Key to the larval stages of common
Odonata of Hindu Kush Himalaya, with short notes on habitats and ecology
Hasko Nesemann 1, Ram Devi
Tachamo Shah 2 & Deep Narayan Shah 3
1 Centre for Environmental Science, Central University
of Bihar, BIT Campus, Patna, Bihar 800014, India
2 Hindu Kush Himalayan Benthological Society, Kausaltar,
Nepal. P.O. Box: 20791, Sundhara, Kathmandu, Nepal
3 Senckenberg Research Institutes and Natural History
Museums, Department of Limnology and Nature Conservation, Clamecystrasse 12,
D-63571, Gelnhausen, Germany.
Email: 1 hnesemann2000@yahoo.co.in, 2 ramdevishah@hkhbenso.org (corresponding
author), 3 Deep-Narayan.Shah@senckenberg.de
Date of publication (online): 26 September 2011
Date of publication (print): 26
September 2011
ISSN 0974-7907 (online) | 0974-7893
(print)
Editor: K.A. Subramanian
Manuscript
details:
Ms # o2759
Received 11
April 2011
Final received
22 July 2011
Finally
accepted 11 August 2011
Citation: Nesemann, H., R.D.T. Shah & D.N. Shah
(2011). Key to the larval stages of common Odonata of Hindu Kush Himalaya, with
short notes on habitats and ecology. Journal of Threatened Taxa 3(9):
2045–2060.
Copyright: © Hasko Nesemann, Ram Devi Tachamo Shah
& Deep Narayan Shah 2011. Creative Commons Attribution 3.0 Unported
License. JoTT allows unrestricted use of this article in any medium for
non-profit purposes, reproduction and distribution by providing adequate credit
to the authors and the source of publication.
Author Details:Hasko
Nesemann, Ram Devi Tachamo Shah & Deep Narayan Shahare aquatic ecologists. They are specialized in aquatic macroinvertebrates diversity with a keen interest
in freshwater ecology, biogeography, conservation, and ecological water quality
monitoring.
Author
Contribution: HN, RDTS and
DNS conducted fieldworks and equally contributed in manuscript preparation. HN
illustrated the specimens.
Acknowledgements: We want to thank Subodh Sharma (Aquatic
Ecology Centre, Kathmandu University, Dhulikhel, Kavre, Nepal), Gopal Sharma
(Zoological Survey of India, Gangetic Plains Regional Station, Patna, India),
and R.K. Sinha (Centre for Environmental Science, Central University of Bihar,
Patna, India) for their help in fieldwork.
Abstract: The order Odonata is one of the most
widely studied groups among insects from the oriental region. They colonize in
both stagnant and running water bodies of wide water quality. Hitherto, the existing literature on
the Odonata contained numerous publications with coloured figures of adults,
helpful for identification. Identification key with figures on larval stages,
using their coloration as distinguishing characters are largely missing. The
current work attempts to provide an identification key to aquatic larvae of the
most common families of Zygoptera, Anisoptera and Anisozygoptera with colour
illustrations. The specimens were collected from Nepal and India (northern
part). Each family is represented by several examples to demonstrate the range
of morphological variability. This key helps determination of aquatic larvae
Odonata up to family level without enormous efforts in field and laboratory.
Keywords: Aquatic insect, damselfly, dragonfly,
ecology, identification key, India, Nepal.
For figures, images, tables --
click here
INTRODUCTION
The
modern order Odonata is highly diversified with 5,680–5,747 (accepted)
extant species, 864 (accepted) extant subspecies and approximately 600 fossil
species (Xylander & Günther 2003; Kalman et al. 2008; van Tol 2008). The highest species number is known
from the Oriental region which has more than 1,000 species. From India, exactly 499 species were
recorded until 2005 by Mitra and 463 species confirmed by Subramanian
(2009). Among all the species and
subspecies within this geographical limit, the figure or description is known
only for 78 taxa (Mitra 2005). For
Nepal the number of species and subspecies was previously 172 published by Vick
(1989). Later Sharma (1998) listed
202 taxa and Kemp & Butler (2001) added a new species for the country. In Bhutan, Mitra (2006) has published
an actualized Odonata list with 31 taxa, to which the occurrence of Epiophlebia laidlawi around Thimpu can be added (Brockhaus
& Hartmann 2009).
The
taxonomy and knowledge of odonates in the Indian subcontinent and in many other
parts of the world is largely based on terrestrial adults. There has been an old tradition in publication
of very high quality colour figures for each species since the 18th century
(Malz & Schröder 1979). In
recent years all known Odonata species from the Japanese Archipelago were
published by Okudaira et al. (2005) giving colour figures of both the larvae
and the adults.
Mitra
(2003) has provided an updated list of the regional species composition for the
different ecoregions of the Indian subcontinent. It allows recognition of the local fauna and the possible
presence of their aquatic larvae for the Himalayan region. In contrast, the distinction of aquatic
odonates from the same territory is poorly known. Even the identification at the family level remains
difficult for many Zygoptera (Superfamilies Coenagrionoidea, Lestoidea) and
some Anisoptera (Libellulidae vs. Corduliidae).
The
classification of the order Odonata at the family level is a matter of
controversy/ discussion. The
number of families recognized by different authors varies largely. The 15 families in St. Quentin &
Beier (1967), 27 families in Trueman & Rowe (2001), 56 families in Xylander
& Günther (2003) demonstrate the different views. The present study follows the proposed system of Kalkman et
al. (2008) with one addition.
The
Odonata represents 7% among a total of 76,000 freshwater insect species of the
world (Balian et al. 2008). Many
species have small distributional ranges, and are habitat specialists;
including inhabitants of alpine mountain bogs, seepage areas in tropical rain
forests, and waterfalls (Kalkman et al. 2008). Larvae are mostly aquatic and predatory in nature. They feed on small odonates,
oligochaetes, chironomids, bettles, bugs, mayflies, molluscs, even tadpoles and
small fishes, thus playing a major role in the aquatic ecosystem. The Odonata richness alone occupies a
major component in freshwater macroinvertebrate assemblages. This aspect is clearly shown in Fig. 1
based on data sets of 250 macroinvertebrate samples from various studies (Shah
2007; Tachamo 2007; Nesemann 2009; Tachamo 2010).
The
order Odonata is an ideal model taxon for the investigation of the impact of
environmental warming and climate change due to its tropical evolutionary
history and adaptations to temperate climates (Hassall & Thompson 2008). Its assemblages are also considered as
surrogates for the insect community structure in water bodies, being capable of
indicating changes in the biological integrity of these ecosystems (Silva et
al. 2010). This can be proven from
the newly developed HKHbios scoring (Ofenböck et al. 2010) list for the Hindu
Kush Himalayan river system (Fig. 2). The Odonata at family level alone occupy about 11% of the scoring list
with tolerance scores ranging from 5 to 10.
The
objective of the present study is to fill the gap in the knowledge of the
odonata larvae and to provide a pictorial catalogue to help in their
identification. Here 31 examples
from recent collections are presented to give their morphological characters as
well as live colour. Colour was studied in living materials.
Identification characters of Odonata
larvae
Srivastava (1990) highlighted that the aquatic phase
of the life cycle comprises eggs, pro-larval and larval stages, and 70–95
% of the whole life span is passed in water. Larvae undergo approximately 10–20 molts (mostly
11–14), over a period of three months (e.g. some Libellulidae) and about
6–10 years (e.g. Epiophlebiidae) depending on the species. One characteristic shared by all
Odonata larvae is the conspicuous grasping labium (mask) (Fig. 3 a–c),
used for capturing the prey. At
rest stage, the labium is held folded underneath the head. During prey-capture, the labium is shot
rapidly forward and the prey is grasped with paired hand-like lateral lobes
(palps). Form, size and number of mental setae can be used for family or even
genus identification but requires a microscope. Even from the above characters and with mask retracted,
identification of larvae to suborder and family is very easy, based on several
other features. These are namely the apices of abdomen, number and form of
caudal gills, presence of abdominal gills, form, size and number of segments of
antennae, presence of teeth along the anterior margin of the lateral lobes
(palps) of labium (mask) and anal pyramid with length relationship of epiproct,
paraprocts and cerci (Fig. 4 a–b).
The identification of larvae (nymphs)
even to genus, is often difficult because of the fact that morphological
differences are so slight (Pennak 1978, p. 557). Therefore, keys must be used
with great care.
The
identification of the collected and figured specimens was mainly based on
descriptions given for the Odonata fauna of Japan (Kawai 2005; Okudaira et al.
2005), Malaysia (Yule & Hoi Sen 2004), and a few available publications
from the western Himalayan region (Kumar 1973; Mitra 2005). The identification result reached in
the present study remains mostly at family level. Only in a few cases the genus or species level could be
reached.
MATERIALS AND METHODS
Study area
The study was carried out in various parts of Nepal and the northern part of India (Fig. 5) between 2005 and 2009. The climate in the region varies from humid sub-tropical to temperate with hot summers from March to early June, the monsoon season from mid-June to September and winter from November to February. There is a dominance of monsoon rainfall pattern with maximum precipitation in the summer. The region is one of the most fertile and densely populated regions of the world.
All
the illustrated specimens are with the authors’ personal collection.
Illustrated catalogue
Zygoptera: Chlorocyphidae
The
medium-sized larvae (Fig. 6) have two forceps-like caudal gills which are
triangular in cross section. They
inhabit unpolluted, fast running streams and rivers (Fraser 1919a; Kumar &
Prasad 1977). Chlorocyphidae occur
from tropical Africa to Australia with the highest diversity in the Oriental
region (Kalkman et al. 2008). The
family is represented with 21 species in India (Subramanian 2009) and five have
been recorded from Nepal (Sharma 1998) based on adults.
Euphaeidae
The
larvae (Image 1 a–b) are medium-sized to large and robust with
stonefly-like, flattened body form. They have three very large caudal gills that are saccoid. In addition, there are filamentous gills
on the underside of abdominal segments II-VIII that are light grey-blue and
un-pigmented (Image 1b). These
characters allow easy identification of the family in the field.
Euphaeidae
(and some related families) are distributed from the Mediterranean in the west
to Japan in the east. These
pollution-sensitive larvae are highly specialized on lotic microhabitats. They are locally common in fast running
streams and smaller rivers of the Himalayan middle mountains. They prefer unpolluted waters with low
organic load. Usually they are
found on the underside of large stones in high water current of riffles and
rapids together with large stoneflies of the family Perlidae. Earlier Euphaeidae were often united
with other similar forms as families Polythoridae and Epallagidae (Xylander
& Günther 2003).
Calopterygidae
The
family is also known as broad-winged Damselflies (Image 2). The larvae of the family has a shorter
middle gill than lateral gills that are triangular in cross section without
visible veins. Prementum is
diamond shaped with deep median cleft. Palpal lobes are deprived of setae. First segment of antenna is longer than or equal to the
combined length of the remaining antennal segments. The body size ranges between 30–40 mm.
The
family has cosmopolitan distribution and contains 171 species worldwide. They
are most often found at the edge of streams with slow flowing water. In the
study area, they were clinging to root masses and overhung on twigs.
Synlestidae
The
caudal gills of Synlestidae (Image 3) are short, broad and leaf like rounded,
with oval apices and a smaller median lobe. Prementum or palps do not hold any setae. The mentum is deeply cleft. The palpal lobes have a long moveable
hook and two robust spines. The
adults are large, metallic green or bronze-black damselflies inhabiting in
forested streams. A distinct
‘breaking joint’ or area of weakness occurs at the base of each caudal
gill. In our study, Synlestidae
occurred in pristine rocky mountain streams at an elevation of 1600m.
Amphipterygidae
(including: Philogangidae)
The deeply pigmented aquatic larvae (Image 4) are
typical running water species which have a flattened body and bear long
7-segmented antennae. They may be
larger than other damselflies and have a stonefly-like appearance. The palpal lobes of the labium have
three spines and one movable hook. Their long gills are of saccoid type. They are rare in the Himalayan region with a few scattered
records from Nepal (Kemp & Butler 2001) and northeastern India from
Darjeeling to Assam and Meghalaya (Prasad & Varshney 1995).
This family includes around 10–12 species of
4–5 genera in the tropical and oriental region. They share generally plesiomorhic characters and might be an
ancient relict line within the Zygoptera (Dudgeon 1999; Kalkman et al. 2008,
2010). The genus Philoganga is often separated
as subfamily or family (Subramanian 2009).
Platystictidae
The
larvae of the Platystictidae family (Image 5) possess more or less saccoid
gills as in Euphaeidae, but do not bear any abdominal gills. The palpal lobes of the labium consist
one spine and one movable hook. The colour pattern of the body is pale and
somewhat spindly with large bulbous eyes.
The
Platystictidae are widespread in South Asia to Southeast Asia (New Guinea) and
are also known from central America and the northern part of South
America. The larvae are found in
small forested streams. Around 191–213 species are known worldwide
(Kalkman et al. 2008; van Tol 2008).
Protoneuridae
The
larvae of this family (Image 6) have two-segmented leaf-like caudal gills of
similar shape and length. The gills are clearly divided into a thickened dark
proximal portion and a thin, paler distal part. The anterolateral margins of the labial mentum are fringed
with tiny teeth. One premental
seta is situated on either side of the midline of the mentum. There are three setae on the palpal
lobes. This family has delicate
aquatic larvae with flattened bodies and relatively long antennae; the long,
slender legs are fringed with setae. The posterior margin of the head forms two lateral horn-like extensions,
whereas it is smoothly rounded in Coenagrionidae.
Protoneuridae
have a wide distribution in tropical and subtropical zones but they are
insufficiently known and not generally recognized as family by traditional
odonatology. Protoneuridae inhabit
in a narrow range of slowly running and stagnant waters. The most abundant fauna is found in
wetlands and lentic zones of rivers and streams in lowlands and plains. In the study area, they occurred
numerously together with Coenagrionidae in submerged macrophytes of ponds,
reservoirs and lakes. The general
color appearance of the observed larval forms is uniform light yellow brown. The identification of genus- or species
level is almost impossible due to the high number of taxa with completely
unknown larvae.
Platycnemididae
The
caudal gills of the larvae are very long, their length is approximately the
same as the abdomen with apices somewhat pointed or attenuated and inconspicuous
tracheal branching (Image 7). The
gills are not usually clearly divided into proximal and distal portions. The third segment of antenna is
slightly longer than the second. The anterolateral margins of the labial mentum are not toothed.
The
records of larvae in the study area are rare. The family occurs at elevations ranging from about 200m to
1900m. The figured specimen (Image
7) was found in the littoral section of Jagadishpur reservoir (197m).
Coenagrionidae
(Synonym: Agrionidae)
The
larvae have leaf-like caudal gills of similar shape and length. The gills are not usually distinctly
divided into proximal and distal portions. The caudal gills are shorter than the abdomen, with rounded
apices and conspicuous tracheal branching. The anterolateral margins of the labial mentum are not
toothed and 3-5 premental setae are usually situated on either side of the
midline of the mentum. The third
segment of antenna is shorter than the second.
This
family has the highest species number among all Zygoptera with 1,080 taxa and a
worldwide distribution. Coenagrionidae (Image 8a–d) inhabit a wide range of running and
stagnant waters; the most diversified fauna is found in wetlands and lentic
zones of rivers and streams. In
the study area, they occurred numerously together with Libellulidae in
submerged macrophytes of ponds, reservoirs and lakes. The general color appearance included light yellow brown
forms, dark striped forms, and bright green to dark brown forms. The distinction between Coenagrionidae
and Protoneuridae is very easy based on the form of head and color but the
identification of genus or species level is almost impossible due to the high
number of taxa with completely unknown larvae.
Anisoptera: Gomphidae
The
general body shape of Gomphidae (Image 9 a–b) is compact, and elongate
with an ovate dorso-ventrally flattened abdomen. The legs and often the whole larval body is covered with
various types of hairs, setae, and spines. The antennae are four-segmented with the third segment
enlarged. The tarsi of the first
two pairs of legs are two-segmented. The labial mentum is more or less quadrate and the anterior margin of
labial mentum is never cleft.
The
larvae mostly inhabit running waters and are highly diversified in lowlands at
floodplains of large rivers. Worldwide there are more than 966 species known. All Gomphidae are burrowers in
sediment. The larvae process various morphological adaptations to different
sediment types. Despite their
burrowing lifestyle some Gomphidae are very good swimmers too.
The
larval body of sand and silt (Psammopelal, Pelal) burrower is covered with fine
hairs. Living specimens have attracting light greenish colour (Image 9a). They occur in moderately polluted water
bodies. In case of fine gravel (akal) burrower, only the legs are covered with
fine hairs. The third antenna
segment is broadened spooned-shaped (Image 9b). Living specimens have yellow-orange brownish colour. They occur in non/slightly and
moderately polluted river stretches. The figured specimen (Image 9b) was found
in the same habitat of Aphelocheirus spp. (Heteroptera: Nepomorpha:
Aphelocheiridae).
Lindeniinae
This
subfamily (Prasad & Varshney 1995, p. 403) or family (Hawking & Theischinger
1999, p. 25) (Image 10) comprises the genera Sieboldius, Ictinogomphus and Gomphidia in
Asia and Australia. Kalkman et
al., (2008) does not include Lindeniinae (or Lindeniidae) as a separate
taxon. The larvae are very large
and robust with circular flattened abdomen. Previously they were placed into the family Gomphidae, but
differ in several characters and life style. The labium is enlarged and much broader than in
Gomphidae. The colour of the body
is dark ochre-brown.
Lindeniinae
larvae are not sediment-inhabiting; they are exclusive climbers on submerged
macrophytes. They colonize large
stagnant water bodies and slowly running rivers from lowlands up to 800m. Larva was found climbing on submerged
macrophytes in a lentic zone of Metapotamon-type (large river). They were recorded in a moderately
polluted water body. It is locally abundant in floating macrophytes, found in
Nepal (Phewa Tal wetlands, Begnas Tal effluent) and India (Jharkhand, upper Subernarekha
River and Maharashtra, Tahoba wetland), preferring Eichhornia crassipes as
substrate.
Lindeniidae
were already separated from the majority of Gomphidae on subfamily level as
Hageniinae by several authors (St. Quentin & Beier 1968, p. 8). More recent publications raise them to
family level (Xylander & Günther 2003, p. 141).
Aeshnidae
Aeshnidae
larvae (Image 11 a–d) are the largest among odonata reaching more than
5cm length. The larvae are rather
elongated with a robust, cylindrical abdomen and very large eyes. The antennae are six or seven-segmented
and filamentous. The tarsi of all
legs have three segments. The
labial mentum is widest in the distal portion and narrowing towards the
posterior part with a cleft in the anterior margin. The body surface of the larvae is smooth, without any hairs,
setae or bristles. The larval
colour display a wide range from light yellow, bright green, ochre brown to
dark brownish often with segmentally arranged dark patterns on the dorsal side
of the abdomen.
Within
the family Aeshnidae, the subfamily Anactinae is mainly confined to the
Ethiopian and Oriental regions with range extension of some species into the
temperate Palearctic. In the
Indian subcontinent, they are found sporadically in various undisturbed,
natural, slightly and moderately polluted waters. They are nowhere abundant or common and only small numbers
of individuals were observed.
Cordulegastridae
The
body of Cordulegastridae larvae (Image 12) is elongate and covered with
bristles or tufts of setae. The distal margin of the palpal lobes of labium is
with large irregular teeth which interlock with those on the corresponding
lobe. The anterior margin of the
mentum is cleft. The colour appearance is dominated by a dark brown background
with some blackish markings, regularly arranged on the dorsal side of the
abdominal segments.
The
family has a limited distribution range in the Palearctic and Oriental
regions. The larvae are crawlers
on sand and muddy sediments of fast running cool streams and rivers, especially
in the Himalaya. They usually lay half buried in the surface sediment layer and
wait for prey. The larvae are
pollution-sensitive and demand highly oxygenated water. They are not common and were recorded
during the present study only from the upper stretches of small rivers and
streams of natural forests above 1500m.
Macromiidae
The
legs of Macromiidae (Image 13) are very long, giving the larvae a “spidery”
appearance. The abdomen is depressed and more or less circular in outline. On the head, a small “horn” is present
between the antennal bases. The
labium bears rather long, regular teeth along the distal margins of the palpal
lobes.
The
family has a worldwide distribution but their occurrence is restricted in the
tropical, subtropical and warm temperate zones except South America. There are approximately 120 species
known. They prefer running water
with low organic input and are found in slightly to moderately polluted
stretches. A few Macromia and Epophthalmiaspecies are recorded from northern India and Nepal (Sharma 1998; Mitra
2003). The Macromiidae are
recently raised to family level, previously they were placed as subfamily
Epophthalmiinae into family Corduliidae.
They
are frequently recorded from the upper regions of undisturbed forest streams in
the Himalayan middle mountains from 800 to 1970 m. The figured specimen might belong to Macromia moorei moorei, which is spread widely over the northern
Indian subcontinent. The larvae
occur on coarse-grained sand or gravel substrate (Psammal, Akal) deposited
behind or under large stones.
Corduliidae
The
larvae of Corduliidae (Image 14 a–b) resemble Libellulidae, but their
size is usually larger and the body is more firm than the latter ones. Their legs are rather short and the apex
of the femur does not extend beyond abdominal segment VIII. The abdomen is not markedly depressed
or circular in outline. The cerci are generally more than one-half as long as
paraprocts.
The
total number of species is 255 worldwide; in Asia the family is less
represented. Historically, there
was no clear distinction between the three families Libellulidae, Corduliidae
and Macromiidae. They all were
placed into a single family Libellulidae. More recently fundamental characters of the anal pyramid allow
distinguishing larvae. In
Corduliidae, the length of cerci exceeds always more than half as long as
epiproct, whereas in Libellulidae the length of cerci is less than half as long
as epiproct (Okudaira et al. 2005, p. 360). They were rarely collected in the study area from slowly
running stretches of stream and river with moderate to heavy pollution. It is not possible to recognize and
separate them in the field from Libellulidae; proper identification can be only
done in a laboratory with a microscope.
Libellulidae
The
larvae (Image 15 a–g) are minute to medium-sized and have a delicate
comparatively soft body. Their legs are rather short and the apex of the femur
does not extend beyond abdominal segment VIII. The abdomen is not markedly depressed or circular in
outline. The cerci generally are
not more than one-half as long as paraprocts.
The
family Libellulidae, the largest family of Anisoptera has a cosmopolitan
distribution with more than 970–1,012 described species. The larvae are very similar in
appearance and shape to Corduliidae but differ by their anal pyramid. In Libellulidae, the length of the
cerci is less than half as long as epiproct. Body colour of the different
species may cover a wide range from bright yellow, light greenish to dark
brown. Larvae are usually very abundant in all types of stagnant waters and are
able to colonize successfully even in small water bodies with low oxygen where
other odonates cannot survive.
Anisozygoptera:
Epiophlebiidae
The
larvae are somewhat slender and elongate; with a slight petiolation at the base
of the wing pad. The minute and very short antennae are with five
segments. The larval body is very
hard and firm covered with tubercles, but lacking any bristles. The family is extremely rare with isolated
discontinuous relict distribution in Japan and the Himalaya only. The family is certainly recorded from
Mesozoic onwards (Nel & Jarzembowski 1996).
There
are only two extant species, regarded as ‘living fossils’. Epiophlebia superstes are recorded only from Japan while Epiophlebia laidlawi are recorded from the Himalayan regions
of Bhutan, India and Nepal. The
life cycle of the Epiophlebia
superstes is
better known, including adults, terrestrial phase, and egg deposition; adults
of E. laidlawi are not yet found. The larvae are limited on natural upper
regions of fast running forest streams with good water quality. Small larvae prefer rapids and riffles
with embedded stream bottom; they are highly pollution-sensitive and live only
in Epirhithron- to Metarhithron-type of biocoenotic zone (Nesemann et al. 2008,
2011).
The
young larvae (Image 16a) differ markedly in dorsal colour, having dark
pigmentation only on abdominal segments 2 to 5, and 9. Large larvae have generally brownish or
nearly blackish appearance with dorsal metameric pattern on abdomen (Image
16b). The distinguishing of male and female individuals by the presence of
ovipositor is only possible for larger larvae from 8mm body length onwards
(Nesemann et al. 2008, 2011).
TAXONOMIC KEY
The
order Odonata can be divided into two distinct groups or suborders: Damselflies
(Zygoptera) and Dragonflies (Anisoptera).
Damselflies
larvae are usually more slender than dragonflies and their abdomen terminates
in three caudal filaments (gills) resembling leaves. Dragonflies larvae are much more robust with an abdomen
terminating in five points consisting of a pair of cerci, a pair of paraprocts,
and a single epiproct. In both
damselflies and dragonflies, the shape of the lower lip (labium) can be a
diagnostic character for separating families. The shape of antennal segments is
also an important character in identification of odonates.
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