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Psocoptera are a paraphyletic group 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 ...
s that are commonly known as booklice, barklice or barkflies. The name Psocoptera has been replaced with
Psocodea Psocodea is a taxonomic group of insects comprising the bark lice, book lice and parasitic lice. It was formerly considered a superorder, but is now generally considered by entomologists as an order. Despite the greatly differing appearance of ...
in recent literature, with the inclusion of the former order
Phthiraptera Louse ( : lice) is the common name for any member of the clade Phthiraptera, which contains nearly 5,000 species of wingless parasitic insects. Phthiraptera has variously been recognized as an order, infraorder, or a parvorder, as a result of ...
into Psocodea (as part of the suborder
Troctomorpha Troctomorpha is one of the three major suborders of Psocodea (formerly Psocoptera)(barklice, booklice, and parasitic lice), alongside Psocomorpha and Trogiomorpha. There are more than 30 families and 5,800 described species in Troctomorpha. Th ...
). These insects first appeared in the
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last ...
period, 295–248 million years ago. They are often regarded as the most primitive of the hemipteroids. Their name originates from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
word ψῶχος, ''psokhos'' meaning gnawed or rubbed and πτερά, ''ptera'' meaning
wings A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expre ...
. There are more than 5,500 species in 41 families in three suborders. Many of these species have only been described in recent years. They range in size from 1–10 millimetres (0.04–0.4 in) in length. The species known as booklice received their common name because they are commonly found amongst old
book A book is a medium for recording information in the form of writing or images, typically composed of many pages (made of papyrus, parchment, vellum, or paper) bound together and protected by a cover. The technical term for this physi ...
s—they feed upon the paste used in binding. The barklice are found on trees, feeding on algae and lichen.


Anatomy and biology

Psocids are small, scavenging insects with a relatively generalized body plan. They feed primarily on fungi, algae, lichen, and organic detritus in nature but are also known to feed on starch-based household items like grains, wallpaper glue and book bindings. They have chewing
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
s, and the central lobe of the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. T ...
is modified into a slender rod. This rod is used to brace the insect while it scrapes up detritus with its mandibles. They also have a swollen forehead, large
compound eyes A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distin ...
, and three
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
. Their bodies are soft with a segmented abdomen. Some species can spin silk from glands in their mouth. They may festoon large sections of trunk and branches in dense swathes of silk. Some psocids have small ovipositors that are up to 1.5 times as long as the hindwings, and all four wings have a relatively simple
venation Venation may refer to: * Venation (botany), the arrangement of veins in leaves * Wing venation, the arrangement of veins in insect wings See also * * Vernation Vernation (from ''vernal'' meaning ''spring'', since that is when leaves spring fort ...
pattern, with few cross-veins. The wings, if present, are held tent-like over the body. The legs are slender and adapted for jumping, rather than gripping, as in the true lice. The abdomen has nine segments, and no cerci. There is often considerable variation in the appearance of individuals within the same species. Many have no wings or ovipositors, and may have a different shape to the thorax. Other, more subtle, variations are also known, such as changes to the development of the
seta In biology, setae (singular seta ; from the Latin word for " bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
e. The significance of such changes is uncertain, but their function appears to be different from similar variations in, for example,
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
s. Like aphids, however, many psocids are
parthenogenic Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
, and the presence of males may even vary between different races of the same species. Psocids lay their eggs in minute crevices or on foliage, although a few species are known to be
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the ...
. The young are born as miniature, wingless versions of the adult. These
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label= Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ...
s typically molt six times before reaching full adulthood. The total lifespan of a psocid is rarely more than a few months. Booklice range from approximately 1 mm to 2 mm in length (″ to ″). Some species are wingless and they are easily mistaken for bedbug nymphs and vice versa. Booklouse eggs take two to four weeks to hatch and can reach adulthood approximately two months later. Adult booklice can live for six months. Besides damaging books, they also sometimes infest food storage areas, where they feed on dry, starchy materials. Although some psocids feed on starchy household products, the majority of psocids are woodland insects with little to no contact with humans, therefore they are of little economic importance. They are scavengers and do not bite humans. Psocids can affect the ecosystems in which they reside. Many psocids can affect decomposition by feeding on detritus, especially in environments with lower densities of predacious micro arthropods that may eat psocids. The nymph of a psocid species, '' Psilopsocus mimulus'', is the first known wood-boring psocopteran. These nymphs make their own burrows in woody material, rather than inhabiting vacated, existing burrows. This boring activity can create habitats that other organisms may use.


Interaction with humans

Some species of psocids, such as '' Liposcelis bostrychophila'', are common pests of stored products. Psocids, among other arthropods, have been studied to develop new pest control techniques in food manufacturing. One study found that modified atmospheres during packing (MAP) helped to control the reoccurrence of pests during the manufacturing process and prevented further infestation in the final products that go to consumers.


Classification

In the 2000s, morphological and molecular
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
evidence has shown that the parasitic lice (
Phthiraptera Louse ( : lice) is the common name for any member of the clade Phthiraptera, which contains nearly 5,000 species of wingless parasitic insects. Phthiraptera has variously been recognized as an order, infraorder, or a parvorder, as a result of ...
) evolved from within the psocopteran suborder
Troctomorpha Troctomorpha is one of the three major suborders of Psocodea (formerly Psocoptera)(barklice, booklice, and parasitic lice), alongside Psocomorpha and Trogiomorpha. There are more than 30 families and 5,800 described species in Troctomorpha. Th ...
, thus making Psocoptera paraphyletic with respect to Phthiraptera. In modern systematics, Psocoptera and Phthiraptera are therefore treated together in the order
Psocodea Psocodea is a taxonomic group of insects comprising the bark lice, book lice and parasitic lice. It was formerly considered a superorder, but is now generally considered by entomologists as an order. Despite the greatly differing appearance of ...
.Bess, Emilie, Vince Smith, Charles Lienhard, and Kevin P. Johnson (2006) Psocodea. Parasitic Lice (=Phthiraptera), Book Lice, and Bark Lice. Version 8 October 2006 (under construction). http://tolweb.org/Psocodea/8235/2006.10.08 in The Tree of Life Web Project, http://tolweb.org/ Here is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
showing the relationships within
Psocodea Psocodea is a taxonomic group of insects comprising the bark lice, book lice and parasitic lice. It was formerly considered a superorder, but is now generally considered by entomologists as an order. Despite the greatly differing appearance of ...
, with the former grouping Psocoptera highlighted:


References


External links


National Barkfly Recording SchemePsoco Net

Tree of Life: Psocodea


on the UF / IFAS Featured Creatures Web site {{Authority control Insect orders Paraphyletic groups Paraneoptera