Protura
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The Protura, or proturans, and sometimes nicknamed coneheads, are very small (0.6-1.5mm long), soil-dwelling animals, so inconspicuous they were not noticed until the 20th century. The Protura constitute an order of hexapods that were previously regarded as insects, and sometimes treated as a
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
in their own right. Some evidence indicates the Protura are basal to all other hexapods, although not all researchers consider them
Hexapoda The subphylum Hexapoda (from Greek for 'six legs') comprises most species of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura (all of these were once considered inse ...
, rendering the
monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...
of Hexapoda unsettled. Uniquely among hexapods, proturans show
anamorphic Anamorphic format is the cinematography technique of shooting a widescreen picture on standard 35 mm film or other visual recording media with a non-widescreen native aspect ratio. It also refers to the projection format in which a distorted ...
development, whereby
body segment Segmentation in biology is the division of some animal and plant body plans into a series of repetitive segments. This article focuses on the segmentation of animal body plans, specifically using the examples of the taxa Arthropoda, Chordata, an ...
s are added during moults. There are close to 800 species, described in seven
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
. Nearly 300 species are contained in a single
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
, ''
Eosentomon ''Eosentomon'' is a genus of proturans in the family Eosentomidae The Eosentomidae are a family of hexapods in the order Protura The Protura, or proturans, and sometimes nicknamed coneheads, are very small (0.6-1.5mm long), soil-dwelling ani ...
''.


Morphology

Proturans have no eyes, wings, or antennae, and, lacking
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...
ation, are usually white or pale brown. The sensory function of the antennae is fulfilled by the first of three pairs of five-segmented
legs A leg is a weight-bearing and locomotive anatomical structure, usually having a columnar shape. During locomotion, legs function as "extensible struts". The combination of movements at all joints can be modeled as a single, linear element ...
, which are held up, pointing forward and have many tarsal sensilla and sensory hairs. They walk with only four legs. The head is conical, and bears two pseudoculi with unknown function. The body is elongated and cylindrical, with a postanal
telson The telson () is the posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on accou ...
at the end. The mouthparts are entognathous (enclosed within the head capsule) and consist of thin
mandibles 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 ...
and maxillae. There are no cerci at the end of the abdomen, which gives the group their name, 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 ...
''proto-'' (meaning "first", in this case implying primitive), and ''ura'', meaning "tail". The first three abdominal segments bear short limb-like
appendage An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including anten ...
s called "styli". First pair of styli is two-segmented, while the second and third pair are two-segmented or unsegmented. The
genitalia A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...
are internal and the genital opening lies between the eleventh segment and the telson of the adult. The genitalia are everted from a chamber in both sexes. Only the two families
Eosentomidae The Eosentomidae are a family of hexapods in the order Protura The Protura, or proturans, and sometimes nicknamed coneheads, are very small (0.6-1.5mm long), soil-dwelling animals, so inconspicuous they were not noticed until the 20th century. ...
and
Sinentomidae ''Sinentomon'' is the only genus in the family Sinentomidae, in the hexapod order Protura. It contains three species found in China, Japan, and North Korea. Species * ''Sinentomon chui ''Sinentomon chui'' is a species of proturan in the famil ...
possess a simple
tracheal The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends from the la ...
system with a pair of
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
s on both
mesothorax The mesothorax is the middle of the three segments of the thorax of hexapods, and bears the second pair of legs. Its principal sclerites (exoskeletal plates) are the mesonotum (dorsal), the mesosternum (ventral), and the mesopleuron (lateral) on ...
and
metathorax The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites ( exoskeletal plates) are the metanotum (dorsal), the metasternum (ventral), and the metapleuron (lateral) ...
(but never on the abdomen), while those in the other families lack these structures and perform
gas exchange Gas exchange is the physical process by which gases move passively by Diffusion#Diffusion vs. bulk flow, diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liqui ...
by
diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
.


Ecology

Proturans live chiefly in
soil Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former te ...
,
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es, and
leaf litter Plant litter (also leaf litter, tree litter, soil litter, litterfall or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that have fallen to the ground. This detritus or dead organic material and its constituent ...
of moist temperate forests that are not too acidic. They have also been found beneath rocks or under the bark of trees, as well as in animal burrows. They are generally restricted to the uppermost , but have been found as deep as . Although they are sometimes considered uncommon, they are probably often overlooked because of their small size: densities of over 90,000 individuals per square metre have been measured. The diet of proturans is not yet sufficiently observed, but they feed on
mycorrhizal fungi   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
, dead
Acari Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
, and mushroom powder in culture, and are thought to feed on decaying vegetable matter and fungi in the wild. The styliform mouthparts suggest the Protura are fluid feeders, with evidence that some species suck out the contents of fungal hyphae. Proturans which live near the soil surface generally have one generation per year and have longer legs, while those that live deeper have shorter legs and reproduce less seasonally, although some migratory species move to deeper layers for the winter and shallower layers for the summer.


Development

The
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 ty ...
has 8 abdominal segments plus telson, but the number increases through
moulting In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
until the full adult number of 11 is reached. Further moults may occur, but do not add any more body segments, and it is not known whether the adults continue to moult through their lives.
Eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
have only been observed in a few species. Five
developmental stages Stages of development may refer to: Biology *Developmental biology, the study of the process by which animals and plants grow and develop *Prenatal development, also called fetal development, or embryology *Human development (biology), the proc ...
follow: the prenymph hatches from the egg and has only weakly developed mouthparts and 8 abdominal segments; nymph I follows and has fully developed mouthparts; nymph II has 9 abdominal segments; maturus junior has 11 abdominal segments and is followed by the adult. The family Acerentomidae differs in having an extra preimago stage, with partially developed genitalia, between the maturus junior and the adult.


History

Proturans were first discovered in the early 20th century, when
Filippo Silvestri Filippo Silvestri (22 June 1873 – 10 June 1949) was an Italian entomologist. He specialised in world Protura, Thysanura, Diplura and Isoptera, but also worked on Hymenoptera, Myriapoda and Italian Diptera. He is also noted for describing an ...
and
Antonio Berlese Antonio Berlese (26 June 1863, in Padua – 24 October 1927, in Florence) was an Italian entomologist. Career Berlese worked on pest insects notably of fruit trees. He published over 300 articles and a book ''Gli insetti loro organizzazione, svil ...
discovered the animals independently. The first species to be described was '' Acerentomon doderoi'', published in 1907 by Silvestri, based on material from near
Syracuse, New York Syracuse ( ) is a City (New York), city in and the county seat of Onondaga County, New York, Onondaga County, New York, United States. It is the fifth-most populous city in the state of New York following New York City, Buffalo, New York, Buffa ...
.


Impact on humans

Proturans aid in decomposition by helping in the breakdown of leaf litter and recycling organic nutrients back into the soil. They thus play a role in soil formation and composition, which can be vital in soil restoration.


References


External links

* *
Proturans
on the
University of Florida The University of Florida (Florida or UF) is a public land-grant research university in Gainesville, Florida. It is a senior member of the State University System of Florida, traces its origins to 1853, and has operated continuously on its ...
/
Institute of Food and Agricultural Sciences The University of Florida Institute of Food and Agricultural Sciences (UF/IFAS) is a teaching, research and Extension scientific organization focused on agriculture and natural resources. It is a partnership of federal, state, and county governmen ...
''Featured Creatures'' website {{Authority control Arthropod orders Taxa described in 1907 Taxa named by Filippo Silvestri