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Odonata attracted by light – a new topic for myth-busters

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Odonata attracted by light – a new topic for myth-busters

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Six Odonata species were collected during night light trapping on the Mambilla Plateau, Taraba State Nigeria. Being predominantly diurnal insects, odonates captured in light traps have always been considered as an anomaly. The new data initiated an extensive interrogation of all records on Odonata collected near artificial light sources. A total of 415 records (402 published and 13 new) are presented here with a summary of previous discussions and new discussion points. The general conclusion is that odonates are mainly confused by, rather than attracted to the light. New avenues for further research in this field are suggested based on previous important studies undertaken on Odonata morphology and physiology.
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International Dragonfly Fund - Report 43 (2012): 1-52 1
Odonata attracted by light a new topic for myth-busters
Danladi M. Umar1, Milen G. Marinov1, Martin Schorr2, Hazel M. Chapman1,
1School of Biological Sciences, University of Canterbury, Private Bag 4800,
Christchurch 8140, New Zealand
2International Dragonfly Fund e.V. / ÖSTLAP, Schulstr. 7B, 54314 Zerf, Germany
Abstract
Six Odonata species were collected during night light trapping on the Mambilla Pla-
teau, Taraba State Nigeria. Being predominantly diurnal insects, odonates captured
in light traps have always been considered as an anomaly. The new data initiated an
extensive interrogation of all records on Odonata collected near artificial light sources.
A total of 415 records (402 published and 13 new) are presented here with a sum-
mary of previous discussions and new discussion points. The general conclusion is
that odonates are mainly confused by, rather than attracted to the light. New ave-
nues for further research in this field are suggested based on previous important stu-
dies undertaken on Odonata morphology and physiology.
Introduction
Odonates are amongst the oldest of distinct insect orders and supremely visual
animals (Yang & Osorio 1996). They have the largest compound eyes of any insect
and the highest number of omatidia (Corbet 1999). As a result it is not surprising
that their vision has evolved to near perfection. Olberg et al. (2005) proved that odo-
nates, perched or flying, could precisely estimate the size of their potential prey and
the distance to it. Moreover they can aim at a point in front of their prey, so that it is
intercepted with a relatively straight flight trajectory. Mizutani et al. (2003) report on
the so called motion camouflage established in Anax papuensis (Burmeister, 1839)
and possibly used by other species as well. This is achieved when the predator mo-
ves towards the prey in such a manner that projects a stationary image in the prey’s
retina. This aerial manoeuvre involves a precise estimation of prey’s position and
movements.
Visual clues play a crucial role in prey detection by Odonata (Labhart & Nilsson 1995,
Olberg et al. 2000, 2005), as well as in habitat selection (Wildermuth 1994), sexual re-
cognition behaviour (Wildermuth 1998) and any aspects of pre- and postcopulatory
behaviour (Corbet 1999). Moreover several studies support the suggestion that the
2 Odonata attracted at light
polarised light reflected by water provides important information on the quality of
freshwater habitats, not only for odonates, but for number of aquatic insects (Kriska
et al. 2006). These authors found that red and black horizontal plastic reflector sheets
were equally highly attractive to water insects, while yellow and white reflectors were
unattractive. In odonates reflective surfaces can aid the orientation from a distance,
particularly where other cues (e.g. atmospheric humidity, dimension and shape of the
water body, undulation of the water surface, water plants on the surface and on the
shore, temperature and odour) are still ineffective (Bernáth et al. 2002).
In spite of their visual superiority odonates have often been confused and trapped
when their natural environment has been altered by humans. Individuals have been
reportedly attracted by the shiny surface of car roofs (Wildermuth & Horváth 2005),
asphalt (Fraser 1936), solar panels (Horváth et al. 2010), and grave stones (Horváth
et al. 2007). Furthermore, choice experiments have shown that crude oil can be mo-
re attractive to odonates than water (Horváth et al. 1998). Therefore, odonates have
been sampled with various traps usually set up for studies on other animal groups.
Aerial nets attached to large vessels have produced interesting findings on the
migratory species (Holzapfel & Harrell 1968, Wolf et al. 1986, Yoshimoto et al. 1962),
as have ornithological and bow nets (Baccetti et all. 1990; Borisov 2008, 2009; Rinte-
le 1997). Hoess & Rezbanyai-Reser (2005) caught the powerful flier Libelulla quadri-
maculata Linnaeus, 1758 with a pitfall trap. Although very rare, Odonata in pitfall
traps have been reported in other occasions, too (Garcia Ruiz 2005, Mertent et al.
2007, Santos et al. 2007). Dogramaci et al. (2010) collected Zygoptera specimens
using pan traps and on several occasions odonates have been collected with Malaise
traps (Askew et al. 1998; Flint 1996, Richards & Windsor, 2007; Rizali et al. 2002,
Sartor et al. 2009). Glotzhober & Riggs (1998) were able to increase their total catch
of Odonata using a modified Malaise trap.
However, the majority of records of trapped Odonata come from collections around
light sources. The total catch per trap varies between regions globally but range from
single individuals to tens of thousands (Feng et al. 2006). There are conflicting views
about the reasons behind this relatively high trapping success at night. Odonates are
predominantly diurnal species, but the significant number collected near light sour-
ces has led many researchers to interpret this behaviour as some kind of attraction
to the light (e.g. Aubert 1964; Averill 1995).
Odonata larvae have also been collected using underwater light-traps (Aiken 1979,
Dommanget 1991, Engelmann 1973, 1974, Espinosa & Clark 1972, Hungerford et al.
1955, Nikolaeva 2008, Schilling et al. 2009, Weber 1987, Williams et al. 1996). Some
laboratory studies on the orientation of the larvae according to the light rays have
been undertaken as well (Abbott 1926, Riley 1912).
The present paper presents new data on odonates captured from light traps. It
investigates the previous records of all Odonata collected by light sources, presents a
International Dragonfly Fund - Report 43 (2012): 1-52 3
discussion on the probable reasons for this behaviour and suggests future research
to clarify this behaviour.
Material and Methods
Light trapping was conducted on the Mambilla Plateau (approx. 1,500 m a.s.l.) near
the village of Yelwa, Taraba State, Nigeria (Fig. 1). The trapping was a part of a larger
ecological study investigating the effect of land use on benthic invertebrate commu-
nities in tropical highland Nigerian streams. Nine streams were selected in three dif-
fering land uses: three forested, three maize fields and three tea plantations (Fig. 1 and
Table 1). The traps consisted of a light source (12 volts DC flash lamps) and a white plas-
tic tray filled with water. Detergent was added to the water to reduce surface tension.
Figure 1. Situation of light traps on Mambilla Plateau, Taraba State Nigeria.
The trays were placed at about 30cm from the water edge and the light source was
attached at about 30cm above the trays’ rim (Fig. 2). Light traps were set up over-
night in four occasions (07 December 2010, 07 January 2011, 07 February 2011 and
07 March 2011). At this stage only Odonata species were identified using Clausnitzer
& Dijkstra (in press). On collection three characteristics were recorded: species affi-
liation, sex, and state of maturity. The specimens were preserved in low grade etha-
4 Odonata attracted at light
nol which made the final identification difficult. Two specimens were particularly
difficult to identify. One was badly damaged, making identification impossible and
the second was identified with supposition because the species affiliation could not
be confirmed. The state of maturity of specimens posed the greatest challenge becau-
se it was unclear if the lack of body colour pattern was due to age or to the method
Table 1. Locations of all nine light traps set up during the experiment.
Trap ID
Land use
Elevation (m a.s.l.)
Latitude
Longitude
F1
forested
7o 5.808’
11o 3.518’
F2
forested
7o 5.829’
11o 3.352’
F3
forested
7o 5.491’
11o 3.331’
M1
maize field
7o 5.284’
11o 4.536’
M3
maize field
7o 5.269’
11o 4.595’
M3
maize field
7o 5.272’
11o 4.637’
Te1
tea plantation
7o 5.614’
11o 5.568’
Te2
tea plantation
7o 5.560’
11o 5.560’
Te3
tea plantation
7o 5.058’
11o 5.181’
Figure 2. The design of the light traps (picture also shows the primary investigator).
of preservation. Immature specimens were generally identified by the body colour
and intensity of the pruinescence on the abdomen, in combination with the
hardness of the wing membrane. No detailed discussion was made on these results
International Dragonfly Fund - Report 43 (2012): 1-52 5
as some of them were based on the “best guesses” from the identification. The state
of maturity was included in the current paper for general orientation only and should
not be interpreted in future studies.
A review of all known records of odonates coming to artificial light sources was pre-
pared by reviewing the published literature and consulting other experts, acknow-
ledged here. The discussion was performed on the information about Odonata imago
only. Responses of larvae to the light sources must be investigated separately as the
results from the experiments are inconsistent in their findings.
Results
A total of 94 specimens belonging to six odonate species were collected with four of
the light traps from Nigeria. Table 2 represents the total catch per trap and date.
All six species were included in another table containing a total of 415 records of
Odonata coming to light. These records are from the literature or provided by experts
(Table 3, see appendix). The species from Nigeria are recorded for the first time from
light traps, which increases the overall number of odonate taxa recorded from light
sources to 198 (Table 4, see appendix). As the list includes a large number of records
from various parts of the world, a unified approach was preferred while assessing
the current taxonomic species status. With one exception, species were included
according to the World List of Odonata (Schorr & Paulson 2011). Thus some original
published records may not match the entries or may not agree completely with the
taxonomy currently suggested by other authors. Orthetrum camerunense Gambles,
1959 is the only species whose taxonomy differs from Schorr & Paulson (2011),
where it is given as synonym of Orthetrum caffrum (Burmeister, 1839). Here a full
separate species rank is preferred following the unpublished key of Clausnitzer &
Dijkstra (in press) which specifically deals with African Odonata.
Discussion
Odonates are considered predominantly diurnal insects with few species active during
morning and evening twilight (eocrepuscular), and few others flying in evening twi-
light (crepuscular) (Corbet 1999). Adults are sometime visitors of moth traps and have
also been found actively flying near lanterns (Feulner 2007), candles (Pemberton
1995, Wesenberg-Lund 1913), lighted buildings or vessels (Askew et al. 1998, Averill
1995, Campos 1931, Dannreuther 1935, Mitra 1974, Morley 1919, Morton 1932,
Paine 1996b, Platt & Harrison 1994, Schneider 1992, Tani 1998, Walker & Pittaway
1987, Yamane & Hashiguchi 1994, Young 1967), street lights (Askew et al. 1998),
searchlights (Bartenef 1933), flashlights (Honda 2003). This type of behaviour is ge-
nerally reported as “surprising” (Morley 1919), “uncharacteristic” (Yosef 1994) or
“curious” (Young 1967). The prevalent opinion is that these species have been at-
6 Odonata attracted at light
Table 2. Odonata species collected during the light trap experiment in Nigeria.
tracted to light, which infers a preference
for specific wave lengths of the light. In-
deed, comparing the efficacy of different
light sources, Ramamurthy et al. (2010) es-
tablished that mercury lamps were better
than UV and black light sources, providing
more than 70% of the total Odonata catch.
Other research has provided stronger indi-
cations of some species being lured to light.
Borisov (1990) used the term phoxene (in-
troduced by Gornostaev 1984) to denote
animals that move towards artificial light.
He distinguished 8 odonate species obliga-
tory photoxenes and 12 facultative photox-
ens from SW Tajikistan. However, the total
sample sizes were strongly influenced by the
type of light source. Theischinger (2003,
2010) suggested that Austrocnemis maccul-
lochi (Tillyard, 1926) and Austrocordulia re-
fracta Tillyard, 1909 respectively have an
attraction to artificial light. Those two spe-
cies have been collected regularly and in
large numbers near artificial light sources.
Sharma et al. (2000) set up an Odonata light
trap experiment and reported 14 species
with Diplacodes trivialis (Rambur, 1842),
Brachythemis contaminata (Fabricius, 1793)
and Agriocnemis pygmaea (Rambur, 1842)
being the most numerous and therefore
appear to be light-attracted. Parr (2006b)
refers to an observation when Aeshna
grandis (Linnaeus, 1758) was lured in the
light trap just few minutes after its initial
operation. Frost (1971) summarised 12
years of sampling records carried out with
light traps at the Archbold Biological Sta-
tion, Florida. He used data on Pachydiplax
longipennis (Burmeister, 1839) to support
the view that some Odonata are attracted
to light, with 96 specimens having been col-
International Dragonfly Fund - Report 43 (2012): 1-52 7
lected in one month. The majority of the specimens were sampled during calm
nights, thus not disturbed by humans or bad weather conditions.
However, a number of theories have been proposed to explain odonate captures by
light traps. Borisov (2007) outlines three groups of obligatory photoxenes:
Individuals, which during warm periods of the year could embark on activities
during the night such as searching for food, flying to roosting places and copula-
tion.
Species whose adults emerge during the night.
Entirely diurnal species active in high temperature, low humidity and direct sun
light. However, some predominantly teneral individuals arrive to the light sources.
In summary, Borisov (2007) suggested that obligatory photoxenes appear to be indi-
viduals behaving differently from their typical behaviour when they experience ex-
tremes in weather conditions or unusual situations. A summary of the theories ex-
plaining this natural phenomenon is provided below.
Night emergence of imago
Borisov (2004) considered night emergence as a means for regulating mating, avoid-
ing predators and desiccation. Adults emerging during the night have no experience
and could easily be confused by the bright rays of the light source. Corbet (1981)
also considered this phenomenon as important because tenerals performing their
maiden flight have been regularly sampled during the experiments done in Trinidad.
Showers & Horsnail (2001) reported on an exceptional situation where 187 Zygo-
ptera exuviae were collected from a light trap over 13 days, with a maximum of 56
individuals collected during one night. Lindenia tetraphylla (Vander Linden, 1825)
was initially identified as a facultative photoxene (Borisov 1990), but transferred to
the group of obligatory photoxene due to its night emergence (Borisov 2007). How-
ever, not all species whose imago emerge during the night seem to be affected by
the light. Sympecma species typically emerge at night, but are not included amongst
the obligatory phoxene (Borisov 2007).
Escaping from a refuge
Night emergence was used by Mazochin-Porchnakov (1960) to develop a theory ex-
plaining light catches. He suggested that the insects travel to light as they come out
of their night refuge because they could see a way for a free flight. The light indica-
tes to the animal an open area to fly to. Therefore it is not the light itself which at-
tracts the insects, but the opportunity they perceive as a path towards an open area.
In nature open spaces usually better lit and brighter than close-canopied areas.
Insects constantly use bright illumination as an indication of a space (emergence from
a hollow, or from dense vegetation). Borisov (2007) supported this view with exam-
8 Odonata attracted at light
ples from his odonatological studies. He emphasises that the flight towards a light
source could be explained as a movement towards a clear area with UV-light. Cam-
pos (1931) concluded that odonates have been captured in the traps while they have
been searching for sheltered places. The electrical light was found to be of no impor-
tance for them and does not attract these insects.
Prey availability
Borisov (1990) emphasised that odonates may have not been attracted to light, but
to the prey which they may find near the light. Some Anax parthenope (Selys, 1839)
adults, for example, tended to withdraw from the light after being confused and lost
orientation close to the source. Corbet (1999) concluded that it was unclear what
drives an adult to fly towards the light the stimulus of the light itself or the poten-
tial prey assemblages. He included this type of behaviour in a special Strategy A.1.2
that is believed to increase the foraging efficiency. The same views are shared by
other researchers (Sluvko 2007, Richards & Rowe 1994, Wright 1944, Yosef 1994,
Young 1967).
Odonates disturbed near light sources
Corbet (1999) suggested that some individuals caught in light traps may have been
disturbed by the trap operator and inadvertently flown in. Wright (1944) reported
that individuals disturbed by workmen were perching on or near the lights of the cars
and feeding on smaller insects. Furthermore, Dumont (2004) suggested that some
species may end up in light traps after having been disturbed while roosting at night,
and may not normally be night-active at all. Both Dunkle (1978) and Parr (2006b)
came to the conclusion that many of the catches by light traps may simply reflect in-
dividuals accidentally disturbed near the trap. However, only species which in late
evening retain their motion at a certain level, referred to as a period of potential
activity (Borisov 1990), could be forced to move within the study area. Species in a
state of complete inactivity could not be chased off the ground and usually fell to the
ground off the perching substrates. Similarly, Bick (1949) collected Libellula need-
hami Westfall, 1943, A. junius and P. longipennis from their roosting grounds with a
simple flashlight.
Further important discussion points
The summary of the above presented theories was prepared following the new re-
cords from Nigeria, which initiated a large literature review. It could possibly be
expanded in the near future with more data from other publications. Light trapping
is a very popular method in entomology. It has provided similar estimates for the
relative abundance of grasshoppers compared to sweeping technique (Evans et al.
1983), proved to be effective in studying migration patterns of Heteroptera (Benedek
& Jászai 1973), not to mention the key role that lighttrapping plays in the study of
International Dragonfly Fund - Report 43 (2012): 1-52 9
moths (Hausmann 2001). Odonata are often part of the by-catch and as such they
may be reported at order level only. Nevertheless they do present in the samples
collected using light traps. That is why a detailed review of any record of dragonflies
coming to light source must consider every other insect group that has been studied
by this method. Such a review is an overwhelming task and is out of focus of the
present review. However, entomologists are encouraged to add their records (Parr
2006a) in order to achieve a thorough and comprehensive list of Odonata species
collected near the light sources. Every piece of new data could open novel discus-
sions to the one presented below.
The literature review considers six other discussion points, either underestimated or
neglected so far, as important in explaining odonates’ behaviour in relation to artifi-
cial light: a) link between Odonata collected by the light traps to the extreme weather
conditions and migration, b) low proportion of species and individual odonates com-
pare to other insect groups, c) low representation in the samples of crepuscular spe-
cies, d) female dominated sex ratio, e) Zygoptera are much underrepresented in the
light trap samples than Anisoptera, and f) atypical behaviour near light sources.
Extreme weather conditions and odonates migration
Odonates have often been sampled near light sources when forced to move long dis-
tances due to severe change in atmospheric conditions or during their regular an-
nual migration. Al-Houty (1985, 1989) reported on significant numbers of Selysiothe-
mis nigra (Vander Linden, 1825) associated with sandstorms. The individuals were
obviously exhausted and easy to catch by hand, which is atypical for this species. S.
nigra is usually difficult to observe and individuals are hard to catch with a net (M.
Marinov, per. observ.). Corbet (1981) stated that the number of individuals collected
by the light traps was high soon after the onset of the rains, while Ramamurthy et al.
(2010) showed a significant positive correlation with average rainfall and Odonata
catches by the light traps. Furthermore, Anax guttatus (Burmeister, 1839), Anax
ephippiger (Burmeister, 1839) and Gynacantha hyalina Selys, 1882 have been col-
lected during monsoons (Asaithambi & Manickavasagam 2002). A. ephippiger is also
a well known migrant and is the fourth most often collected species from light sour-
ces (Table 4). Averill (1995) reported on a large number of A. ephippiger hovering
around the decks of a lighted ship when the moon was obscured but they did not
occur on cloudless nights. Other species commonly caught in light traps are known
for their long distance dispersal which happens in assemblages of tens of thousands
individuals. Barrois (1896) reported a single observation of about 60,000 individuals
sitting on the electric wires running along roadsides for about 12 km. These were
probably Sympetrum striolatum (Charpentier, 1840) and Sympetrum sanguineum
(Müller, 1764), which rank as the first and fifth most often collected species re-
spectively (Table 4). Furthermore, Aeshna mixta Latreille, 1805, which ranks second, is
a regular migrant bearing the colloquial named Migrant Hawker. Riddiford (1992)
10 Odonata attracted at light
refered to a huge number of A. mixta described by local people as clouds of dragon-
flies on the night preceding light-trapping. Parr (1996) commented on the high num-
ber of reports of migration events involving hundreds of A. mixta individuals. Parr
(2006b) also attributed all UK records of S. sanguineum, S. striolatum and A. mixta
from light traps to the migration mode of those species. Pantala flavescens (Fabri-
cius, 1798), third place in Table 4, is perhaps the best example. Its aerial distapersal
has been witnessed many times with some extraordinary figures. Feng et al. (2006)
reported on the most striking catch ever 42,161 specimens collected with a search-
light trap for one night. They related this massive movement to the nights charac-
terised by fog or slight wind.
Odonates underrepresented in light traps
Despite the high figures presented above odonates usually represent a very low (if
anything at all) proportion of the total number of insects caught per light trap. During
his 30-years life-time of experience with Noctuidae, the lepidopterologist Stoyan
Beshkov has barely found any Odonata in his moth trap. He has been working very
intensively in one of the Europe’s biodiversity hot spots, (the Rhodopes Mountain in
Bulgaria and Greece) with total of 61 Odonata species known so far and at least 3
more expected (Marinov 2004, 2007). Farrow (1984) operated a light trap for 351
nights and collected total of 105,480 specimens with only 19 of them being
odonates. A 12-years period of light-trapping on the Florida Peninsula produced as
many as 200 specimens from 17 species (Frost 1971). These are a small fraction of
the 143 species reported by Frost (1970) for the entire Florida Peninsula. While the
total catch of 200 specimens sounds high, it was obtained over a 12-years period.
During the same period Caenis diminuta Walker, 1853 was represented in the samp-
les with 44,079 specimens collected over only one month (Frost 1963). Moreover, in
some years no odonates were collected at all (Frost 1962). It is true that in this
occasion the observation was done during the winter months, but Odonata are
active on the Peninsula all year around (Chelmick 2005). McGregor et al. (1987)
collected total of 1,126,946 insects from the North Island hill country, New Zealand
with only 12 individual odonates. New Zealand is generally poor in Odonata species
and this is the only record for the country of any imago coming to light.
Low representation of crepuscular and eocrepuscular species
Surprisingly the data indicate that species caught in light traps are mainly known to
have well expressed diurnal activities. However crepuscular (feeding) behaviour is
known of many species from all continents: Reeves (2003) listed 23 crepuscular spe-
cies for Australia; Belevich & Yurchenko (2010) presented observations form Eurasia
for eight crepuscular Aeshnidae; Williams (1937) discussed the situation for the
Americas; Samraoui (1999) provided details for Africa, and Asian crepuscular species
were treated in Arai (1985), Orr (2011), Wilson (2001). In contrast eocrepuscular
International Dragonfly Fund - Report 43 (2012): 1-52 11
species, like T. tillarga and Gynacanthini, have been recorded less often light sources
(Table 4). Corbet (1961) found it interesting that a crepuscular species like Gyna-
cantha villosa Grünberg, 1902 was not collected in light traps which caught four
other Odonate species. Münchberg (1966) interpreted the ocelli at the vertex region
of caput (“frontal eyes”) as "receptors for light of low intensity". “Their arrangement
in space and dimension indicates that they inform also of the direction of the in-
cident light rays. Probably those ocelli play an important role both in crepuscular
activity and attraction by artificial light sources. Adults do need light for orientation
and in twilight they probably use their ocelli in a similar manner as Wellington (1974)
described for bumblebees. In those insects polarised light and ocelli can prolong
foraging, but homing was a problem for bumblebees when they depend on landmarks
and have insufficient light for orienteering. That is perhaps why crepuscular and eo-
crepuscular odonates were very rarely collected with light traps. Those species are
probably able to distinguish between the dim light (their natural environment) and
atypical light source (artificial light).
Female odonates dominate in light traps
Females outnumbering males around the light sources is another interesting disco-
very that has been documented by other authors. Dumont (2004) attributed this fact
to the cryptic behaviour of the female. High male density near the water edge is
commonly observed in odonates, while females visit water significantly less fre-
quently (Corbet 1999). In the present review, females only samples were reported in
57 occasions, while males only were found in 52 samples; females’ prevalence in the
light traps was observed in 17 instances and males were found to be dominant in
nine of the traps. Eleven catches produced equal number for males and females.
This perhaps reflects the location in which the light traps have usually been set up,
that is in forest clearings away from the water edge. Borisov (1990) established that
female Ischnura evansi Morton, 1919 were twice as likely to be trapped as males.
Similarily females of Ischnura fountainei Morton, 1905 were four times as likely to be
caught as males when the trap was not operating close to any wetland.
A significant dominance of Anisoptera over Zygoptera
Tunmore (2005) observed that damselflies do not often appear at the light sources.
Figure 3 presents a summary of the ratio between Zygoptera and Anisoptera repre-
sentatives trapped by the artificial light. The dominance of Anisoptera families (83%)
may be explained with the observed tendency of trapping mainly migratory species
that embark on long distance dispersals. However, Parr (2006b) sought the explana-
tion of this fact in other than biological underlying mechanism because the migra-
tion alone does not explain the much lower records of Zygoptera (17%) coming to
light compare to Anisoptera. As mentioned above Parr (2006b) did not consider the
attraction to light as typical behaviour for this insect order but rather suggested that
12 Odonata attracted at light
most of the captures are due to accidental individuals chased from their temporal
refuges.
Figure 3. Zygoptera versus Anisoptera families recorded from light traps.
Atypical behaviour near light sources
Atypical behaviour near a light source was reported for the first time by Bayford
(1911). On this occasion powerful fliers such as A. cyanea were sitting motionless
and easily picked up by hand. The research suggested that the atypical behaviour
near electric lamps could be triggered by the light source which alters the otherwise
powerful sight of odonates. The light may act in the same manner as it does on hu-
mans staring directly into a light source. Zieba & Buczynski (2007) have observed
Aeshna viridis Eversmann, 1836 individuals sitting head on to the lamp so that their
body axes were forwardly directed to the source of light. They were easy to collect
by hand although the authors do not believe the odonates were completely dazzled
(P. Zieba & P. Buczynski per. comm.). Young (1967) has observed Epitheca princeps
Hagen, 1861 on lighted wall, again easy to pick up by hand and individuals did not fly
away when let back on the substrate. An anomalous behavior of curving the abdo-
men upwards when perched on the wall was observed. This atypical behaviour of E.
princeps was interesting also for the individuals had repeatedly aggregated at a
particular light source although other lamps on the street obviously illuminated the
same wave lengths. The dragonfly persistence was attributed to the physical en-
vironment of the surroundings, like vegetation. A similar relationship between the
vegetation and dragonfly body position at rest was observed by Bartenef (1930). The
position and orientation of the adults was determined by the transmitted sun light
regardless of the cloudiness and the direct exposure to the sun light of the insects’
International Dragonfly Fund - Report 43 (2012): 1-52 13
bodies. Orientation of the adults on defoliated trees was more constant than those
sitting on fully leaved trees. The light transmitted by the leaves obviously has in-
fluenced the orientation of the adults in various directions.
Platt & Harrison (1994) provide an anecdotal observation for the atypical behaviour
which may have been provoked by the effect which light has on odonates’ senses. A
male Anax junius (Drury, 1773) was observed circling around the lamp bulb for seve-
ral minutes. The authors believe that the male may have been responding to the
prismatic spectral rainbows created by the faceted glass edges of the lamp housing.
The green and blue light bands could have created an image of another male. There-
fore the flight around the house lamp could be a behavioural response to confront a
conspecific rival male. Reflected light could act as a very strong stimulus. Hooper et
al. (2006) showed that polarised reflection from a wing surface of Aeshna cyanea
(Müller, 1764) acted as an intraspecific signalling channel. The responses to this
reflection depended on the thickness of the wing membrane and the nature of wax
pruinosity. Wiesenborn (2011) concluded that nitrogen concentration in the insect
exoskeleton appeared to increase as abundances of resilin and other fluorescent,
elastic proteins increased. In P. longipennis these structural compounds are respon-
sible for emitting blue fluorescence in the UV light at the area of wing articulation
and are probably important nitrogen sources for insectivorous vertebrates. Indeed,
Mitra (1974) found that geckoes attack Crocothemis servilia servilia (Drury, 1773)
and B. contaminata, but never Tholymis tillarga (Fabricius, 1798) even when
individuals coming to light are sitting very close to the lizard. This reaction is linked
to the colouration of the T. tillarga wings which superficially resemble hymenopte-
rous insects.
A new theory following the Nigerian data
For the purposes of this research it is important to understand how the Nigerian
data fit into the discussion. The total sample consists of six species belonging to the
same family Libellulidae (Table 2). For light trap Te1 odonates were the only insects
collected during the night. All of the species are very common and known as exclusi-
ve inhabitants of stagnant water bodies. No crepuscular, eocrepuscular or significant
long dispersal activities have been recorded for them. Males totally dominate and as
far as the preservation method allows us to tell, the immature individuals were less
abundant than mature ones. The general conclusion is that the sample is typical of
what one could find during the day along the banks of vegetated pools. However,
the light traps were set up at the sample streams and at first sight all new data do
not comply entirely with what already has been discussed in the literature. No rain
storms or other extreme weather conditions were observed during the time of trap-
ping, no atypical activities were noted around the light traps, odonates totally do-
minated in some light traps, so did males over females. It is unlikely that the indivi-
duals were newly emerged searching for an open space as not any truly teneral
14 Odonata attracted at light
specimens were present in the sample. Those identified as immature possess much
harder wing membrane from what are typically observed for tenerals. It is un-
doubtedly a situation never encountered before which deserves a special attention
in future studies. This finding could be explained with the proximity of an irrigation
pool occurred within 50m from the light trap Te1. It is possible that the effect of the
light in combination with the reflection from the water surface of the tray, which
was reported above as an extremely powerful signal for odonates, had created an
image of a possible new habitat and lured the males for establishing new territories.
It is not difficult to imagine that the warm climate of tropical Nigeria could keep the
individuals in the discussed above situation of period of potential activity. Perhaps
the territorial and predation mode of odonates do not drop so significantly during
the night and they could react to such a strong aerial ecological photopollutant
(after Horváth et al 2009) like bright artificial light sources.
Conclusions and recommendations
All main discussion points presented above are supportive for the view that odona-
tes are probably not attracted, but confused by the light. Those insects are a by-pro-
duct of the total catch per light trap rather than obligatory visitors to light sources.
Even when very large catches were made by light traps it does not necessarily mean
that they have been attracted to the wave length of the source. It seems more likely
that they are caught in light traps due to some combination of other factors. Yamane
& Hashiguchi (1994) report on a large number of P. flavescens that in a fine windless
weather dived into the sea. Some of them were picked up and when released went
to the sea again. In no circumstances could such behaviour be attributable to any
kind of attraction to death.
However, more studies are needed to establish the true nature of the phenomenon
discussed here. Generally light traps are not popular in Odonata studies. Borisov
(1990) and Sharma et al. (2000) are so far the only examples of light trap studies spe-
cifically designed for capturing adult Odonata. People are encouraged to use this
method in odonatological studies as well (Wada & Inoue 1997, Zieba & Buczynski
2007), because light-trapping was found important in studying the phenology and
ecology of the adult Anisoptera (Corbet 1981). The study case from Nigeria is suppor-
tive of this general conclusion. If indeed the adult odonates in the tropical regions
sustain their activity during the night, then the light trapping within those areas is
something that must be considered in the odonatological studies. Moreover there are
already some very important results that have been obtained during occasional
samples using light-trapping method. Holotypes of Melanocacus mungo (Needham,
1940), Hemicordulia gracillima Fraser, 1944, Orthetrum icteromelas cinctifrons Pinhey,
1970, Gynacantha rammohani Mitra & Lahiri, 1975, are known from this type of study.
International Dragonfly Fund - Report 43 (2012): 1-52 15
Future studies in this field must consider two additional points not discussed in de-
tails by previous research: a) Odonata population dynamics and ratio, and b) indivi-
dual species perception of light.
The bulk of the data from light-trap investigations was collected from research done
on migratory species and with just few exceptions, Odonata species and individuals
were far underrepresented in the total catch of insect specimens. This fact could well
be a reflection of the total ratio of adult odonates towards other insect orders in na-
ture. This could be supported only with evidences from population studies involving
large scale insect sampling encompassing species from various orders within the
same investigated area. Such information, is not however available at the moment,
but would make a very interesting study for future analyses.
The insect perception of wave length would be another important part of a study to-
wards clarifying whether odonates are indeed confused by light rather than attract-
ed by it. It is well established that the insect vision varies between the species even
within the same order or family. Yang & Osorio (1996) have found substantial dif-
ferences between the laminas in the eyes of Hemicordulia and Sympetrum, which
may perhaps explain the different responses to light of the representatives of the
Corduliidae and Libellulidae. Similar comparisons need to be made between crepus-
cular/eocrepuscular species versus those with well expressed diurnal flying modes.
Such research is believed to shed some more light on the dilemma discussed here. It
will determine whether the so called attraction to light sources of odonates has its
origin in the morphology of the eyes, is a physiological reaction or purely an acciden-
tal event as inferred here for the majority of species reported so far.
Acknowledgements
The study was supported by the International Dragonfly Fund. Many people contri-
buted with literature, advises and valuable comments at various stages during the
work. Here their names are arranged in alphabetical order: Adrian Parr, Akihiko Sasa-
moto, Attila Kalmár, Bastiaan Kiauta, Elizabeth Graham, Jean-Piere Boudot, Jon Hard-
ing, Jon O’Brien, Jurg De Marmels, Marianne Kiauta, Naoji Katatani, Naoya Ishizawa,
Natalia von Ellenrieder, Pam Taylor, Richards Rowe, Richard Seidenbusch, Sergey Bo-
risov, Shyamkant Talmale, Stephen Richards, Tim Beynon, Viola Clausnitzer.
16 Odonata attracted at light
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28 Odonata attracted at light
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Appendix
Table 3. A summary of Odonata records (published and new data) from light traps.
Counrty
Family
Species
Sex
per
site
Notes
Reference
Japan
Calopterygi-
dae
Atrocalopteryx atrata
(Selys, 1853)
1
Tani (1998)
French
Guyana
Calopterygi-
dae
Hetaerina moribunda
Hagen in Selys, 1853
1
Geijskes (1971)
India
Caloptery-
gidae
Vestalaria smaragdina
(Selys, 1879)
2
Shull & Nadkerny
(1967)
Bots-
wana
Coenagrion-
idae
Agriocnemis exilis
Selys, 1872
Pinhey (1976)
India
Coenagrion-
idae
Agriocnemis pygmaea
(Rambur, 1842)
Sharma et al. (2000)
Ivory
Coast
Coenagrion-
idae
Agriocnemis zerafica
Le Roi, 1915
2 ♀♀
Dumont (2004)
USA
Coenagrion-
idae
Anomalagrion
hastatum (Say, 1839)
Frost (1964)
USA
Coenagrion-
idae
Argia fumipennis
(Burmeister, 1839)
Frost (1975)
Australia
Coenagrion-
idae
Austrocnemis maccul-
lochi (Tillyard, 1926)
55 ♂♂, 22 ♀♀
Theischinger (2003)
China
Coenagrion-
idae
Cecrion sexlineatum
(Selys, 1883)
Easton & Liang
(2000)
China
Coenagrion-
idae
Cecrion sexlineatum
(Selys, 1883)
new for the
country
Wilson (1996)
India
Coenagrion-
idae
Ceriagrion coroman-
delianum (Fabricius,
1798)
occasionally
at the light
source
Andrew & Tembhare
(1997)
India
Coenagrion-
idae
Ceriagrion coroman-
delianum (Fabricius,
1798)
Sharma et al. (2000)
International Dragonfly Fund - Report 43 (2012): 1-52 29
Counrty
Family
Species
Sex
per
site
Notes
Reference
Namibia
Coenagrion-
idae
Ceriagrion suave Ris,
1921
4 ♂♂
Martens et al. (2003)
Russia
Coenagrion-
idae
Coenagrion lanceo-
latum (Selys, 1872)
Kosterin & Dubatolov
(2005)
USA
Coenagrion-
idae
Enallagma cardenium
Selys, 1876
Frost (1975)
USA
Coenagrion-
idae
Enallagma civile
(Hagen, 1861)
1
Wright (1944)
USA
Coenagrion-
idae
Enallagma concisum
Williamson, 1922
Frost (1964)
Russia
Coenagrion-
idae
Enallagma cyathige-
rum (Charpentier,
1840)
Sluvko (2007)
UK
Coenagrion-
idae
Enallagma cyathige-
rum (Charpentier,
1840)
Odin (2006)
UK
Coenagrion-
idae
Enallagma cyathige-
rum (Charpentier,
1840)
Paine (1992b)
UK
Coenagrion-
idae
Enallagma cyathige-
rum (Charpentier,
1840)
Parr (2006b)
UK
Coenagrion-
idae
Enallagma cyathige-
rum (Charpentier,
1840)
1
Parr (2007)
UK
Coenagrion-
idae
Enallagma cyathige-
rum (Charpentier,
1840)
exuviae
Showers & Horsnail
(2001)
UK
Coenagrion-
idae
Enallagma cyathige-
rum (Charpentier,
1840)
5
Tunmore (2005)
USA
Coenagrion-
idae
Enallagma laurenti
Calvert, 1919
Frost (1969)
Russia
Coenagrion-
idae
Erythromma najas
(Hansemann, 1823)
Borisov (2007)
UK
Coenagrion-
idae
Erythromma najas
(Hansemann, 1823)
exuviae
Showers & Horsnail
(2001)
Russia
Coenagrion-
idae
Erythromma viridulum
Charpentier, 1840
Borisov (1990)
UK
Coenagrion-
idae
Erythromma viridulum
Charpentier, 1840
Jones (2004)
UK
Coenagrion-
idae
Erythromma viridulum
Charpentier, 1840
Parr (2006b)
Brazil
Coenagrion-
idae
Homeoura ? nepos
(Selys, 1876)
3 ♂♂, 1
Longfield (1929)
Japan
Coenagrion-
idae
Ischnura asiatica
(Brauer, 1865)
1 , 1
Coll. 27. 07.
2002 at the
Omurasaki
Center, Ho-
kuto City
N. Ishizawa (per.
comm.)
30 Odonata attracted at light
Counrty
Family
Species
Sex
per
site
Notes
Reference
Bulgaria
Coenagrion-
idae
Ischnura elegans
(Vander Linden, 1820)
1
S. Beshkov (per.
observ.)
Poland
Coenagrion-
idae
Ischnura elegans
(Vander Linden, 1820)
1
Buczyński &
Buczyńska (2010)
Russia
Coenagrion-
idae
Ischnura elegans
(Vander Linden, 1820)
11
Maximum
catch for one
night
Borisov (1990)
Tadzhi-
kistan
Coenagrion-
idae
Ischnura elegans
(Vander Linden, 1820)
1
Borisov (2007)
UK
Coenagrion-
idae
Ischnura elegans
(Vander Linden, 1820)
Parr (2006b)
UK
Coenagrion-
idae
Ischnura elegans
(Vander Linden, 1820)
2
Tunmore (2005)
Arab
Penin-
sula
Coenagrion-
idae
Ischnura evansi
Morton, 1919
thousands of
immature indi-
viduals at light
source far
from the near-
est known
larval habitat
(T. Pittaway
per. comm.)
Corbet (1999)
Arab
Penin-
sula
Coenagrion-
idae
Ischnura evansi
Morton, 1919
to lighted
windows
Walker & Pittaway
(1987)
Oman
Coenagrion-
idae
Ischnura evansi
Morton, 1919
large number
attracted to
light
Waterson & Pittaway
(1991)
Russia
Coenagrion-
idae
Ischnura evansi
Morton, 1919
146
maximum
catch for one
night
Borisov (1990)
Russia
Coenagrion-
idae
Ischnura fountainei
Morton, 1905
411
maximum
catch for one
night
Borisov (1990)
USA
Coenagrion-
idae
Ischnura posita
(Hagen, 1861)
Frost (1966)
USA
Coenagrion-
idae
Ischnura posita
(Hagen, 1861)
Frost (1969)
Russia
Coenagrion-
idae
Ischnura pumilio
(Charpentier, 1825)
1
Borisov (1990)
Mexico
Coenagrion-
idae
Ischnura ramburii
(Selys in Sagra, 1857)
collected at
32, 74 and
106 km dis-
tance off the
shore
Sparks et al. (1986)
USA
Coenagrion-
idae
Ischnura ramburii
(Selys in Sagra, 1857)
Frost (1964)
USA
Coenagrion-
idae
Ischnura ramburii
(Selys in Sagra, 1857)
1
Wright (1944)
India
Coenagrion-
idae
Ischnura senegalensis
(Rambur, 1842)
occasionally
at the light
source
Andrew & Tembhare
(1997)
International Dragonfly Fund - Report 43 (2012): 1-52 31
Counrty
Family
Species
Sex
per
site
Notes
Reference
India
Coenagrion-
idae
Ischnura senegalensis
(Rambur, 1842)
Sharma et al. (2000)
USA
Coenagrion-
idae
Nehalennia integri-
collis Calvert, 1913
Frost (1964)
USA
Coenagrion-
idae
Nehalennia pallidula
Calvert, 1913
Frost (1969)
Russia
Coenagrion-
idae
Nehalennia speciosa
(Charpentier, 1840)
Kosterin & Dubatolov
(2005)
India
Coenagrion-
idae
Pseudagrion decorum
(Rambur, 1842)
Sharma et al. (2000)
India
Coenagrion-
idae
Rhodischnura nursei
(Morton, 1907)
occasionally
at the light
source
Andrew & Tembhare
(1997)
Mexico
Coenagrion-
idae
Coenagrionidae indet.
9 ind.
from
1 spe-
cies
collected at
32, 74 and
106 km dis-
tance off the
shore
Wolf et al. (1986)
USA
Lestidae
Archilestes grandis
(Rambur, 1942)
Platt & Harrison
(1994)
Japan
Lestidae
Indolestes peregrinus
(Ris, 1916)
1
collected on
28. 07.2007
at the
Omurasaki
Center, Ho-
kuto City
N. Ishizawa (per.
comm.)
Japan
Lestidae
Indolestes peregrinus
(Ris, 1916)
1
collected on
22. 07.2000
at the Omu-
rasaki Cen-
ter, Hokuto
City
N. Ishizawa (per.
comm.)
UK
Lestidae
Lestes sponsa
(Hansemann, 1823)
Paine (1992a)
UK
Lestidae
Lestes sponsa
(Hansemann, 1823)
Parr (2006b)
USA
Lestidae
Lestes vidua Hagen,
1861
Frost (1975)
USA
Lestidae
Lestes sp.
Platt & Harrison
(1994)
Russia
Lestidae
Sympecma gobica
Förster, 1900
Borisov (1990)
Russia
Lestidae
Sympecma paedisca
(Brauer, 1877)
1
Borisov (1990)
Russia
Lestidae
Sympecma paedisca
(Brauer, 1877)
Kosterin & Dubatolov
(2005)
Russia
Lestidae
Sympecma paedisca
(Brauer, 1877)
1 , 1
Malikova et al.
(2007)
Poland
Platycne-
mididae
Platycnemis pennipes
(Pallas, 1771)
2 ♂♂
Buczyński &
Buczyńska (2010)
Switzer-
Platycne-
Platycnemis pennipes
2 ♂♂, 2 ♀♀
Hoess & Rezbanyai-
32 Odonata attracted at light
Counrty
Family
Species
Sex
per
site
Notes
Reference
land
mididae
(Pallas, 1771)
Reser (2005)
Switzer-
land
Aeshnidae
Aeshna cyanea
(Müller, 1764)
20 ♂♂
Hoess & Rezbanyai-
Reser (2005)
Switzer-
land
Aeshnidae
Aeshna cyanea
(Müller, 1764)
1
Schiess (1982)
UK
Aeshnidae
Aeshna cyanea
(Müller, 1764)
Bayford (1911)
UK
Aeshnidae
Aeshna cyanea
(Müller, 1764)
1
possibly mi-
grating indi-
vidual
Bland (1997)
UK
Aeshnidae
Aeshna cyanea
(Müller, 1764)
Moorley (1919)
UK
Aeshnidae
Aeshna cyanea
(Müller, 1764)
Parr (2006b)
UK
Aeshnidae
Aeshna cyanea
(Müller, 1764)
1
Roddis (2007)
Denmark
Aeshnidae
Aeshna grandis
(Linnaeus, 1758)
1
catching flies
around the
altar candles
Wesenberg-Lund
(1913)
UK
Aeshnidae
Aeshna grandis
(Linnaeus, 1758)
1
Dannreuther (1937a)
UK
Aeshnidae
Aeshna grandis
(Linnaeus, 1758)
Paine (1996b)
UK
Aeshnidae
Aeshna grandis
(Linnaeus, 1758)
Parr (2006b)
Bulgaria
Aeshnidae
Aeshna mixta Latreille,
1805
1
S. Beshkov (per.
observ.)
Spain
Aeshnidae
Aeshna mixta Latreille,
1805
79
Riddiford (1992)
Switzer-
land
Aeshnidae
Aeshna mixta Latreille,
1805
21 ♂♂
Hoess & Rezbanyai-
Reser (2005)
Switzer-
land
Aeshnidae
Aeshna mixta Latreille,
1805
Schiess (1982)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
Brown (2000)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
Hundreds in-
dividuals fly-
ing pass the
ship
Dannreuther (1935)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
Dannreuther (1937a)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
Dannreuther (1937b)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
Deans (2005)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
1
Dewick (2000)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
1
Dewick (2006)
International Dragonfly Fund - Report 43 (2012): 1-52 33
Counrty
Family
Species
Sex
per
site
Notes
Reference
UK
Aeshnidae
Aeshna mixta Latreille,
1805
1
Hadley (1980)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
Jones (2004)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
Odin (2006)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
1
Parr (2000b)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
20+ records
Parr (2006b)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
1
Perrin (2011)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
1
Tunmore (2006)
UK
Aeshnidae
Aeshna mixta Latreille,
1805
1
Tunmore (2010)
Poland
Aeshnidae
Aeshna viridis
Eversmann, 1836
1 , 4 ♀♀
Zieba & Buczynski
(2007)
UK
Aeshnidae
Aeshna viridis
Eversmann, 1836
1
Mattila (2001)
Cayman
islands
Aeshnidae
Anax amazili
(Burmeister, 1839)
1
Askew et al. (1998)
not
specified
Aeshnidae
Anax amazili
(Burmeister, 1839)
2 ♀♀
Campos (1931)
Cyprus
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
on the decks
of a ship
Averill (1995)
Dubai
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
1
Chalmers (2010)
Egypt
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
Silsby (1993)
India
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
collected dur-
ing monsoon
Asaithambi &
Manickavasagam
(2002)
Iran
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
Sutton (1966)
Monte-
negro
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
1
Dumont (1977)
Russia
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
Borisov (1990)
UAE
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
1
attracted by
the light of a
lantern
Feulner (2007)
Uganda
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
20 ♂♂, 46 ♀♀
all but 1
were near
the light trap
Corbet (1984)
UK
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
1
Paine (1996b)
UK
Aeshnidae
Anax ephippiger
(Burmeister, 1839)
Parr (2006b)
India
Aeshnidae
Anax guttatus
predominant-
Andrew & Tembhare
34 Odonata attracted at light
Counrty
Family
Species
Sex
per
site
Notes
Reference
(Burmeister, 1839)
ly collected at
light source
(1997)
India
Aeshnidae
Anax guttatus
(Burmeister, 1839)
Arulprakash &
Gunathilagaraj
(2010)
India
Aeshnidae
Anax guttatus
(Burmeister, 1839)
collected dur-
ing monsoon
Asaithambi & Manic-
kavasagam (2002)
India
Aeshnidae
Anax guttatus
(Burmeister, 1839)
18
Patil et al. (1982)
Laos
Aeshnidae
Anax guttatus
(Burmeister, 1839)
lighted
candle
Pemberton (1995)
Republic
of
Maldives
Aeshnidae
Anax guttatus
(Burmeister, 1839)
1
Mahlendorf &
Martens (2004)
Singa-
pore
Aeshnidae
Anax guttatus
(Burmeister, 1839)
1
Norma-Rashid et al.
(2008)
Tailand
Aeshnidae
Anax guttatus
(Burmeister, 1839)
1
Hämäläinen (1988)
India
Aeshnidae
Anax immaculifrons
Rambur, 1842
predominant-
ly collected at
light source
Andrew & Tembhare
(1997)
India
Aeshnidae
Anax immaculifrons
Rambur, 1842
Sharma et al. (2000)
Russia
Aeshnidae
Anax imperator Leach,
1815
Sluvko (2007)
Switzerla
nd
Aeshnidae
Anax imperator Leach,
1815
1
Hoess & Rezbanyai-
Reser (2005)
UK
Aeshnidae
Anax imperator Leach,
1815
Parr (2006b)
UK
Aeshnidae
Anax imperator Leach,
1815
1
Parr (2007)
Mexico
Aeshnidae
Anax junius (Drury,
1773)
8
collected at
74, 106 & 160
km distance
off the shore
Sparks et al. (1986)
USA
Aeshnidae
Anax junius (Drury,
1773)
Beckemeyer (2003)
USA
Aeshnidae
Anax junius (Drury,
1773)
Frost (1966)
USA
Aeshnidae
Anax junius (Drury,
1773)
1
Platt & Harrison
(1994)
USA
Aeshnidae
Anax junius (Drury,
1773)
Wright (1944)
Willis
Island,
Australia
Aeshnidae
Anax papuensis
(Burmeister, 1839)
11
Farrow (1984)
Dubai
Aeshnidae
Anax parthenope
(Selys, 1839)
2 ♀♀
Chalmers (2010)
Egypt
Aeshnidae
Anax parthenope
(Selys, 1839)
Silsby (1993)
International Dragonfly Fund - Report 43 (2012): 1-52 35
Counrty
Family
Species
Sex
per
site
Notes
Reference
India
Aeshnidae
Anax parthenope
(Selys, 1839)
sitting near
light sources
Asaithambi & Manic-
kavasagam (2002)
Oman
Aeshnidae
Anax parthenope
(Selys, 1839)
1
Waterson & Pittaway
(1991)
Russia
Aeshnidae
Anax parthenope
(Selys, 1839)
Borisov (1990)
UK
Aeshnidae
Anax parthenope
(Selys, 1839)
1
Parr (2006b)
China
Aeshnidae
Anax parthenope
julius Brauer, 1865
Easton & Liang
(2000)
China
Aeshnidae
Anax parthenope
julius Brauer, 1865
59,
342
caught during
2003 field
season
Feng et al. (2006)
Gambia
Aeshnidae
Anax tristis Hagen,
1867
Silsby (1999)
off
Angola
Aeshnidae
Anax tristis Hagen,
1867
1
60 km from
mainland
Schnieder (1992)
Tanganyi
ka
Aeshnidae
Anax tristis Hagen,
1867
Pinhey (1961)
France
Aeshnidae
Boyeria irene
(Fonscolombe, 1838)
1
Morton (1932)
Canada
Aeshnidae
Boyeria vinosa (Say,
1840)
1
Hutchingso (2001)
not
specified
Aeshnidae
Coryphaeschna
adnexa (Hagen, 1861)
1
Campos (1931)
USA
Aeshnidae
Coryphaeschna
ingens (Rambur,
1842)
Frost (1964)
French
Guyana
Aeshnidae
Coryphaeschna
viriditas Calvert, 1952
1 , 1
Geijskes (1971)
Trinidad
Aeshnidae
Coryphaeschna
viriditas Calvert, 1952
3 ♂♂, 5 ♀♀
Corbet (1981)
USA
Aeshnidae
Epiaeschna heros
(Fabricius, 1798)
1
Platt & Harrison
(1994)
USA
Aeshnidae
Gomphaeschna
antilope (Hagen,
1874)
Frost (1964)
India
Aeshnidae
Gynacantha bayadera
Selys, 1891 (= G.
furcata?)
predominant-
ly collected at
light source
Andrew & Tembhare
(1997)
Ivory
Coast
Aeshnidae
Gynacantha bullata
Karsch, 1891
4 ♂♂, 1
Dumont (2004)
Sarawak,
Malaysia
Aeshnidae
Gynacantha ? dohrni
Krüger, 1899
Dow (2005)
India
Aeshnidae
Gynacantha hyalina
Selys, 1882
Arulprakash &
Gunathilagaraj
(2010)
India
Aeshnidae
Gynacantha hyalina
Selys, 1882
collected dur-
ing monsoon
Asaithambi & Manic-
kavasagam (2002)
Trinidad
Aeshnidae
Gynacantha mexicana
1
Corbet (1981)
36 Odonata attracted at light
Counrty
Family
Species
Sex
per
site
Notes
Reference
Selys, 1868
Cayman
Islands
Aeshnidae
Gynacantha nervosa
Rambur, 1842
1
Askew et al. (1998)
Trinidad
Aeshnidae
Gynacantha nervosa
Rambur, 1842
4 ♂♂, 1
Corbet (1981)
USA
Aeshnidae
Gynacantha nervosa
Rambur, 1842
Frost (1964)
USA
Aeshnidae
Gynacantha nervosa
Rambur, 1842
2
Ihssen (1997)
India
Aeshnidae
Gynacantha rammo-
hani Mitra & Lahiri,
1975
1
holotype
Mitra & Lahiri (1975)
Tailand
Aeshnidae
Gynacantha saltatrix
Martin, 1909
Hämäläinen (1987a)
Borneo,
Malaysia
Aeshnidae
Gynacantha sp.
Orr (1995)
Ivory
Coast
Aeshnidae
Gynacantha sp.
1
Dumont (2004)
not
specified
Aeshnidae
Gynacantha sp.
1
Campos (1931)
Sarawak,
Malaysia
Aeshnidae
Gynacantha sp.
1
Dow (2005)
Sarawak,
Malaysia
Aeshnidae
Heliaeschna crassa
Krüger, 1899
Dow (2005)
Borneo,
Malaysia
Aeshnidae
Heliaeschna sp.
Orr (1995)
Sarawak,
Malaysia
Aeshnidae
Heliaeschna sp.
1
Dow (2005)
Malay
Penin-
sula &
Indonesia
Aeshnidae
Indaeschna grubaueri
(Förster, 1904)
Lieftinck (1954)
French
Guyana
Aeshnidae
Neuraeschna clavifor-
cipata Martin, 1909
1
Geijskes (1971)
French
Guyana
Aeshnidae
Neuraeschna costalis
(Burmeister, 1839)
1
Geijskes (1971)
Borneo,
Malaysia
Aeshnidae
Oligaeschna sp.
Orr (1995)
Malay
Penin-
sula
Aeshnidae
Periaeschna laidlawi
(Förster, 1908)
immature
Lieftinck (1954)
Sarawak,
Malaysia
Aeshnidae
Tetracanthagyna brun-
nea McLachlan, 1898
1
Dow (2005)
Malay
Penin-
sula
Aeshnidae
Tetracanthagyna
plagiata (Waterhouse,
1877)
Lieftinck (1954)
Cayman
Islands
Aeshnidae
Triacanthagyna septi-
ma (Selys in Sagra,
1857)
1
Askew et al. (1998)
not
Aeshnidae
Triacanthagyna septi-
Campos (1931)
International Dragonfly Fund - Report 43 (2012): 1-52 37
Counrty
Family
Species
Sex
per
site
Notes
Reference
specified
ma (Selys in Sagra,
1857)
Trinidad
Aeshnidae
Triacanthagyna sept-
ima (Selys in Sagra,
1857)
1
Corbet (1981)
USA
Aeshnidae
Triacanthagyna trifida
(Rambur, 1842)
Frost (1964)
Mexico
Aeshnidae
Aeshnidae indet.
8 ind.
from
1 sp.
collected at
74, 106 & 160
km distance
from the shore
Wolf et al. (1986)
Australia
Corduliidae
Austrocordulia refracta
Tillyard, 1909
Theischinger (2010)
USA
Corduliidae
Epitheca cynosura
(Say, 1840)
Frost (1966)
USA
Corduliidae
Epitheca princeps
Hagen, 1861
63 ♂♂, 15 ♀♀
for 17 conse-
quitive nights
Young (1967)
USA
Corduliidae
Epitheca sepia (Gloyd,
1933)
Frost (1966)
Malaysia
Corduliidae
Hemicordulia
gracillima Fraser, 1944
1
holotype
Fraser (1944)
USA
Corduliidae
Neurocordulia yama-
skanensis
(Provancher, 1875)
1
Montgomery (1955)
Switzer-
land
Corduliidae
Oxygastra curtisii
(Dale, 1834)
1
Hoess & Rezbanyai-
Reser (2005)
Thailand
Gomphidae
Acrogomphus minor
Laidlaw, 1931
1
paratype
Laidlaw (1931)
Russia
Gomphidae
Anormogomphus
kiritschenkoi Bartenev,
1913
1
teneral
female
Borisov (1990)
Australia
Gomphidae
Austrogomphus aus-
tralis Dale in Selys,
1854
Corbet (1999)
Malaysia
Gomphidae
Burmagomphus
arthuri Lieftinck, 1953
1
new for the
country
Hämäläinen (2000)
French
Guyana
Gomphidae
Cacoides latro
(Erichson, 1848)
2 ♀♀
Geijskes (1971)
Russia
Gomphidae
Gomphus flavipes li-
neatus Bartenev, 1929
tenerals
Borisov (2007)
Peru
Gomphidae
Heterogomphus sp.
Burmeister (2006)
Thailand
Gomphidae
Ictinogomphus de-
coratus melaenops
Selys, 1857
1
Hämäläinen (1987b)
India
Gomphidae
Ictinogomphus rapax
(Rambur, 1842)
Predominant-
ly collected at
light source
Andrew & Tembhare
(1997)
India
Gomphidae
Ictinogomphus rapax
(Rambur, 1842)
sitting near
light sources
Asaithambi &
Manickavasagam
(2002)
38 Odonata attracted at light
Counrty
Family
Species
Sex
per
site
Notes
Reference
India
Gomphidae
Ictinogomphus rapax
(Rambur, 1842)
Sharma et al. (2000)
Kuwait
Gomphidae
Lindenia tetraphylla
(Vander Linden, 1825)
sampled dur-
ing a sand
storm
Al-Houty (1985)
Kuwait
Gomphidae
Lindenia tetraphylla
(Vander Linden, 1825)
sampled dur-
ing a sand
storm
Al-Houty (1989)
Russia
Gomphidae
Lindenia tetraphylla
(Vander Linden, 1825)
1 , 1
teneral
female
Borisov (1990)
Russia
Gomphidae
Lindenia tetraphylla
(Vander Linden, 1825)
1
Skvortsov & Kutaev
(2010)
Indone-
sia
Gomphidae
Macrogomphus p.
parallelogramma
(Burmeister, 1839)
♀♀
Lieftinck (1954)
USA
Gomphidae
Melanocacus mungo
(Needham, 1940)
2 ♂♂
teneral +
holotype
Needham (1940)
Uganda
Gomphidae
Notogomphus lujai
(Schouteden, 1934)
1
mature
individual
Corbet (1961)
Russia
Gomphidae
Onychogomphus
flexuosus (Schneider,
1845)
1 , 1
teneral
female
Borisov (1990)
Bulgaria
Gomphidae
Onychogomphus
forcipatus (Linnaeus,
1758)
1
S. Beshkov & M.
Beshkova (per.
observ.)
Switzer-
land
Gomphidae
Onychogomphus f.
forcipatus (Linnaeus,
1758)
1 , 1
Hoess & Rezbanyai-
Reser (2005)
Russia
Gomphidae
Ophiogomphus
reductus Calvert, 1898
1
Borisov (1990)
Namibia
Gomphidae
Paragomphus cogna-
tus (Rambur, 1842)
1
Martens et al. (2003)
India
Gomphidae
Paragomphus lineatus
(Selys, 1850)
predominant-
ly collected at
light source
Andrew & Tembhare
(1997)
Sudan
Gomphidae
Paragomphus pumilio
(Rambur, 1842)
Ris (1924)
French
Guyana
Gomphidae
Phyllogomphoides
andromeda (Selys,
1869)
1 , 1
Geijskes (1971)
French
Guyana
Gomphidae
Phyllogomphoides
fuliginosus (Hagen in
Selys, 1854)
1
Geijskes (1971)
Peru
Gomphidae
Phyllogomphoides
selysi (Navás, 1924)
2
Dunkle (1989)
Uganda
Gomphidae
Phyllogomphus selysi
Schouteden, 1933
1
Dijkstra et al. (2006)
French
Guyana
Gomphidae
Progomphus brachy-
cnemis Needham,
1944
1
juvenile
Geijskes (1971)
International Dragonfly Fund - Report 43 (2012): 1-52 39
Counrty
Family
Species
Sex
per
site
Notes
Reference
China
Gomphidae
Sinictinogomphus
clavatus (Fabricius,
1775)
Easton & Liang
(2000)
Ivory
Coast
Libellulidae
Aethriamanta rezia
Kirby, 1889
1
Dumont (2004)
French
Guyana
Libellulidae
Anatya guttata
(Erichson, 1848)
1
Geijskes (1971)
Trinidad
Libellulidae
Anatya guttata
(Erichson, 1848)
1
Corbet (1981)
India
Libellulidae
Brachydiplax sobrina
(Rambur, 1842)
Sharma et al. (2000)
Trinidad
Libellulidae
Brachymesia furcata
(Hagen, 1861)
1
Corbet (1981)
Trinidad
Libellulidae
Brachymesia herbida
(Gundlach, 1889)
1
Corbet (1981)
India
Libellulidae
Brachythemis conta-
minata (Fabricius,
1793)
prey of gecko
Mitra (1974)
India
Libellulidae
Brachythemis conta-
minata (Fabricius,
1793)
Sharma et al. (2000)
India
Libellulidae
Brachythemis conta-
minata (Fabricius,
1793)
3
Shull & Nadkerny
(1967)
Uganda
Libellulidae
Brachythemis leuco-
sticta (Burmeister,
1839)
Corbet (1959)
India
Libellulidae
Bradinopyga geminata
(Rambur, 1842)
occasionally
at the light
source
Andrew & Tembhare
(1997)
India
Libellulidae
Bradinopyga geminata
(Rambur, 1842)
1
Shull & Nadkerny
(1967)
Malay
Penin-
sula &
Indonesia
Libellulidae
Camacinia gigantea
(Brauer, 1867)
Lieftinck (1954)
USA
Libellulidae
Celithemis eponina
(Drury, 1773)
Frost (1975)
USA
Libellulidae
Celithemis ornata
(Rambur, 1842)
Frost (1964)
Ivory
Coast
Libellulidae
Chalcostephia flavi-
frons Kirby, 1889
4 ♂♂, 2 ♀♀
Dumont (2004)
Nigeria
Libellulidae
Crocothemis erythraea
(Brullé, 1832)
this paper
Nigeria
Libellulidae
Crocothemis sangui-
nolenta (Burmeister,
1839)
this paper
India
Libellulidae
Crocothemis servilia
(Drury, 1773)
occasionally
at the light
source
Andrew & Tembhare
(1997)
40 Odonata attracted at light
Counrty
Family
Species
Sex
per
site
Notes
Reference
India
Libellulidae
Crocothemis servilia
(Drury, 1773)
prey of gecko
Mitra (1974)
India
Libellulidae
Crocothemis servilia
(Drury, 1773)
Sharma et al. (2000)
Russia
Libellulidae
Crocothemis servilia
(Drury, 1773)
30
maximum
catch for one
night
Borisov (1990)
Russia
Libellulidae
Diplacodes lefebvrii
(Rambur, 1842)
Borisov (1990)
Uganda
Libellulidae
Diplacodes luminans
(Karsch, 1893)
21 ♂♂, 27 ♀♀
Corbet (1984)
India
Libellulidae
Diplacodes trivialis
(Rambur, 1842)
Sharma et al. (2000)
USA
Libellulidae
Erythemis simplici-
collis (Say, 1840)
Frost (1975)
USA
Libellulidae
Erythemis simplici-
collis (Say, 1840)
Wright (1944)
French
Guyana
Libellulidae
Erythrodiplax famula
(Erichson, 1848)
12 ♂♂, 24 ♀♀
Geijskes (1971)
not
specified
Libellulidae
Erythrodiplax fusca
(Rambur, 1842)
1
Campos (1931)
Trinidad
Libellulidae
Erythrodiplax fusca
(Rambur, 1842)
1
Corbet (1981)
French
Guyana
Libellulidae
Erythrodiplax haema-
togastra (Burmeister,
1839)
1
Geijskes (1971)
French
Guyana
Libellulidae
Erythrodiplax longitu-
dinalis (Ris, 1919)
6 ♂♂, 3 ♀♀
Geijskes (1971)
USA
Libellulidae
Erythrodiplax minus-
cula (Rambur, 1842)
Frost (1964)
Trinidad
Libellulidae
Erythrodiplax umbrata
(Linnaeus, 1758)
1
Corbet (1981)
Ivory
Coast
Libellulidae
Hemistigma albipuncta
(Rambur, 1842)
1
Dumont (2004)
Ukraine
Libellulidae
Leucorrhinia pectoralis
(Charpentier, 1825)
1 , 1
Bartenef (1933)
USA
Libellulidae
Libellula auripennis
Burmeister, 1839
Wright (1944)
USA
Libellulidae
Libellula axilena
Westwood, 1837
Yosef (1994)
UK
Libellulidae
Libellula quadrimacu-
lata Linnaeus, 1758
Dannreuther (1935)
Thailand
Libellulidae
Macrodiplax cora
(Kaup in Brauer, 1867)
1
Hämäläinen (1987b)
French
Guyana
Libellulidae
Micrathyria aequalis
(Hagen, 1861)
1
Geijskes (1971)
French
Guyana
Libellulidae
Micrathyria atra
(Martin, 1897)
1
Geijskes (1971)
Trinidad
Libellulidae
Micrathyria atra
(Martin, 1897)
1
Corbet (1981)
International Dragonfly Fund - Report 43 (2012): 1-52 41
Counrty
Family
Species
Sex
per
site
Notes
Reference
Trinidad
Libellulidae
Micrathyria laevigata
Calvert, 1909
1
Corbet (1981)
French
Guyana
Libellulidae
Micrathyria mengeri
Ris, 1919
1
Geijskes (1971)
Trinidad
Libellulidae
Micrathyria ocellata
Martin, 1897
1 , 1
Corbet (1981)
French
Guyana
Libellulidae
Nephepeltia flavifrons
(Karsch, 1889)
1
Geijskes (1971)
Malaysia
Libellulidae
Neurothemis tullia
tullia (Drury, 1773)
1
Lieftinck (1954)
Uganda
Libellulidae
Notiothemis robertsi
Fraser, 1944
1
mature
individual
Corbet (1961)
Ivory
Coast
Libellulidae
Olpogastra lugubris
(Karsch, 1895)
1
Dumont (2004)
French
Guyana
Libellulidae
Orthemis ferruginea
(Fabricius, 1775)
1
Geijskes (1971)
Trinidad
Libellulidae
Orthemis ferruginea
(Fabricius, 1775)
8 ♂♂, 5 ♀♀
Corbet (1981)
Trinidad
Libellulidae
Orthemis sulphurata
Hagen, 1868
2 ♀♀
Corbet (1981)
Ivory
Coast
Libellulidae
Orthetrum brachiale
(Palisot de Beauvois,
1805)
2 ♂♂, 4 ♀♀
Dumont (2004)
Russia
Libellulidae
Orthetrum brunneum
(Fonscolombe, 1837)
Borisov (1990)
Nigeria
Libellulidae
Orthetrum camerunen-
se Gambles, 1959
this paper
Switzer-
land
Libellulidae
Orthetrum cancellatum
(Linnaeus, 1758)
1
Hoess & Rezbanyai-
Reser (2005)
Nigeria
Libellulidae
Orthetrum chrysostig-
ma (Burmeister, 1839)
this paper
Nigeria
Libellulidae
Orthetrum ? guineen-
se Ris, 1909
this paper
Uganda
Libellulidae
Orthetrum hintzi
Schmidt, 1951
1
mature
individual
Corbet (1961)
not
specified
Libellulidae
Orthetrum icteromelas
cinctifrons Pinhey,
1970
1
holotype
Dijkstra (2007)
Uganda
Libellulidae
Orthetrum julia Kirby,
1900
1 , 2 ♀♀
probably of
this species
mature
individual
Corbet (1961)
India
Libellulidae
Orthetrum sabina
(Drury, 1770)
occasionally
at the light
source
Andrew & Tembhare
(1997)
India
Libellulidae
Orthetrum sabina
(Drury, 1770)
Sharma et al. (2000)
Russia
Libellulidae
Orthetrum sabina
(Drury, 1770)
Borisov (1990)
Uganda
Libellulidae
Orthetrum ? trinacria
(Selys, 1841)
identified by
supposition
Corbet (1959)
42 Odonata attracted at light
Counrty
Family
Species
Sex
per
site
Notes
Reference
Japan
Libellulidae
Orthetrum triangulare
melania(Selys, 1883)
1 , 1
collected on
27.07. 2002
at the Omu-
rasaki Cen-
ter, Hokuto
City
N. Ishizawa (per.
comm.)
India
Libellulidae
Orthetrum sp.
3
Shull & Nadkerny
(1967)
USA
Libellulidae
Pachydiplax longi-
pennis (Burmeister,
1839)
Frost (1964)
USA
Libellulidae
Pachydiplax longi-
pennis (Burmeister,
1839)
43 ♂♂, 53 ♀♀
collected for
one month
Frost (1971)
USA
Libellulidae
Pachydiplax longi-
pennis (Burmeister,
1839)
Frost (1975)
USA
Libellulidae
Pachydiplax longi-
pennis (Burmeister,
1839)
Wright (1944)
Nigeria
Libellulidae
Palpopleura portia
(Drury, 1773)
this paper
China
Libellulidae
Pantala flavescens
(Fabricius, 1798)
96
caught during
2003 field
season
Feng et al. (2006)
French
Guyana
Libellulidae
Pantala flavescens
(Fabricius, 1798)
1
Geijskes (1971)
India
Libellulidae
Pantala flavescens
(Fabricius, 1798)
Sharma et al. (2000)
India
Libellulidae
Pantala flavescens
(Fabricius, 1798)
Shull & Nadkerny
(1967)
Japan
Libellulidae
Pantala flavescens
(Fabricius, 1798)
10
Asahina & Turuoka
(1967)
Japan
Libellulidae
Pantala flavescens
(Fabricius, 1798)
Honda (2003)
Japan
Libellulidae
Pantala flavescens
(Fabricius, 1798)
Inoue (1998)
Japan
Libellulidae
Pantala flavescens
(Fabricius, 1798)
tens of kilome-
tres off shore
Yamane & Hashi-
guchi (1994)
Republic
of
Maldives
Libellulidae
Pantala flavescens
(Fabricius, 1798)
1
Mahlendorf &
Martens (2004)
Russia
Libellulidae
Pantala flavescens
(Fabricius, 1798)
Borisov (1990)
Trinidad
Libellulidae
Pantala flavescens
(Fabricius, 1798)
1
Corbet (1981)
Willis
Island,
Australia
Libellulidae
Pantala flavescens
(Fabricius, 1798)
3
Farrow (1984)
Cayman
islands
Libellulidae
Pantala hymenaea
Say, 1840
1
Askew et al. (1998)
International Dragonfly Fund - Report 43 (2012): 1-52 43
Counrty
Family
Species
Sex
per
site
Notes
Reference
Mexico
Libellulidae
Pantala hymenaea
Say, 1840
1
collected at
160 km off
shore
Sparks et al. (1986)
India
Libellulidae
Pantala sp.
36
Patil et al. (1982)
Namibia
Libellulidae
Parazyxomma flavi-
cans (Martin, 1908)
1
Martens et al. (2003)
Uganda
Libellulidae
Parazyxomma flavi-
cans (Martin, 1908)
Corbet (1959)
Trinidad
Libellulidae
Perithemis mooma
Kirby, 1889
1
Corbet (1981)
USA
Libellulidae
Perithemis tenera
(Say, 1840)
Montgomery (1955)
Singa-
pore
Libellulidae
Pornothemis serrata
Krüger, 1902
1
Norma-Rashid et al.
(2008)
Kuwait
Libellulidae
Selysiothemis nigra
(Vander Linden, 1825)
great
num-
ber
sampled dur-
ing a sand
storm
Al-Houty (1985)
Kuwait
Libellulidae
Selysiothemis nigra
(Vander Linden, 1825)
sampled dur-
ing a sand
storm
Al-Houty (1989)
Russia
Libellulidae
Selysiothemis nigra
(Vander Linden, 1825)
Borisov (1990)
Russia
Libellulidae
Selysiothemis nigra
(Vander Linden, 1825)
1
Skvortsov & Kutaev
(2010)
USA
Libellulidae
Sympetrum ambiguum
(Rambur, 1842)
Montgomery (1955)
Russia
Libellulidae
Sympetrum arenicolor
Jödicke, 1994
Borisov (2007)
Russia
Libellulidae
Sympetrum danae
(Sulzer, 1776)
Kosterin & Dubatolov
(2005)
UK
Libellulidae
Sympetrum danae
(Sulzer, 1776)
Parr (2006b)
Russia
Libellulidae
Sympetrum
depressiusculum
(Selys, 1841)
Kosterin & Dubatolov
(2005)
Russia
Libellulidae
Sympetrum eroticum
(Selys, 1883)
1
Malikova et al.
(2007)
Japan
Libellulidae
Sympetrum eroticum
eroticum (Selys, 1883)
1
collected on
22.07.2000
at the Omu-
rasaki Cen-
ter, Hokuto
City
N. Ishizawa (per.
comm.)
UK
Libellulidae
Sympetrum flaveolum
(Linnaeus, 1758)
1
Mendel & Marsh
(1995)
UK
Libellulidae
Sympetrum flaveolum
(Linnaeus, 1758)
Paine (1995)
UK
Libellulidae
Sympetrum flaveolum
(Linnaeus, 1758)
1
Paine (1996a)
UK
Libellulidae
Sympetrum flaveolum
Parr (2006b)
44 Odonata attracted at light
Counrty
Family
Species
Sex
per
site
Notes
Reference
(Linnaeus, 1758)
Russia
Libellulidae
Sympetrum fonsco-
lombii (Selys, 1840)
Borisov (1990)
UK
Libellulidae
Sympetrum fonsco-
lombii (Selys, 1840)
Parr (2006b)
UK
Libellulidae
Sympetrum fonsco-
lombii (Selys, 1840)
1
Tunmore (2002)
UK
Libellulidae
Sympetrum fonsco-
lombii (Selys, 1840)
small
numbers
Tunmore (2007)
Switzer-
land
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
1
Hoess & Rezbanyai-
Reser (2005)
UK
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
1
Mendel & Marsh
(1995)
UK
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
Odin (2006)
UK
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
1
Paine (1992a)
UK
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
Paine (1995)
UK
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
9
Parr (2000a)
UK
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
Parr (2001)
UK
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
12
Parr (2005)
UK
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
20+ records
Parr (2006b)
UK
Libellulidae
Sympetrum sangui-
neum (Müller, 1764)
single indivi-
duals
Parr (2007)
Spain
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
4
Riddiford (1992)
Switzer-
land
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
1 , 3 ♀♀
Hoess & Rezbanyai-
Reser (2005)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
Cade (2004)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
Dannreuther (1937b)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
Deans (2005)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
1
Deans (2006a)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
Deans (2006b)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
Deans (2008)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
Deans (2009)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
1
Deans (2010)
International Dragonfly Fund - Report 43 (2012): 1-52 45
Counrty
Family
Species
Sex
per
site
Notes
Reference
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
10
for a 20-days
period
Dewick (1999)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
19
total for the
season
Dewick (2000)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
Dewick (2006)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
1
Dewick (2011)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
Moore (2009)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
42
Odin (2006)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
1
Paine (1996a)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
19
Parr (2000a)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
1
Parr (2001)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
26
Parr (2005)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
approxima-
tely 100 re-
cords
Parr (2006b)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
occasionally
at the light
source
Tunmore (2005)
UK
Libellulidae
Sympetrum striolatum
(Charpentier, 1840)
Tunmore (2010)
Russia
Libellulidae
Sympetrum vulgatum
(Linnaeus, 1758)
Borisov (2007)
Switzer-
land
Libellulidae
Sympetrum vulgatum
(Linnaeus, 1758)
2
Hoess & Rezbanyai-
Reser (2005)
Russia
Libellulidae
Sympetrum vulgatum
decoratum Selys,
1884
Borisov (1990)
Switzer-
land
Libellulidae
Sympetrum sp.
Aubert (1964)
UK
Libellulidae
Sympetrum sp.
Dewick (2001)
UK
Libellulidae
Sympetrum sp.
18
Parr (2001)
USA
Libellulidae
Sympetrum sp.
3
Platt & Harrison
(1994)
Trinidad
Libellulidae
Tauriphila australis
(Hagen, 1867)
1
Corbet (1981)
Trinidad
Libellulidae
Tholymis citrina
Hagen, 1867
1
Corbet (1981)
India
Libellulidae
Tholymis tillarga
(Fabricius, 1798)
Arulprakash & Guna-
thilagaraj (2010)
India
Libellulidae
Tholymis tillarga
(Fabricius, 1798)
mostly collec-
ted sitting
Asaithambi &
Manickavasagam
46 Odonata attracted at light
Counrty
Family
Species
Sex
per
site
Notes
Reference
near the light
source
(2002)
India
Libellulidae
Tholymis tillarga
(Fabricius, 1798)
Mitra (1974)
India
Libellulidae
Tholymis tillarga
(Fabricius, 1798)
Sharma et al. (2000)
Ivory
Coast
Libellulidae
Tholymis tillarga
(Fabricius, 1798)
15 ♂♂, 34 ♀♀
Dumont (2004)
off
Angola
Libellulidae
Tholymis tillarga
(Fabricius, 1798)
2 ♂♂, 1
60 km from
mainland
Schnieder (1992)
Willis
Island,
Australia
Libellulidae
Tholymis tillarga
(Fabricius, 1798)
4
Farrow (1984)
not
specified
Libellulidae
Tramea basilaris
(Palisot de Beauvois,
1805)
Campos (1931)
Uganda
Libellulidae
Tramea basilaris
(Palisot de Beauvois,
1805)
6 ♂♂, 9 ♀♀
Corbet (1984)
India
Libellulidae
Tramea basilaris
burmeisteri Kirby,
1889
occasionally
at the light
source
Andrew & Tembhare
(1997)
Trinidad
Libellulidae
Tramea calverti
Muttkowski, 1910
2 ♂♂, 4 ♀♀
Corbet (1981)
USA
Libellulidae
Tramea carolina
(Linnaeus, 1763)
Frost (1964)
Republic
of
Maldives
Libellulidae
Tramea limbata
(Desjardins, 1832)
1
Mahlendorf &
Martens (2004)
Willis
Island,
Australia
Libellulidae
Tramea loewii Kaup in
Brauer, 1866
1
Farrow (1984)
Mexico