1986, 1989, 1992, 1994a and b,
1996, 1999, 2000, 2002), Chandra et al., (2012), Jehamalar and Chandra (2013a, b).
The revisionary work of Andersen (1975, 1980, 1990 and 1993), Den Boer (1965),
BASU and SUBRAMANIAN
362
Hennig (1981), Polhemus and Andersen (1984), Hungerford and Matsuda (1958a
and b, 1960, 1962, 1965), Polhemus (1994), Polhemus and Polhemus (1994, 1995),
Chen and Nieser (1993a and b), Chandra and Jehamalar (2012a), Jehamalar and
Chandra (2013c, d), Basu et al., (2014, 2016) on few genera of Gerridae; Andersen
(1981a, 1983, 1989), Polhemus (1990a), Lundblad (1936), Jehamalar (2015)
on Veliidae; Andersen and Polhemus (1980) on Mesoveliidae; Hungerford and
Evans (1934), Polhemus and Polhemus (1995), Jehamalr and Chandra (2014b) on
Hydrometridae; Andersen (1981b) and Zettel (1998) on Hebridae of the World are
important to understand the present scenario of Indian Gerromorpha.
The true water bugs or Nepomorpha occur all over the world except
Antarctica, with about 2000 species altogether, was rst recognized as a group by
Latreille (1810) under the name Hydrocorisae. The taxa historically were placed
in the Hydrocorisae (Dufour, 1833; Leston et al., 1954) and are characterized by
short antennae that are concealed in part or entirely by the eyes. Taxonomy,
nomenclature, higher classication and phylogeny of Nepomorpha were reviewed
by many workers and important contributions have been made by Popov (1971),
Stys and Kerzhner (1975), Rieger (1976), Stys and Jansson (1988), Mahner (1993),
Andersen (1995), Polhemus et al. (1995) and Hebsgaard et al. (2004). Thirumalai
(2007) published a checklist on the Indian Nepomorpha, which enlisted 153 species
under 34 genera of aquatic bugs found in India. However, the number increased
to 157 species with the discovery of more new species under different families in
recent years in India. The earlier knowledge on Indian aquatic bugs is mainly
limited to taxonomic contributions by Distant (1903, 1906, 1910a and b), Annandale
(1919), Paiva (1919a, b), Dover (1928), Hutchinson (1933, 1940), Haz and Mathai
(1938), Haz and Riberio (1939), Haz and Pradhan (1947) and Pradhan (1950a).
The revisionary work of Lundblad (1933), Chen (1960) and Wroblewski (1960,
1962, 1972) on the genus Micronecta (Micronectidae), Todd (1955) on the genus
Nerthra (Gelastocoridae), Esaki and China (1928), Polhemus (1990b) and Papacek
and Zettel (2003 and 2004) on the family Helotrephidae. Polhemus and Polhemus
(1988), Sites et al., (2011), Basu et al. (2013) on Aphelocheiridae, La Rivers (1971a
and b, 1974) on Naucoridae, Lansbury (1972 and 1973) on the genera Ranatra,
Montonepa and Cercotmetus (Nepidae), Hungerford (1933), Brooks (1951) and
Lansbury (1968) on the genera Notonecta, Anisops, Enithares (Notonectidae)
and Jaczewski (1934), Kormilev (1971), Chandra and Jehamalar (2012b) on the
family Ochteridae of the world are noteworthy to understand the nepomorphan
fauna of India. Consolidated information on the occurrence as well as the present
status of aquatic bugs in India pertains to the works of Bal and Basu (1994, 1997,
2000, 2002, 2003, 2004), Ghosh et al. (1989), Nahar (2004), Papacek and Zettel
(2004), Polhemus and Starmuhlner (1990). Ravishankar and Venkatesan (1988),
Thirumalai (1983, 1986, 1989, 1994a, 1998, 1999, 2001, 2002, 2004 and 2007),
Thirumalai and Raghunathan (1988), Thirumalai and Kumar (2005), Thirumalai
et al. (2004, 2006), Venkatesan and Rao (1980) and Zettel (1997a, b, 1998, 2000
and 2001), Chandra et al., (2012), Jehamalar and Chandra (2013a, b).
The major taxonomic work on the infraorder Leptopodomorpha has been
carried out by Reuter (1912), Drake and Chapman (1952), Schuh et al. (1987),
Cobben (1959, 1980, 1985), Polhemus and Polhemus (2007, 2012), Zettel et al.,
(2013).
Insecta : Hemiptera (Aquatic bugs)
363
Diversity
Polhemus and Polhemus (2008) and Alder and Foottit (2009) mentioned that
approximately 4656 species of aquatic and semi-aquatic bugs belonging to three
infraorders, 20 families and 326 genera inhabit freshwater globally. In addition,
more than 1100 species remain undescribed (Polhemus and Polhemus, 2007). Of
these, 1103 species occur in the Oriental region. Whereas, the species richness
is higher in Neotropical region and it is represented by 1289 species. India is
represented by 318 species under 82 genera and 18 families (Thirumalai, 1999,
2002, 2007). Among these, the families like Veliidae, Gerridae, Corixidae,
Micronectidae, Hebridae, Helotrephidae and Naucoridae are high species rich
families. However the current number of species is 4940 globally. Global freshwater
aquatic Heteroptera diversity is provided in Table 1.
Table 1. Global diversity of freshwater aquatic Heteroptera
(Polhemus and Polhemus, 2008).
Infraorder Family Species
Gerromorpha Gerridae 777
Veliidae 1011
Hebridae 230
Hydrometridae 128
Mesoveliidae 47
Hermatobatidae 12
Macroveliidae 3
Paraphrynoveliidae 2
Nepomorpha Belostomatidae 160
Nepidae 271
Notonectidae 406
Corixidae and Micronectidae 613
Pleidae 38
Aphelocheiridae 84
Naucoridae 404
Ochteridae 69
Gelastocoridae 111
Helotrephidae 181
Potamocoridae 10
Leptopodomorpha Leptopodidae 41
Saldidae 335
Omaniidae 6
Aepophilidae 1
Total 4940
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Classication of Aquatic bugs of Indian Subregion
Infraorder Gerromorpha
Superfamily Gerroidea
Family GERRIDAE
Subfamily Rhagadotarsinae
Subfamily Trepobatinae
Subfamily Gerrinae
Subfamily Eotrechinae
Subfamily Cylindrostethinae
Subfamily Ptilomerinae
Subfamily Halobatinae
Family VELIIDAE
Subfamily Microveliinae
Subfamily Rhagveliinae
Subfamily Haloveliinae
Subfamily Veliinae
Subfamily Perittopinae
Superfamily Mesovelioidea
Family MESOVELIIDAE
Subfamily Mesoveliinae
Superfamily Hydrometroidea
Family HYDROMETRIDAE
Subfamily Hydrometrinae
Superfamily Hebroidea
Family HEBRIDAE
Subfamily Hebrinae
Subfamily Hyrcaninae
Infraorder Nepomorpha
Superfamily Nepoidea
Family NEPIDAE
Subfamily Nepinae
Subfamily Ranatrinae
Family BELOSTOMATIDAE
Subfamily Belostomatinae
Subfamily Lethocerinae
Superfamily Ochteroidea
Family GELASTOCORIDAE
Subfamily Nerthrinae
Family OCHTERIDAE
Superfamily Corixoidea
Family CORIXIDAE
Subfamily Corixinae
Subfamily Cymatinae
Family MICRONECTIDAE
Subfamily Micronectinae
Superfamily Naucoroidea
Family APHELOCHEIRIDAE
Subfamily Aphelocheirinae
Family NAUCORIDAE
Subfamily Cheirochelinae
Subfamily Laccocorinae
Subfamily Naucorinae
Superfamily Notonectoidea
Family NOTONECTIDAE
Subfamily Anisopinae
Subfamily Notonectinae
Superfamily Pleoidea
Family PLEIDAE
Family HELOTREPHIDAE
Subfamily Fischerotrephinae
Subfamily Helotrephinae
Infraorder Leptopodomorpha
Family LEPTOPODIDAE
Family SALDIDAE
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365
Family wise diversity is summarized Table 2.
Table 2. Aquatic bugs diversity across family, genera and species
in the Indian subregion
Sl. No. Infraorder Family No. Genera No. Species
I Gerromorpha
1 Gerridae 26 93
2 Veliidae 13 33
3 Hydrometridae 1 5
4 Hebridae 4 22
5 Mesoveliidae 1 3
II Nepomorpha
6 Aphelocheiridae 1 8
7Nepidae 3 24
8 Naucoridae 720
9Notonectidae 4 32
10 Corixidae 720
11 Micronectidae 2 27
12 Pleidae 1 4
13 Helotrephidae 6 13
14 Ochteridae 1 2
15 Gelastocoridae 1 5
16 Belostomatidae 2 5
III Leptopodomorpha
17 Leptopodidae 2 2
18 Saldidae 2 7
Total 84 325
Distribution
Aquatic and semi-aquatic hemipterans are distributed in a wide range of freshwater
habitats ranging from pond, lake, forested small temporary pool, rivers, hill
streams, canal within agricultural eld, waterfalls, cascades, mountainous lake
or stream etc. They occur on all continents except Antarctica and more diverse
in tropical areas. High diversity of water bugs is found in the pristine habitats
of Eastern Himalaya and Western Ghats. However, many families especially of
the Infraorder Leptopodomorpha are yet to be completely documented. They are
also found in the high altitudinal regions and show variance in diversity across
altitudinal gradients and land use pattern. Hence, they are the ideal candidates
for zoogeographical, ecological and biomonitoring studies. Nevertheless, the
documentation of the updated list of species is still incomplete and many more
species are yet to be described.
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Endemism
A high percentage of tropical endemic species are found. Although several species
are cold adapted and associated with stream or waterfalls with high structural
complexity (Polhemus et al., 1992) Due to poor dispersal capabilities, they serve
as zoogeographical indicators (Thirumalai, 1999). Some species has wide habitat
distribution and few are restricted to either lentic or lotic habitats. The Oriental
realm especially the Indian subregion supports a wide variety of aquatic bugs due
to its complex biogeographic past. Though an intensive study is presently focused
on this area, resulting in a good number of new taxa, which are mostly endemics.
Biology
Aquatic and semi-aquatic bugs mainly inhabit inland freshwater habitats, though
several species are associated with intertidal brackish water and marine habitats.
They are predaceous or scavengers except family Corixidae (detritivore), feeding
on other aquatic insects. Few species of the family Belostomatidae, Aphlocheiridae,
Naucoridae, Nepidae and Gerridae are known to predate on other aquatic insects
of their own size, and even tadpoles, sh, molluscs, frogs etc. by immobilizing
preys. They have sexual dimorphism and males are often smaller than females.
All members of the aquatic bugs migrate to new habitats and some of them are
attracted to light. They mostly encountered in apterous forms. However, the
temporary pools are mainly dominated by macropterous population. Species
diversity is affected with the land use pattern altitude, habitat and microhabitat.
Life cycle
The aquatic Hemipterans are hemimetabolous insects i.e., they undergo incomplete
metamorphosis. The life cycle stages consist of egg, nymph and adult. They produce
one or two or sometimes several generations per year. In general, the overwintered
adults lay eggs in springs, develop in summer and further replicate their cycle.
Though, several studies have been undertaken on life histories of these bugs in
laboratory conditions. Usually, eggs are laid and glued on the rocks and plants
or sometimes on the back of the males (Belostomatidae). The eggs are relatively
tough, hexagonal, oval or spindle shaped with button-like or peg like micropylar
processes. There are ve nymphal instars and the nymphs are morphologically
alike to adults, but without reproductive development. The entire life cycle of
these bugs may take 2 to 3 months (Leong, 1962). In some species life cycle starts
from mid May to July and new generations appear in late August to October and
overwinter until March to April (Ohba and Goodwyn, 2010). However, their life
cycle depends on the temperature and availability of food resources. They usually
found to mate throughout the year, but mating increased during February. They
stridulate to attract their mates. Most of them have well-developed forelegs with
tooth or spines for grasping females. Although, surface ripples also attract mates
(Wilcox, 1969), in a few species of water striders, mating occur on the surface water
and the pair copulate for several hours or even days (Birch et al., 1979). In true
aquatic bugs (Nepomorpha), they complete their entire life cycle including mating
underwater.
Insecta : Hemiptera (Aquatic bugs)
367
Longevity
The life span of adult usually lasts for one to six months. However, it varies
according to species and depends on temperature of the aquatic habitat.
Threats
The major threat is the impact of anthropogenic disturbances on inland freshwater
ecosystems. Habitat destruction, deforestation, rapid urbanization, changes in
the land use pattern such as dam construction, encroachment, siltation, weed
infestation, industrial or pesticidal pollution, over exploitation and electroshing
to catch shes in streams or rivers, which also affects other aquatic insects are
the real threats to these bugs inhabiting in such ecosystems. On the other hand,
introduction of exotic insect species to control weeds causes loss of native insect
population. Climate change and global warming are immediate threats to this
freshwater ecosystem, especially in Tropics.
Conservation and Signicance
The rst and principal actions necessary for conservation of these bugs, are the
restoration and management of inland freshwater habitats through protection and
preservation of bio-resources of the habitats. The effective conservation of species
requires understanding of how and where population occurs in a landscape and
what are the factors affecting this population. One of the major attempts to protect
the aquatic fauna is to set priorities among the several deserving candidates of
aquatic habitats and to nd out the factor responsible for their decline. The major
approaches to conserve should be a combination of ecological, biotechnological and
socio-cultural and legal. The ecological and biotechnological approach involves
rapid identication through taxonomic studies and DNA Barcoding of them and
their different life stages. The awareness creation among local communities and to
involve them in management of freshwater habitats is also required.
Gaps in research
Several species especially of the family Leptopodidae, Saldidae, Helotrephidae,
Micronectidae, Veliidae are still unknown from India. Many freshwater
ecosystems of India are under surveyed and unexplored, mainly the Eastern
Himalayan Biodiversity Hotspots and the Western Himalaya. However, major
work on wetlands of Western Ghats has already been undertaken by Thirumalai
(1986, 1989, 1992, 1994, 2004) and Thirumalai and Kumar (2005). However, the
molecular studies involving DNA barcoding technique is still lacking and initiative
is required. Though, few species were barcoded recently and published at NCBI
databases by Basu and Venkatesan (2016).
Future dimensions
An intensive survey in the unexplored areas of India is required to update the
baseline information and to understand their pattern of distribution.
Combined approaches of both morphological and molecular systematic studies
and phylogenetic analysis are needed.
Studies on impact of several ecological and environmental factors and the
BASU and SUBRAMANIAN
368
impact of anthropogenic disturbances on species of freshwater Hemipterans should
be one of the priorities.
Studies on cryptic species population and to resolve the problem through
molecular study is also obligatory.
ACKNOWLEDGEMENTS
The authors wish to express their deep felt gratitude and thanks to Dr. Kailash
Chandra, Director, Zoological Survey of India, Kolkata, for providing facilities to
complete this work.
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