Megaloptera
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Megaloptera is an
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s. It contains the alderflies, dobsonflies and fishflies, and there are about 300 known
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
. The order's name comes from
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...
, from ''mega-'' (μέγα-) "large" + ''pteryx'' (πτέρυξ) "wing", in reference to the large, clumsy wings of these insects. Megaloptera are relatively unknown insects across much of their range, due to the adults' short lives, the aquatic larvae's often-high tolerance of
pollution Pollution is the introduction of contaminants into the natural environment that cause adverse change. Pollution can take the form of any substance (solid, liquid, or gas) or energy (such as radioactivity, heat, sound, or light). Pollutants, th ...
(so they are not often encountered by swimmers etc.), and the generally crepuscular or nocturnal habits. However, in the
Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North America, North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. ...
the dobsonflies are rather well known, as their males have tusk-like mandibles. These, while formidable in appearance, are relatively harmless to humans and other animals; much like a
peacock Peafowl is a common name for three bird species in the genera '' Pavo'' and '' Afropavo'' within the tribe Pavonini of the family Phasianidae, the pheasants and their allies. Male peafowl are referred to as peacocks, and female peafowl are r ...
's feathers, they serve mainly to impress females. However, the mandibles are also used to hold females during mating, and some male dobsonflies spar with each other in courtship displays, trying to flip each other over with their long mandibles. Dobsonfly larvae, commonly called hellgrammites, are often used for angling
bait Bait may refer to: General * Bait (luring substance), bait as a luring substance ** Fishing bait, bait used for fishing Film * ''Bait'' (1950 film), a British crime film by Frank Richardson * ''Bait'' (1954 film), an American noir film by Hugo ...
in North America. The Megaloptera were formerly considered part of a group then called Neuroptera, together with lacewings and
snakeflies Snakeflies are a group of predatory insects comprising the order Raphidioptera with two extant families: Raphidiidae and Inocelliidae, consisting of roughly 260 species. In the past, the group had a much wider distribution than it does now; sn ...
, but these are now generally considered to be separate orders, with Neuroptera referring to the lacewings and relatives (which were formerly called Planipennia). The former Neuroptera, particularly the lacewing group, are nonetheless very closely related to each other, and the new name for this group is
Neuropterida The Neuropterida are a clade, sometimes placed at superorder level, of holometabolous insects with over 5,700 described species, containing the orders Neuroptera (lacewings, antlions), Megaloptera (alderflies, dobsonflies), and Raphidiopter ...
. This is either placed at superorder rank, with the Endopterygota—of which they are part—becoming an unranked clade above it, or the Endopterygota are maintained as a superorder, with an unranked Neuropterida being a part of them. Within the endopterygotes, the closest living relatives of the neuropteridan clade are the
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s. The Asian dobsonfly '' Acanthacorydalis fruhstorferi'' can have a wingspan of up to , making it the largest aquatic insect in the world by this measurement.


Anatomy and life cycle

Adult megalopterans closely resemble the lacewings, except for the presence of a pleated region on their hindwings, helping them to fold over the abdomen. They have strong mandibles and mouthparts apparently adapted for chewing, although many species do not eat as adults. They have large compound eyes, and, in some species, also have ocelli. The wings are large and subequal. The female may lay up to 3,000 eggs in a single mass, placing them on vegetation overhanging water. Megaloptera undergo the most rudimentary form of
complete metamorphosis Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygota. ...
among the insects. There are fewer differences between the larval and adult forms of Megaloptera than in any other order of holometabolous insects, and their aquatic
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e dwell in fresh water, around which the adults also live. The larvae are carnivorous, and are known to feed on small invertebrates, such as crustaceans, clams, worms and other insects. They possess strong jaws that they use to capture their prey. They have large heads and elongated bodies. The abdomen bears a number of fine tactile filaments, which, in some species, may include
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they ar ...
s. The final segment of the abdomen bears either a pair of prolegs, or a single, tail-like appendage. The larvae grow slowly, taking anywhere from 1 to 5 years to reach the last larval stage. When they reach maturity, the larvae crawl out onto land to
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
te in damp soil or under logs. Unusually, the pupa is fully motile, with large mandibles that it can use to defend itself against predators. The short-lived adults emerge from the pupa to mate - many species never feed as adults, living only a few days or hours.


Evolution

Apart from the two living families, there are a few prehistoric taxa in the Megaloptera, only known from
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s. Some of these occupy a more basal position: * Genus '' Corydasialis'' (sometimes considered
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...
family Corydasialidae) * Family Parasialidae (probably paraphyletic) * Family Euchauliodidae * Family Nanosialidae The Megaloptera are monophyletic and are a sister clade of the
Neuroptera The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megaloptera and Raphidioptera in th ...
.Wang, Y.; Liu, X.; Winterton, S. L. & Yang, D. (2012). "The first mitochondrial genome for the fishfly subfamily Chauliodinae and implications for the higher phylogeny of Megaloptera". ''PLoS One''. 7 (10): e47302. Within the Megaloptera, Corydalinae and
Chauliodinae Fishflies are members of the subfamily Chauliodinae, belonging to the megalopteran family Corydalidae. They are most easily distinguished from their closest relatives, dobsonflies, by the jaws (mandibles) and antennae. In contrast to the large j ...
are sister clades. The oldest fossils confidently identifiable as megalopterans date to the Early
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
.


Footnotes


References

* * (2005): ''Evolution of the Insects''.
Cambridge University Press Cambridge University Press is the university press of the University of Cambridge. Granted letters patent by King Henry VIII in 1534, it is the oldest university press in the world. It is also the King's Printer. Cambridge University Pr ...
. * (2007): The neuropterid fauna of Dominican and Mexican amber (Neuropterida, Megaloptera, Neuroptera). ''American Museum Novitates'' 3587: 1-58
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{{Authority control Insect orders Cisuralian first appearances Extant Permian first appearances Neuropterida