Pseudostigmatidae
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The Pseudostigmatidae are a family of
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
damselflies Damselflies are flying insects of the suborder Zygoptera in the order Odonata. They are similar to dragonflies (which constitute the other odonatan suborder, Epiprocta) but are usually smaller and have slimmer bodies. Most species fold the win ...
, known as helicopter damselflies, giant damselflies, or forest giants. The family includes the largest of all damselfly species. They specialize in preying on web-building
spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
s, and breed in
phytotelmata Phytotelma (plural phytotelmata) is a small water-filled cavity in a terrestrial plant. The water accumulated within these plants may serve as the habitat for associated fauna and flora. A rich literature in German summarised by Thienemann (19 ...
, the small bodies of water held by plants such as
bromeliads The Bromeliaceae (the bromeliads) are a family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the tropical Americas, with several species found in the American subtropics and one in tropical west Africa, ...
.


Range

The species traditionally placed in Pseudostigmatidae are all
Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeogra ...
. Two range as far as northeastern Mexico: ''Mecistogaster ornata'' occurs in
Tamaulipas Tamaulipas, officially the Free and Sovereign State of Tamaulipas, is a state in Mexico; one of the 31 states which, along with Mexico City, comprise the 32 federal entities of Mexico. It is divided into 43 municipalities. It is located in nor ...
and ''Pseudostigma aberrans'' in both Tamaulipas and
Nuevo León Nuevo León, officially the Free and Sovereign State of Nuevo León, is a Administrative divisions of Mexico, state in northeastern Mexico. The state borders the Mexican states of Tamaulipas, Coahuila, Zacatecas, and San Luis Potosí, San Luis ...
. In 2006, molecular phylogenetic analysis confirmed that the African damselfly ''
Coryphagrion grandis ''Coryphagrion grandis'' is a species of damselfly found in coastal forests and on the lower slopes of the Eastern Arc Mountains in Kenya, Tanzania and Mozambique.Clausnitzer, V. 2010''Coryphagrion grandis''.The IUCN Red List of Threatened Species ...
'', previously often classified within
Megapodagrionidae Megapodagrionidae is a family of damselflies. They are commonly called flatwings for their habit of spreading out their wings horizontally when at rest. A 2013 phylogenetic analysis pares down this family into only three genera, and numerous ot ...
or in a monotypic family Coryphagrionidae, belonged within family Pseudostigmatidae, close to genus ''Mecistogaster'', as was proposed already ten years before. This finding suggests that the family dates back to before the breakup of the supercontinent
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
. More recent studies, using a denser taxon sampling, have shown this family was paraphyletic, and that ''C. grandis'' and New World taxa have rather followed a fantastic ecomorphological
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
.


Naiad

As with other damselflies, the young, known as
naiad In Greek mythology, the naiads (; ), sometimes also hydriads, are a type of female spirit, or nymph, presiding over fountains, wells, springs, streams, brooks and other bodies of fresh water. They are distinct from river gods, who embodied ...
s, have gills and live in
fresh water Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salt (chemistry), salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include ...
. The tropical forests where pseudostigmatids live typically have few ponds and lakes, so the water that collects in or on plants is an important habitat. Water-filled
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...
holes and
bromeliad The Bromeliaceae (the bromeliads) are a Family (biology), family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the Tropics, tropical Americas, with several species found in the American subtropics and on ...
tanks are the most dependable and widespread habitats available to pseudostigmatid naiads, and a majority of species use tree holes. Tree hole species are rarely found in bromeliads and vice versa, probably because bromeliad phytotelmata have much higher oxygen content than those in tree holes. One species apparently specializes in
bamboo Bamboos are a diverse group of mostly evergreen perennial plant, perennial flowering plants making up the subfamily (biology), subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family, in th ...
stems that have filled with water after being breached by other insects. In areas where tree holes are uncommon, naiads of a few species can be found in fallen
fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
husks, though these phytotelmata may dry out quickly or be overturned by animals. Fallen
palm Palm most commonly refers to: * Palm of the hand, the central region of the front of the hand * Palm plants, of family Arecaceae ** List of Arecaceae genera **Palm oil * Several other plants known as "palm" Palm or Palms may also refer to: Music ...
bract In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...
s provide an even more unstable habitat, and no pseudostigmatid naiads have ever been found in them. There remain six species known as adults whose larvae have not yet been identified.Fincke, "Use of Forest", 105–108. The naiads are
top predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
s in their
aquatic Aquatic means relating to water; living in or near water or taking place in water; does not include groundwater, as "aquatic" implies an environment where plants and animals live. Aquatic(s) may also refer to: * Aquatic animal, either vertebrate ...
habitats, feeding on the larvae of
mosquito Mosquitoes, the Culicidae, are a Family (biology), family of small Diptera, flies consisting of 3,600 species. The word ''mosquito'' (formed by ''Musca (fly), mosca'' and diminutive ''-ito'') is Spanish and Portuguese for ''little fly''. Mos ...
es and other
flies Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwin ...
,
tadpole A tadpole or polliwog (also spelled pollywog) is the Larva, larval stage in the biological life cycle of an amphibian. Most tadpoles are fully Aquatic animal, aquatic, though some species of amphibians have tadpoles that are terrestrial animal, ...
s, and the naiads of their own and other
odonate Odonata is an order of predatory flying insects that includes the dragonflies and damselflies (as well as the '' Epiophlebia'' damsel-dragonflies). The two major groups are distinguished with dragonflies (Anisoptera) usually being bulkier with l ...
species.


Adult

Adults are exceptionally large for damselflies, with wingspans as high as 19 cm (7.5 in.) reported for ''
Megaloprepus ''Megaloprepus'' is a genus of large Neotropical damselflies in the family Coenagrionidae, commonly known as helicopter damsels. There are four species distributed from Mexico to Ecuador and Peru.Feindt, W., & Hadrys, H. (2022)The damselfly genus ...
'' and body length up to 13 cm (5.1 in.) for '' Pseudostigma aberrans''. The
pterostigma The pterostigma (plural: pterostigmata) is a group of specialized cells in the outer wings of insects, which are often thickened or coloured, and thus stand out from other cells. It is particularly noticeable in dragonflies, but present also in ...
—a thickened, pigmented cell found on the leading edge of the wing in other odonates—is either missing or else modified into a pseudostigma of several cells. In some species the pseudostigma is a large colored spot covering most of the tip of the wing. Mary and William Beebe described the appearance of ''Mecistogaster'' flying in the rainforest of
Guyana Guyana, officially the Co-operative Republic of Guyana, is a country on the northern coast of South America, part of the historic British West Indies. entry "Guyana" Georgetown, Guyana, Georgetown is the capital of Guyana and is also the co ...
:
Spinning through the aisles made by the giant columns of tree-trunks, were curious translucent pin-wheels .... the wing spots revolved rapidly while the rest of the wings became a mere gray haze.
Adult lifespans as long as seven months have been recorded (in ''Megaloprepus'').


Foraging

As far as is known, all pseudostigmatids feed on web-building spiders, which they pluck out of their webs. The only recorded exception to this uniform diet is that they occasionally take wrapped prey from a web rather than the spider who wrapped it. Most
Odonata Odonata is an order of predatory flying insects that includes the dragonflies and damselflies (as well as the '' Epiophlebia'' damsel-dragonflies). The two major groups are distinguished with dragonflies (Anisoptera) usually being bulkier with ...
are generalist predators; the Pseudostigmatidae may be the only true specialist predators in the order.Corbet, 355–357. Spiders provide a large amount of nutrition per unit of prey, and since empty webs are often taken over by other spiders, a pseudostigmatid that learns web locations may be able to take prey from them again and again. They forage in sun flecks and the gaps created by fallen trees, where there is enough light to see spider webs. Several species have been observed searching for webs by flying up one side of a tree and down the other side. When they locate one they hover in front of it. To catch a spider they first fly backward, then quickly fly forward to grab it in their forelegs. Then they back away again and perch to consume the spider, removing the legs before eating the body. Though this careful forward-and-back approach minimizes contact with the web, they often preen after eating to remove any strands that do adhere to them. Pseudostigmatids prey on spiders about 3 to 6 mm in body length, avoiding larger ones. They prefer soft-bodied spiders, which can be eaten more quickly. They have never been observed feeding on ground-dwelling spiders, though they do catch web-building spiders that drop to the ground or retreat to a rolled leaf in an attempt to escape. The complex barrier webs that most '' Nephila'' spiders build protect them from predation by pseudostigmatids.


Reproduction

Some species mate opportunistically while foraging in light gaps. In others, males briefly hold mating territories in sunny gaps, or defend a large phytotelma and mate with females who lay eggs there. All pseudostigmatids have
ovipositor The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
s capable of cutting into plant tissue to insert eggs, which is the most common mode of
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
in damselflies. Yet one species, ''Mecistogaster martinezi'', has been observed apparently "tossing" eggs from its abdomen onto the surface of the water while in flight. This behavior may allow the insect to avoid predators such as spiders from which it would otherwise be unable to escape in the small, confined space of a phytotelma. However, Ola Fincke has suggested that the ''M. martinezi'' female may simply have been using its long abdomen to test for water in the tree hole before alighting to lay eggs—a behavior observed in a related species—and that the eggs seen floating on the water afterward were laid by mosquitoes.


Species list

The family contains the following species:Schorr et al. *'' Anomisma abnorme'' *''
Coryphagrion grandis ''Coryphagrion grandis'' is a species of damselfly found in coastal forests and on the lower slopes of the Eastern Arc Mountains in Kenya, Tanzania and Mozambique.Clausnitzer, V. 2010''Coryphagrion grandis''.The IUCN Red List of Threatened Species ...
'' *'' Mecistogaster amalia'' – Amalia Helicopter *'' Mecistogaster amazonica'' *'' Mecistogaster asticta'' *'' Mecistogaster buckleyi'' – Blue-tipped Helicopter *'' Mecistogaster jocaste'' *'' Mecistogaster linearis'' *'' Mecistogaster lucretia'' *'' Mecistogaster martinezi'' *'' Mecistogaster modesta'' *'' Mecistogaster ornata'' – Ornate Helicopter *'' Mecistogaster pronoti'' – Atlantic Helicopter *''
Megaloprepus caerulatus ''Megaloprepus caerulatus'', also known as the blue-winged helicopter, is a forest giant damselfly of the family Coenagrionidae. Forest giant damselflies were previously recognized as their own family, Pseudostigmatidae. ''M. caerulatus'' is fou ...
'' *''
Microstigma anomalum ''Microstigma anomalum'' is a species of narrow-winged damselfly in the family Coenagrionidae. It is found in South America. References Further reading

* Coenagrionidae Articles created by Qbugbot Insects described in 1842 {{Coenag ...
'' *'' Microstigma maculatum'' *''
Microstigma rotundatum ''Microstigma rotundatum'', the helicopter damselfly, is a species of damselfly belonging to the family Coenagrionidae. Description These damselflies have a long, thin body and large wings with a dense venation. The apex of the forewings has lar ...
'' *'' Pseudostigma aberrans'' *'' Pseudostigma accedens''


Notes


References

* * * * * * * {{Taxonbar, from=Q2462707 Taxa named by Robert John Tillyard Odonata families Obsolete arthropod taxa