begin quote from:
damselflies
(Zygoptera)
Damselfly
From Wikipedia, the free encyclopedia
Damselflies are
insects of the suborder
Zygoptera in the order
Odonata. They are similar to
dragonflies, which constitute the other odonatan suborder,
Anisoptera,
but are smaller, have slimmer bodies, and most species fold the wings
along the body when at rest. An ancient group, damselflies have existed
since at least the
Lower Permian, and are found on every continent except
Antarctica.
All damselflies are
predatory;
both nymphs and adults eat other insects. The nymphs are aquatic, with
different species living in a variety of freshwater habitats including
acid bogs, ponds, lakes and rivers. The nymphs moult repeatedly, at the
last moult climbing out of the water to undergo
metamorphosis.
The skin splits down the back, they emerge and inflate their wings and
abdomen to gain their adult form. Their presence on a body of water
indicates that it is relatively unpolluted, but their dependence on
freshwater makes them vulnerable to damage to their wetland habitats.
Some species of damselfly have elaborate courtship behaviours. Many species are
sexually dimorphic, the males often being more brightly coloured than the females. Like dragonflies, they reproduce using indirect
insemination and delayed
fertilisation.
A mating pair form a shape known as a "heart" or "wheel", the male
clasping the female at the back of the head, the female curling her
abdomen down to pick up sperm from secondary genitalia at the base of
the male's abdomen. The pair often remain together with the male still
clasping the female while she lays eggs within the tissue of plants in
or near water using a robust
ovipositor.
Fishing flies that mimic damselfly nymphs are used in wet-fly
fishing. Damselflies sometimes provide the subject for personal
jewellery such as brooches.
Classification
The Zygoptera are an ancient group, with fossils known from the lower
Permian,
at least 250 million years ago. All the fossils of that age are of
adults, similar in structure to modern damselflies, so it is not known
whether their larvae were aquatic at that time. The earliest larval
odonate fossils are from the
Mesozoic.
[2] Fossils of damselfly-like Protozygoptera date back further to 311–30 Mya.
[3] Well-preserved
Eocene damselfly larvae and exuviae are known from fossils preserved in
amber in the Baltic region.
[4]
Molecular analysis in 2013 confirms that most of the traditional families are
monophyletic, but shows that the
Amphipterygidae,
Megapodagrionidae and
Protoneuridae are
paraphyletic
and will need to be reorganised. The Protoneuridae in particular is
shown to be composed of six clades from five families. The result so far
is 27 damselfly families, with 7 more likely to be created. The
discovered
clades
did not agree well with traditional characteristics used to classify
living and fossil Zygoptera such as wing venation, so fossil taxa will
need to be revisited. The 18 extant traditional families are
provisionally rearranged as follows (the 3 paraphyletic families
disappearing, and many details not resolved):
[5]
Dashed lines indicate unresolved relationships.
General description
Illustration of damselfly wing. Both fore and hind wings are similar
The general body plan of a damselfly is similar to that of a dragonfly. The
compound eyes
are large but are more widely separated and relatively smaller than
those of a dragonfly. Above the eyes is the frons or forehead, below
this the clypeus, and on the upper lip the
labrum, an extensible organ used in the capture of
prey. The top of the head bears three simple eyes (
ocelli), which may measure light intensity, and a tiny pair of
antennae that serve no olfactory function but may measure air speed.
[6] Many species are
sexually dimorphic; the males are often brightly-coloured and distinctive, while the females are plainer,
cryptically coloured, and harder to identify to species. For example, in
Coenagrion,
the Eurasian bluets, the males are bright blue with black markings,
while the females are usually predominantly green or brown with black.
A few dimorphic species show female-limited polymorphism, the females
being in two forms, one form distinct and the other with the patterning
as in males. The ones that look like males, andromorphs, are usually
under a third of the female population but the proportion can rise
significantly and a theory that explains this response suggests that it
helps overcome harassment by males.
[8] Some Coenagrionid damselflies show male-limited polymorphism, an even less understood phenomenon.
[9]
In general, damselflies are smaller than dragonflies, the smallest being members of the genus
Agriocnemis (wisps).
[10] However, members of the
Pseudostigmatidae (helicopter damselflies or forest giants) are exceptionally large for the group, with wingspans as much as 19 cm (7.5 in) in
Megaloprepus[11] and body length up to 13 cm (5.1 in) in
Pseudostigma aberrans.
[12]
Terminal segments of the abdomen of a male damselfly (
Pseudagrion caffrum)
showing segments 8–10 (S8, S9, S10), the upper or superior appendages
or cerci (c) and the inferior appendages or paraprocts (p)
The first
thoracic
segment is the prothorax, bearing the front pair of legs. The joint
between head and prothorax is slender and flexible, which enables the
damselfly to swivel its head and to manoeuvre more freely when flying.
The remaining thoracic segments are the fused mesothorax and metathorax
(together termed the
synthorax),
each with a pair of wings and a pair of legs. A dark stripe known as
the humeral stripe runs from the base of the front wings to the second
pair of legs, and just in front of this is the pale-coloured,
antehumeral stripe.
[6]
The forewings and hindwings are similar in appearance and are
membranous, being strengthened and supported by longitudinal veins that
are linked by many cross-veins and that are filled with
haemolymph.
[13] Species markers include quadrangular markings on the wings known as the
pterostigma or stigma, and in almost all species, there is a
nodus near the leading edge. The thorax houses the flight muscles.
[6]
Many damselflies (e.g. Lestidae, Platycnemidae, Coenagrionidae) have
clear wings, but some (Calopterygidae, Euphaeidae) have coloured wings,
whether uniformly suffused with colour or boldly marked with a coloured
patch. In species such as the banded demoiselle,
Calopteryx splendens
the males have both a darker green body and large dark violet-blue
patches on all four wings, which flicker conspicuously in their aerial
courtship dances; the females have pale translucent greenish wings.
The abdomen is long and slender and consists of ten segments. The
secondary genitalia in males are on the undersides of segments two and
three and are conspicuous, making it easy to tell the sex of the
damselfly when viewed from the side. The female genital opening is on
the underside between segments eight and nine. It may be covered by a
subgenital plate, or extended into a complex ovipositor that helps them
lay eggs within plant tissue. The tenth segment in both sexes bears
cerci and in males, its underside bears a pair of paraprocts.
[6]
Damselflies (except spreadwings,
Lestidae)
rest their wings together, above their bodies, whereas dragonflies rest
with their wings spread diametrically apart; the spreadwings rest with
their wings slightly apart. Damselflies have slenderer bodies than
dragonflies, and their eyes do not overlap. Damselfly nymphs differ from
dragonflies nymphs in that they possess caudal gills (on the abdomen)
whereas dragonflies breathe through the rectum. Damselfly nymphs swim by
fish-like undulations, the gills functioning like a tail. Dragonfly
nymphs can forcibly expel water in their rectum for rapid escape.
[15]
Distribution and diversity
Odonates are found on all the continents except Antarctica.
[16]
Although some species of dragonfly have wide distributions, damselflies
tend to have smaller ranges. Most odonates breed in fresh-water; a few
damselflies in the family Caenagrionidae breed in brackish water (and a
single dragonfly species breeds in seawater).
[17][18]
Dragonflies are more affected by pollution than are damselflies. The
presence of odonates indicates that an ecosystem is of good quality. The
most species-rich environments have a range of suitable microhabitats,
providing suitable water bodies for breeding.
[19][20]
Although most damselflies live out their lives within a short
distance of where they were hatched, some species, and some individuals
within species, disperse more widely.
Forktails in the family
Coenagrionidae seem particularly prone to do this, large male
boreal bluets (
Enallagma boreale) in British Columbia often migrating, while smaller ones do not.
[21]
These are known to leave their waterside habitats, flying upwards till
lost from view, and presumably being dispersed to far off places by the
stronger winds found at high altitudes.
[21] In this way they may appear in a locality where no damselflies were to be seen the day before.
Rambur's forktail (
Ischnura ramburii) has been found, for example, on
oil rigs far out in the
Gulf of Mexico.
[6]
The distribution and diversity of damselfly species in the
biogeographical regions is summarized here. (There are no damselflies in the
Antarctic.) Note that some species are widespread and occur in multiple regions.
[20]
Overall, there are about 2942 extant species of damselflies placed in 309 genera.
[20]
Biology
Adult
damselflies catch and eat flies, mosquitoes, and other small insects.
Often they hover among grasses and low vegetation, picking prey off
stems and leaves with their spiny legs.
[21] Although predominantly using vision to locate their prey, adults may also make use of olfactory cues.
[22]
No species are known to hunt at night, but some are crepuscular,
perhaps taking advantage of newly hatched flies and other aquatic
insects at a time when larger dragonflies are roosting.
[23] In tropical South America, helicopter damselflies (
Pseudostigmatidae) feed on
spiders, hovering near an orb web and plucking the spider, or its entangled prey, from the web.
[24]
There are few pools and lakes in these habitats, and these damselflies
breed in temporary water bodies in holes in trees, the rosettes of
bromeliads and even the hollow stems of
bamboos.
[25]
The nymphs of damselflies have been less researched than their
dragonfly counterparts, and many have not even been identified. They
choose their prey according to size and seem less able to overpower
larger prey than can dragonfly nymphs. The major part of the diet of
most species appears to be
crustaceans such as
water fleas.
[23]
Ecology
Fine damselfly habitat: panorama of
Thursley Common, looking over the acid bog pools
Damselflies exist in a range of habitats in and around the wetlands
needed for their larval development; these include open spaces for
finding mates, suitable perches, open aspect, roosting sites, suitable
plant species for ovipositing and suitable water quality, and odontates
have been used for
bio-indication purposes regarding the quality of the
ecosystem. Different species have different requirements for their larvae with regard to water depth, water movement and pH.
[26] The European
common blue damselfly (
Enallagma cyathigerum) for example can occur at high densities in
acid waters where fish are absent, such as in bog pools. The
scarce blue-tailed damselfly (
Ischnura pumilio) in contrast requires
base-rich
habitats and water with a slow flow-rate. It is found in ditches,
quarries, seeps, flushes, marshes and pools. It tolerates high levels of
zinc and copper in the sediment but requires suitable emergent plants
for egg-laying without the water being choked by plants.
[26]
Damselflies' dependence on freshwater habitats makes them very
vulnerable to damage to wetlands through drainage for agriculture or
urban growth.
[28]
In the tropics, the helicopter damselfly
Mecistogaster modesta (
Pseudostigmatidae) breeds in
phytotelmata, the small bodies of water trapped by
bromeliads,
epiphytic plants of the
rainforest of northwest
Costa Rica, at the high density of some 6000 larvae per hectare in patches of secondary forest.
[29] Another tropical species, the cascade damselfly
Thaumatoneura inopinata (
Megapodagrionidae), inhabits
waterfalls in Costa Rica and Panama.
[30][31]
Damselflies, both nymphs and adults, are eaten by a range of
predators including birds, fish, frogs, dragonflies, other damselflies,
water spiders,
water beetles,
backswimmers and
giant water bugs.
[21]
Damselflies have a variety of internal and external parasites. Particularly prevalent are the
gregarine protozoans
found in the gut. In a study of the European common blue damselfly,
every adult insect was infected at the height of the flying season. When
present in large numbers, these parasites can cause death by blocking
the gut.
[21] Bright red water mites
Hydracarina are often seen on the outside of both nymphs and adults, and can move from one to the other at metamorphosis.
[21]
They suck the body fluids and may actually kill young nymphs, but
adults are relatively unaffected, it being necessary for the completion
of the mite's life cycle that it returns to water, a feat accomplished
when the adult damselfly breeds.
[32]
Behaviour
Female blue-fronted dancer
Many damselflies have elaborate courtship behaviours. These are
designed to show off the male's distinctive characteristics, bright
colouring or flying abilities, thus demonstrating his fitness.
Calopteryx
males will hover in front of a female with alternating fast and slow
wingbeats; if she is receptive she will remain perched, otherwise she
will fly off. The male
river jewelwing (
Calopteryx aequabilis)
performs display flights in front of the female, fluttering his
forewings while keeping his hindwings still, and raising his abdomen to
reveal the white spots on his wings.
Platycypha males will hover in front of a female, thrusting their bright white legs forward in front of their heads.
Rhinocypha
will bob up and down, often low over fast-flowing streams, displaying
their bright-coloured bodies and wings. Male members of the family
Protoneuridae with vividly-coloured wings display these to visiting females.
[34] Swift forktail (
Ischnura erratica) males display to each other with their blue-tipped abdomens raised.
Other behaviours observed in damselflies include wing-warning,
wing-clapping, flights of attrition and abdominal bobbing. Wing-warning
is a rapid opening and closing of the wings and is aggressive, while
wing-clapping involves a slower opening of the wings followed by a rapid
closure, up to eight times in quick succession, and often follows
flight; it may serve a thermo-regulatory function.
[36] Flights of attrition are engaged in by the
ebony jewelwing (
Calopteryx maculata)
and involve males bouncing around each other while flying laterally and
continuing to do so, sometimes over a considerable distance, until one
insect is presumably exhausted and gives up.
At night, damselflies usually roost in dense vegetation, perching
with the abdomen alongside a stem. If disturbed they will move around to
the other side of the stem but will not fly off. Spreadwings fully fold
their wings when roosting.
[6] The
desert shadowdamsel (
Palaemnema domina)
aggregates to roost in thick places near streams in the heat of the
day. While there it engages in wing-clapping, the exact function of
which is unknown. Some species such as the rubyspot damselfly,
Hetairina americana,
form night roosting aggregations, with a preponderance of males; this
may have an anti-predator function or may be simply the outcome of
choosing safe roosting sites.
[39]
Reproduction
Second stage: "heart" or "wheel" position, the female receiving sperm from the male secondary sex organ
Mating in damselflies, as in dragonflies, is a complex, precisely
choreographed process involving both indirect insemination and delayed
fertilisation.
[41]
The male first has to attract a female to his territory, continually
driving off rival males. When he is ready to mate, he transfers a packet
of sperm from his primary genital opening on segment 9, near the end of
his abdomen, to his secondary genitalia on segments 2–3, near the base
of his abdomen. The male then grasps the female by the head with the
claspers at the end of his abdomen; the structure of the claspers varies
between species, and may help to prevent interspecific mating.
[41][42]
The pair fly in tandem with the male in front, typically perching on a
twig or plant stem. The female then curls her abdomen downwards and
forwards under her body to pick up the sperm from the male's secondary
genitalia, while the male uses his "tail" claspers to grip the female
behind the head: this distinctive posture is called the "heart" or
"wheel";
[43] the pair may also be described as being "in cop".
Males may transfer the sperm to their secondary genitalia either before
a female is held, in the early stage when the female is held by the
legs or after the female is held between the terminal claspers. This can
lead to variations in the tandem postures.
[45] The spermatophore may also have nutrition in addition to sperms as a "
nuptial gift".
[46] Some cases of
sexual cannibalism exist where females (of
Ischnura graellsii) eat males while in copula.
[47]
Parthenogenesis (reproduction from unfertilised eggs) is exceptional, and has only been recorded in nature in female
Ischnura hastata on the
Azores Islands.
[20][48]
Yellow-striped blue dart,
Pseudagrion indicum, laying eggs; the male (above) continues to hold the female with his claspers.
Egg-laying (ovipositing) involves not only the female darting over
floating or waterside vegetation to deposit eggs on a suitable
substrate, but the male hovering above her, mate-guarding, or in some
species continuing to clasp her and flying in tandem. The male attempts
to prevent rivals from removing his sperm and inserting their own,
[49] a form of
sperm competition (the sperms of the last mated male have the greatest chance of fertilizing the eggs, also known as sperm precedence
[50]) made possible by delayed fertilisation
[43] and driven by
sexual selection.
[41][42]
If successful, a rival male uses his penis to compress or scrape out
the sperm inserted previously; this activity takes up much of the time
that a copulating pair remain in the heart posture.
Flying in tandem has the advantage that less effort is needed by the
female for flight and more can be expended on egg-laying, and when the
female submerges to deposit eggs, the male may help to pull her out of
the water.
[49]
All damselflies are known to lay their eggs inside plant tissues;
those that lay eggs underwater may submerge themselves for 30 minutes at
a time, climbing along the stems of aquatic plants and laying eggs at
intervals.
[51] For example, the red-eyed damselfly
Erythromma najas lays eggs, in tandem, into leaves or stems of floating or sometimes emergent plants; in contrast, the scarce bluetail
Ischnura pumilio
oviposits alone, the female choosing mostly emergent grasses and
rushes, and laying her eggs in their stems either above or just below
the waterline.
[52] The willow emerald
Chalcolestes viridis
(a spreadwing) is unusual in laying eggs only in woody plant tissue,
choosing thin twigs of trees that hang over water, and scarring the bark
in the process. A possible exception is an apparent instance of
ovo-viviparity, in which
Heliocypha perforata
was filmed in western China depositing young larvae (presumably hatched
from eggs inside the female's body) onto a partly-submerged branch of a
tree.
[54]
Many damselflies are able to produce more than one brood per year (
voltinism); this is negatively correlated with
latitude, becoming more common towards the equator, except in the Lestidae.
[55]
Life cycle
Damselflies are
hemimetabolous insects that have no pupal stage in their development.
[56] The female inserts the eggs by means of her ovipositor into slits made in water plants or other underwater
substrates and the larvae, known as naiads or nymphs, are almost all completely aquatic.
[6] Exceptions include the Hawaiian
Mealagrion oahuense and an unidentified Megapodagrionid from New Caledonia,
[57] which are terrestrial in their early stages.
[49] The
spreadwings
lay eggs above the waterline late in the year and the eggs overwinter,
often covered by snow. In spring they hatch out in the meltwater pools
and the nymphs complete their development before these temporary pools
dry up.
[21]
Blue-tailed damselfly (
Ischnura elegans) nymph, showing the three tail appendages
The nymphs are voracious
predators and feed by means of a flat
labium (a toothed mouthpart on the lower jaw) that forms the so-called mask; it is rapidly extended to seize and pierce the
Daphnia (water fleas),
mosquito
larvae, and other small aquatic organisms on which damselfly nymphs
feed. They breathe by means of three large external, fin-like gills on
the tip of the abdomen, and these may also serve for locomotion in the
same manner as a fish's tail.
[6]
Compared to dragonfly larvae, the nymphs show little variation in form.
They tend to be slender and elongate, many having morphological
adaptations for holding their position in fast flowing water. They are
more sensitive than dragonfly nymphs to oxygen levels and suspended fine
particulate matter, and do not bury themselves in the mud.
[23]
The nymphs proceed through about a dozen moults as they grow. In the
later stages, the wing pads become visible. When fully developed, the
nymphs climb out of the water and take up a firm stance, the skin on the
thorax splits and the adult form wriggles out. This has a soft body at
first and hangs or stands on its empty larval case. It pumps haemolymph
into its small limp wings, which expand to their full extent. The
haemolymph is then pumped back into the abdomen, which also expands
fully. The
exoskeleton
hardens and the colours become more vivid over the course of the next
few days. Most damselflies emerge in daytime and in cool conditions the
process takes several hours. On a hot day, the cuticle hardens rapidly
and the adult can be flying away within half an hour.
[6]
Conservation
Conservation
of Odonata has usually concentrated on the more iconic suborder
Anisoptera, the dragonflies. However, the two suborders largely have the
same needs, and what is good for dragonflies is also good for
damselflies. The main threats experienced by odonates are the clearance
of forests, the pollution of waterways, the lowering of groundwater
levels, the damming of rivers for
hydroelectric schemes and the general degradation of wetlands and marshes.
[58]
The clearance of tropical rainforests is of importance because the rate
of erosion increases, streams and pools dry up and waterways become
clogged with silt. The presence of alien species can also have
unintended consequences.
[58] In Hawaii, the introduction of the
mosquitofish (
Gambusia affinis) was effective in controlling mosquitoes but nearly exterminated the island's endemic damselflies.
[59] The ancient greenling
Hemiphlebia mirabilis has been an important flagship species for conservation action in preserving its habitat in Australia.
[60]
In human culture
Damselfly is a 2005 short film directed by Ben O'Connor.
[61] Damselfly is also the title of a 2012 novel in the
Faeble series by S. L. Naeole,
[62] and of a 1994 poem by
August Kleinzahler,
which contains the lines "And that blue there, cobalt / a moment, then
iridescent, / fragile as a lady's pin / hovering above the nasturtium?"
[63] The poet
John Engels published
Damselfly, Trout, Heron in his 1983 collection
Weather-Fear: New and Selected Poems.
[64]
Fishing flies
that mimic damselfly nymphs are sometimes used in wet-fly fishing,
where the hook and line are allowed to sink below the surface.
[65]
Damselflies have formed subjects for personal
jewellery such as brooches since at least 1880.
[66]
See also
References
Selys-Longchamps, E. (1854). Monographie des caloptérygines (in French). Brussels and Leipzig: C. Muquardt. pp. 1-291 [2]. doi:10.5962/bhl.title.60461.
Sources
 |
Wikimedia Commons has media related to Zygoptera. |
External links
 |
Look up damselfly in Wiktionary, the free dictionary. |
Grimaldi, David; Engel, Michael S. (2005). Evolution of the Insects. Cambridge University Press. pp. 174, 178. ISBN 978-0-521-82149-0.
Jarzembowski, E. A.; A. Nel (2002). "The earliest damselfly like insect and the origin of modern dragonflies (Insecta: Odonatoptera: Protozygoptera)". Proceedings of the Geologists' Association. 113: 165–169. doi:10.1016/s0016-7878(02)80018-9.
Bechly, Günter; Wichard, Wilfried (30 December 2008). "Damselfly and dragonfly nymphs in Eocene Baltic amber (Insecta: Odonata), with aspects of their palaeobiology" (PDF). Palaeodiversity. 1: 37–73.
Dijkstra, Klaas-Douwe B.; Kalkman, Vincent J.; Dow, Rory A.; Stokvis, Frank R.; van Tol, Jan (2013). "Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata)". Systematic Entomology. 39 (1): 68–96. doi:10.1111/syen.12035.
Paulson, Dennis (2011). Dragonflies and Damselflies of the East. Princeton University Press. pp. 10–32. ISBN 1-4008-3966-1.
Gossum,
Hans Van; Sherratt, Thomas N. (2008). "A dynamical model of sexual
harassment in damselflies and its implications for female-limited
polymorphism". Ecological Modelling. 210: 212–200. doi:10.1016/j.ecolmodel.2007.07.023.
Gossum,
Hans Van; Sherratt, Tom N.; Cordero-Rivera, Adolfo (2008). "The
evolution of sex-limited colour polymorphism". In Cordoba-Aguilar, Alex.
Dragonflies and Damselflies. Model organisms for ecological and evolutionary research. Oxford University Press. pp. 219–229.
Kipping, Jens; Martens, Andreas; Suhling, Frank (2012). "Africa's smallest damselfly – a new Agriocnemis from Namibia". Organisms Diversity & Evolution. 12: 301–306. doi:10.1007/s13127-012-0084-4.
Groenevelda, Linn F.; Viola Clausnitzerb; Heike Hadrysa (2007). "Convergent
Evolution of Gigantism in Damselflies of Africa and South America?
Evidence from Nuclear and Mitochondrial Sequence Data". Molecular Phylogenetics and Evolution. 42 (2): 339–46. PMID 16945555. doi:10.1016/j.ympev.2006.05.040.
Hedström, Ingemar; Göran Sahlén (2001). "A
key to the adult Costa Rican "helicopter" damselflies (Odonata:
Pseudostigmatidae) with notes on their phenology and life zone
preferences". Rev. Biol. Trop. 49 (3–4): 1037–1056.
Silsby, Jill (2001). Dragonflies of the World. Csiro Publishing. p. 9. ISBN 978-0-643-10249-1.
Borror, Donald J.; Triplehorn, Charles A.; Triplehorn, Norman F. Study of Insects (6 ed.). Philadelphia: Saunders College Publishing. pp. 187–201.
Nilsson, Anders (1997). Aquatic insects of North Europe: A taxonomic handbook. Apollo Books. p. 14. ISBN 978-87-88757-07-1.
Osburn, Raymond C. (1906). "Observations and Experiments on Dragon-Flies in Brackish Water". The American Naturalist. 40 (474): 395–399. doi:10.1086/278632.
Dunson,
William A. (1980). "Adaptations of Nymphs of a Marine Dragonfly,
Erythrodiplax berenice, to Wide Variations in Salinity". Physiological Zoology. 53 (4): 445–452.
"Introduction to the Odonata". UCMP Berkeley. Retrieved 13 March 2015.
Suhling,
F.; Sahlén, G.; Gorb, S.; Kalkman, V.J.; Dijkstra, K-D.B.; van Tol, J.
(2015). "Order Odonata". In Thorp, James & D. Christopher Rogers. Ecology and general biology. Thorp and Covich's Freshwater Invertebrates (4 ed.). Academic Press. pp. 893–932. ISBN 978-0-12-385026-3.
Acorn, John (2004). Damselflies of Alberta: Flying Neon Toothpicks in the Grass. University of Alberta. pp. 9–15. ISBN 978-0-88864-419-0.
Piersanti,
Silvana; Frati, Francesca; Conti, Eric; Gaino, Elda; Rebora, Manuela;
Salerno, Gianandrea (2014). "First evidence of the use of olfaction in
Odonata behaviour". Journal of Insect Physiology. 62: 26–31. doi:10.1016/j.jinsphys.2014.01.006.
Heckman, Charles W. (2008). Encyclopedia
of South American Aquatic Insects: Odonata - Zygoptera: Illustrated
Keys to Known Families, Genera, and Species in South America. Springer. pp. 17, 31–33. ISBN 978-1-4020-8176-7.
Ingley,
Spencer J.; Bybee, Seth M.; Tennessen, Kenneth J.; Whiting, Michael F.;
Branham, Marc A. (2012). "Life on the fly: phylogenetics and evolution
of the helicopter damselflies (Odonata, Pseudostigmatidae)". Zoologica Scripta. 41 (6): 637–650. doi:10.1111/j.1463-6409.2012.00555.x.
Fincke, Ola M. (2006). "Use of Forest and Tree Species, and Dispersal by Giant Damselflies (Pseudostigmatidae): Their Prospects in Fragmented Forests" (PDF). In Adolfo Cordero Rivera. Fourth WDA International Symposium of Odonatology, Pontevedra (Spain), July 2005. Sofia—Moscow: Pensoft Publishers. pp. 103–125.
Allen, Katherine (2009). The ecology and conservation of threatened damselflies. The Environment Agency. pp. 1–6. ISBN 978-1-84911-093-8.
Corbet, P. S. (1980). "Biology of Odonata". Annual Review of Entomology. 25: 189–217. doi:10.1146/annurev.en.25.010180.001201.
Srivastava, Diane S.; Melnychuk, Michael C.; Ngai, Jacqueline T. (2005). "Landscape
variation in the larval density of a bromeliad-dwelling zygopteran,
Mecistogaster modesta (Odonata: Pseudostigmatidae)". International Journal of Odonatology. 8 (1): 67–79. doi:10.1080/13887890.2005.9748244.
Calvert, Philip P. (1914). "Studies on Costa Rican Odonata. V. The waterfall-dwellers: Thaumatoneura imagos and possible male dimorphism". Entomological News and Proceedings of the Entomological Section. 25 (8): 337–348.
"Thaumatoneura inopinata". The IUCN Red List of Threatened Species. IUCN. Retrieved 16 April 2015.
Proctor, Heather (2004). Aquatic Mites from Genes to Communities: From Genes to Communities. Springer Science & Business Media. pp. 79–84. ISBN 978-1-4020-2703-1.
Silsby, Jill (2001). Dragonflies of the World. Csiro Publishing. pp. 58–59. ISBN 978-0-643-10249-1.
Bick, George H.; Bick, Juanda C. (1961). "Demography and Behavior of the Damselfly, Argia Apicalis (Say), (Odonata: Coenagriidae)". Ecology. 46 (4): 461–472. JSTOR 1934877. doi:10.2307/1934877.
Switzer, Paul V.; Grether, Gregory F. (2000). "Characteristics and Possible Functions of Traditional Night Roosting Aggregations in Rubyspot Damselflies" (PDF). Behaviour. 137: 401–416. doi:10.1163/156853900502141.
Battin, Tom (1993). "The odonate mating system, communication, and sexual selection: A review". Italian Journal of Zoology / Bolletino di zoologia. 60 (4): 353–360. doi:10.1080/11250009309355839.
Cordero-Rivera,
Adolfo; Cordoba-Aguilar, Alex (2010). "Selective Forces Propelling
Genitalic Evolution in Odonata". In Leonard, Janet and Alex
Córdoba-Aguilar. The Evolution of Primary Sexual Characters in Animals (PDF). Oxford University Press. p. 343.
Trueman & Rowe 2009, p. Life Cycle and Behavior.
Bick, G. H.; Bick, J. C. (1965). "Sperm Transfer in Damselflies (Odonata: Zygoptera)". Annals of the Entomological Society of America. 58 (4): 592. PMID 5834678. doi:10.1093/aesa/58.4.592.
Cordero
Rivera, A; Cordoba-Aguilar, A (2010). "Selective forces propelling
genitalic evolution in Odonata". In Leonard J; Cordoba-Aguilar, A. The evolution of primary characters in animals. New York: Oxford University Press. pp. 332–352.
Cordero, Adolfo (1992). "Sexual Cannibalism in the Damselfly Species Ischnura graellsii (Odonata: Coenagrionidae)" (PDF). Entomologia Generalis. 17 (1): 17–20. doi:10.1127/entom.gen/17/1992/17.
Lorenzo-Carballa, M. O.; Cordero-Rivera, A. (2009). "Thelytokous parthenogenesis in the damselfly Ischnura hastata (Odonata, Coenagrionidae): genetic mechanisms and lack of bacterial infection". Heredity. 103 (5): 377–384. PMID 19513091. doi:10.1038/hdy.2009.65.
Cardé, Ring T.; Resh, Vincent H. (2012). A World of Insects: The Harvard University Press Reader. Harvard University Press. pp. 195–197. ISBN 978-0-674-04619-1.
Tennessen, K.J. (2009). "Odonata (Dragonflies, Damselflies)". In Resh, Vincent H. & Ring T. Cardé. Encyclopedia of Insects (2 ed.). Academic Press. pp. 721–729.
Lawlor, Elizabeth P. (1999). Discover Nature in Water & Wetlands: Things to Know and Things to Do. Stackpole Books. pp. 88, 94–96. ISBN 978-0-8117-2731-0.
Smallshire, Dave; Swash, Andy (2014). Britain's
Dragonflies: A Field Guide to the Damselflies and Dragonflies of
Britain and Ireland: A Field Guide to the Damselflies and Dragonflies of
Britain and Ireland. Princeton University Press. pp. 94–96. ISBN 978-1-4008-5186-7.
Salindra,
H. G.; Dayananda, K.; Kitching, Roger L. (2014). "Ovo-viviparity in the
Odonata? The case of Heliocypha perforata (Zygoptera: Chlorocyphidae)".
International Journal of Odonatology. 17 (4): 181–185. doi:10.1080/13887890.2014.959076.
Corbet, Philip S.; Suhling, Frank; Soendgerath, Dagmar (2006). "Voltinism of Odonata: a review" (PDF). International Journal of Odonatology. 9 (1): 1–44. doi:10.1080/13887890.2006.9748261.
A Dictionary of Entomology. CABI. 2011. p. 679. ISBN 978-1-84593-542-9.
Willey, Ruth Lippitt (1955). "A terrestrial damselfly nymph (Medapodarionidae) from New Caledonia" (PDF). Psyche. 62 (4): 137–144. doi:10.1155/1955/39831.
Moore, N.W. (1997). "Dragonflies: status survey and conservation action plan" (PDF). International Union for Conservation of Nature. Retrieved 17 March 2015.
Gagné, W.C. (1981). "Status of Hawaii endangered species: insects and land snails". Elepaio: Journal of the Hawaii Audubon Society. 42: 31–36.
New, Timothy Richard. "The
Hemiphlebia damselfly Hemiphlebia mirabilis Selys (Odonata, Zygoptera)
as a flagship species for aquatic insect conservation in south-eastern
Australia". The Victorian Naturalist. 124 (4): 269–272.
"Ben O'Connor". British Council. Retrieved 13 March 2015.
Naeole, S. L. (2012). Damselfly. Crystal Quill.
Kleinzahler, August (August 1994). "The Damselfly". Poetry Magazine. Retrieved 13 March 2015.
Engels, John (1983). Damselfly, Trout, Heron. Weather-Fear: New and Selected Poems. University of Georgia Press. Retrieved 13 March 2015.
Wada, Wes (2012). "Fishing Tips for the Juicebug Damsel Nymph". Fly foundry. Retrieved 13 March 2015.
Media related to Zygoptera at Wikimedia Commons
Data related to Zygoptera at Wikispecies
No comments:
Post a Comment