Psocodea

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Psocodea
Temporal range: 163–0  Ma
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Late Jurassic – Recent
PsocopteraWynaad.jpg
An unidentified bark louse in the family Stenopsocidae
Body lice.jpg
Human body louse
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
(unranked): Paraneoptera
Order: Psocodea
Hennig, 1966
Suborders [1]

Psocodea is a taxonomic group of insects comprising the bark lice, book lice and parasitic lice. [2] It was formerly considered a superorder, but is now generally considered by entomologists as an order. [1] [3] [4] Despite the greatly differing appearance of parasitic lice (Phthiraptera), they are believed to have evolved from within the former order Psocoptera , which contained the bark lice and book lice, now found to be paraphyletic. [5] [6] They are often regarded as the most primitive of the hemipteroids. [7] Psocodea contains around 11,000 species, divided among four suborders and more than 70 families. [1] [2] [8] They range in size from 1–10 millimetres (0.04–0.4 in) in length.

Contents

The species known as booklice received their common name because they are commonly found amongst old books—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. [9] They have chewing mandibles, and the central lobe of the maxilla 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, and three ocelli. Their bodies are soft with a segmented abdomen. [10] Some species can spin silk from glands in their mouth. [11] They may festoon large sections of trunk and branches in dense swathes of silk. [12]

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 pattern, with few cross-veins. The wings, if present, are held tent-like over the body. [10] The legs are slender and adapted for jumping, rather than gripping, as in the true lice. The abdomen has nine segments, and no cerci. [11]

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 setae. The significance of such changes is uncertain, but their function appears to be different from similar variations in, for example, aphids. Like aphids, however, many psocids are parthenogenic, and the presence of males may even vary between different races of the same species. [11]

Psocids lay their eggs in minute crevices or on foliage, although a few species are known to be viviparous. The young are born as miniature, wingless versions of the adult. These nymphs typically molt six times before reaching full adulthood. The total lifespan of a psocid is rarely more than a few months. [11]

Booklice range from approximately 1 mm to 2 mm in length (1/25″ to 1/13″). 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. [13]

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. [14] 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. [15]

Interaction with humans

Some species of psocids, such as Liposcelis bostrychophila , are common pests of stored products. [16] 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. [17]

External phylogeny

Psocodea has been recovered as a monophyletic group in recent studies. Their next closest relatives are traditionally recognized as the monophyletic grouping Condylognatha that contains Hemiptera (true bugs) and Thysanoptera (thrips), which all combined form the group Paraneoptera. However, this is somewhat unclear, as analysis has shown that Psocodea could instead be the sister taxon to Holometabola, which would render Paraneoptera as paraphyletic. [6]

Here is a simple cladogram showing the traditional relationships with a monophyletic Paraneoptera: [6]

Neoptera

Polyneoptera

Eumetabola

Holometabola

Paraneoptera

Psocodea

Condylognatha

Thysanoptera (thrips)

Hemiptera (true bugs)

Here is an alternative cladogram showing Paraneoptera as paraphyletic, with Psocodea as sister taxon to Holometabola: [6]

Neoptera

Polyneoptera

Eumetabola

Holometabola

Psocodea

Condylognatha

Thysanoptera (thrips)  

Hemiptera (true bugs)  

Paraneoptera

Internal phylogeny

Here is a cladogram showing the relationships within Psocodea: [3]

Psocodea
Troctomorpha
Phthiraptera

Philopteridae

Anoplura

Rhynchophthirina

Trichodectidae

Amblycera

Liposcelididae

Pachytroctidae

Sphaeropsocidae

Amphientometae

Psocomorpha

Homilopsocidea

Caeciliusetae

Psocetae

Epipsocetae

Philotarsetae

Archipsocetae

Trogiomorpha

Atropetae

Psyllipsocetae

Prionoglaridetae (paraphyletic)

Classification

The order Psocodea (formerly 'Psocoptera') is divided into three extant suborders.

Suborder Trogiomorpha

Trogiomorpha have antennae with many segments (22–50 antennomeres) and always three-segmented tarsi. [18]

Trogiomorpha is the smallest suborder of the Psocoptera sensu stricto (i.e., excluding Phthiraptera), with about 340 species in 7 families, ranging from the fossil family Archaeatropidae with only a handful of species to the speciose Lepidopsocidae (over 200 species). Trogiomorpha comprises infraorder Atropetae (extant families Lepidopsocidae, Psoquillidae and Trogiidae, and fossil families Archaeatropidae and Empheriidae) and infraorder Psocathropetae (families Psyllipsocidae and Prionoglarididae).

Suborder Troctomorpha

Troctomorpha have antennae with 15–17 segments and two-segmented tarsi.

Troctomorpha comprises the Infraorder Amphientometae (families Amphientomidae, Compsocidae, Electrentomidae, Musapsocidae, Protroctopsocidae and Troctopsocidae) and Infraorder Nanopsocetae (families Liposcelididae, Pachytroctidae and Sphaeropsocidae). Troctomorpha are now known to also contain the order Phthiraptera (lice), and are therefore paraphyletic, as are Psocoptera as a whole.

Some Troctomorpha, such as Liposcelis (which are similar to lice in morphology), are often found in birds' nests, and it is possible that a similar behavior in the ancestors of lice is at the origin of the parasitism seen today. [18]

Suborder Psocomorpha

Psocomorpha are notable for having antennae with 13 segments. They have two- or three-segmented tarsi, this condition being constant (e.g., Psocidae) or variable (e.g., Pseudocaeciliidae) within families. Their wing venation is variable, the most common type being that found in the genus Caecilius (rounded, free areola postica, thickened, free pterostigma, r+s two-branched, m three-branched). Additional veins are found in some families and genera (Dicropsocus and Goja in Epipsocidae, many Calopsocidae, etc.)

Psocomorpha is the largest suborder of the Psocoptera sensu stricto (i.e., excluding Phthiraptera), with about 3,600 species in 24 families, ranging from the species-poor Bryopsocidae (2 spp.) to the speciose Psocidae (about 900 spp). [18] Psocomorpha comprises Infraorder Epipsocetae (families Cladiopsocidae, Dolabellopsocidae, Epipsocidae, Neurostigmatidae and Ptiloneuridae), Infraorder Caeciliusetae (families Amphipsocidae, Asiopsocidae, Caeciliusidae, Dasydemellidae and Stenopsocidae), Infraorder Homilopsocidea (families Archipsocidae, Bryopsocidae, Calopsocidae, Ectopsocidae, Elipsocidae, Lachesillidae, Mesopsocidae, Peripsocidae, Philotarsidae, Pseudocaeciliidae and Trichopsocidae) and Infraorder Psocetae (families Hemipsocidae, Myopsocidae, Psilopsocidae and Psocidae).

Related Research Articles

<span class="mw-page-title-main">Louse</span> Order of insects

Louse 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 developments in phylogenetic research.

<span class="mw-page-title-main">Psocoptera</span> Order of insects

Psocoptera are a paraphyletic group of insects that are commonly known as booklice, barklice or barkflies. The name Psocoptera has been replaced with Psocodea in recent literature, with the inclusion of the former order Phthiraptera into Psocodea.

<span class="mw-page-title-main">Psocomorpha</span> Suborder of booklice

Psocomorpha is a suborder of barklice, booklice, and parasitic lice in the order Psocodea. There are more than 20 families and 5,300 described species in Psocomorpha.

<span class="mw-page-title-main">Trogiomorpha</span> Group of booklice

Trogiomorpha is one of the three major suborders of barklice, booklice, and parasitic lice in the order Psocodea, alongside Troctomorpha and Psocomorpha. There are about 8 families and more than 430 described species in Trogiomorpha. Trogiomorpha is widely agreed to be the earliest diverging of the three suborders, and retains the most primitive characteristics.

<span class="mw-page-title-main">Troctomorpha</span> Suborder of booklice

Troctomorpha is one of the three major suborders of Psocodea (barklice, booklice, and parasitic lice), alongside Psocomorpha and Trogiomorpha. There are more than 30 families and 5,800 described species in Troctomorpha. The order includes parasitic lice, which are most closely related to the booklice family Liposcelididae.

<span class="mw-page-title-main">Psocetae</span> Group of booklice

Psocetae is an infraorder of bark lice in the order Psocodea, within the suborder Psocomorpha. It includes the families Hemipsocidae, Myopsocidae, Psilopsocidae and Psocidae.

<span class="mw-page-title-main">Compsocidae</span> Family of booklice

Compsocidae is a family of Psocodea belonging to the suborder Troctomorpha. The family comprises two extant species in two genera, both found in Mesoamerica. Compsocus elegans is found in Mexico and Central America, while Electrentomopsis variegata is found in Mexico. The antennae of each species have 13 or 14 segments. Two extinct genera, Burmacompsocus and Paraelectrentomopsis are known from the Cenomanian aged Burmese amber of Myanmar and Albian aged Spanish amber.

<span class="mw-page-title-main">Liposcelididae</span> Family of booklice

Liposcelididae is a family of booklice of the order Psocodea, belonging to the suborder Troctomorpha. Members of this family are small and flattened, and often wingless. Mesothorax and metathorax fused in wingless forms.

<span class="mw-page-title-main">Ectopsocidae</span> Family of booklice

Ectopsocidae is a family of Psocodea belonging to the suborder Psocomorpha. The family includes fewer than 200 species, most of them in the genus Ectopsocus.

<span class="mw-page-title-main">Homilopsocidea</span> Group of booklice

Homilopscocidea is an infraorder of Psocodea. It is probably a paraphyletic group, still in use for lack of a better solution. There are about 7 families and more than 1,200 described species in Homilopsocidea.

<span class="mw-page-title-main">Paraneoptera</span> Superorder of insects

Paraneoptera or Acercaria is a superorder of insects which includes lice, thrips, and hemipterans, the true bugs. It also includes the extinct order Permopsocida, known from fossils dating from the Early Permian to the mid-Cretaceous.

<i>Liposcelis</i> Genus of booklice

Liposcelis is a genus of insects in the order Psocoptera, the booklice and barklice. There are about 126 species. Many species are associated with human habitation and several are well known as pests of stored products. The genus is distributed nearly worldwide.

<span class="mw-page-title-main">Epipsocetae</span> Infraorder of booklice

Epipsocetae is an infraorder of psocids in the order Psocodea. There are about 5 families and more than 480 described species in Epipsocetae.

<span class="mw-page-title-main">Ptyctini</span> Tribe of barklice

Ptyctini is a tribe of common barklice in the family Psocidae. There are about 6 genera and at least 20 described species in Ptyctini.

<span class="mw-page-title-main">Psocinae</span> Subfamily of booklice

Psocinae is a subfamily of common barklice in the family Psocidae. There are about 11 genera and at least 40 described species in Psocinae.

<span class="mw-page-title-main">Paracaeciliidae</span> Family of barklice

Paracaeciliidae is a family of bark lice in the order Psocodea. There are about 5 genera and more than 100 described species in Paracaeciliidae.

Spurostigmatidae is a family of lice in the order Psocodea. There is at least one genus, Spurostigma, in Spurostigmatidae.

<span class="mw-page-title-main">Nanopsocetae</span> Infraorder of booklice

Nanopsocetae is one of two major divisions of Troctomorpha in the order Psocodea, alongside Amphientometae. There are more than 20 families and 5,200 described species in Nanopsocetae.

<span class="mw-page-title-main">Amphientometae</span> Infraorder of booklice

Amphientometae is an infraorder of psocids, one of two major division of the Troctomorpha within the order Psocodea. There are about 7 families and at least 230 described species in Amphientometae.

<span class="mw-page-title-main">Philotarsetae</span> Group of booklice

Philotarsetae is an infraorder of bark lice in the order Psocodea, within the suborder Psocomorpha.

References

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