Protura

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Protura
Protura (Acerentomon species) micrograph.jpg
Acerentomon species under stereo microscope
Scientific classificatione
Kingdom: Animalia
Phylum: Arthropoda
Class: Entognatha
Order: Protura
Silvestri, 1907
Families [1]
Acerentomata

Hesperentomidae
Protentomidae
Acerentomidae
Eosentomata

Antelientomidae
Eosentomidae
Sinentomata

Fujientomidae
Sinentomidae
The Protura, or proturans, and sometimes nicknamed coneheads,[2][3] are very small
(<2 mm 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 in their own right.[1][4][5]

Some evidence indicates the Protura are basal to all other hexapods,[6] although
not all researchers consider them Hexapoda, rendering the monophyly of Hexapoda
unsettled.[7] Uniquely among hexapods, proturans show anamorphic development,
whereby body segments are added during moults.[8]

There are close to 800 species, described in seven families. Nearly 300 species are
contained in a single genus, Eosentomon.[1][9]

Contents
1 Morphology
2 Ecology
3 Development
4 History
5 Impact on humans
6 References
7 External links
Morphology
Proturans have no eyes, wings, or antennae, and, lacking pigmentation, are usually
white or pale brown. The sensory function of the antennae is fulfilled by the first
of three pairs of five-segmented legs, which are held up, pointing forward and have
many tarsal sensilla and sensory hairs. They walk with only four legs.[10] The head
is conical, and bears two pseudoculi with unknown function. The body is elongated
and cylindrical,[11] with a postanal telson at the end. The mouthparts are
entognathous (enclosed within the head capsule) and consist of thin mandibles and
maxillae.[8] There are no cerci at the end of the abdomen, which gives the group
their name, from the Greek proto- (meaning "first", in this case implying
primitive), and ura, meaning "tail".[12] The first three abdominal segments bear
limb-like appendages[10] called "styli".[12] The genitalia are internal and the
genital opening lies between the eleventh segment and the telson of the adult.[10]
The genitalia are everted from a chamber in both sexes.[13] Members of Eosentomidae
possess spiracles and a simple tracheal system, while those in the Acerentomoidea
lack these structures and perform gas exchange by diffusion.[10]

Ecology

Protura photographed in Durham, NC

Proturans live chiefly in soil, mosses, and leaf litter[8] of moist temperate
forests[12] that are not too acidic.[14] They have also been found beneath rocks or
under the bark of trees,[11] as well as in animal burrows.[10] They are generally
restricted to the uppermost 0.1 m (3.9 in),[14] but have been found as deep as 0.25
m (9.8 in).[15] Although they are sometimes considered uncommon,[12] they are
probably often overlooked because of their small size:[11] densities of over 90,000
individuals per square metre have been measured.[16]

The diet of proturans is not yet sufficiently observed, but they feed on
mycorrhizal fungi, dead Acari, and mushroom powder in culture,[10] and are thought
to feed on decaying vegetable matter and fungi in the wild.[11][12] The styliform
mouthparts suggest the Protura are fluid feeders, with evidence that some species
suck out the contents of fungal hyphae.[14]

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.[14]

Development
The nymph has 9 abdominal segments, but the number increases through moulting until
the full adult number of 12 is reached. Further moults may occur, but do not add
any more body segments,[12] and it is not known whether the adults continue to
moult through their lives.[10] Eggs have only been observed in a few species.[10]
Five developmental stages follow: the prenymph hatches from the egg and has only
weakly developed mouthparts and 9 abdominal segments; nymph I follows and has fully
developed mouthparts; nymph II has ten abdominal segments; maturus junior has 12
abdominal segments and is followed by the adult.[10] The family Acerentomidae
differs in having an extra preimago stage, with partially developed genitalia,
between the maturus junior and the adult.[10]

History
Proturans were first discovered in the early 20th century, when Filippo Silvestri
and Antonio Berlese discovered the animals independently.[14] The first species to
be described was Acerentomon doderoi, published in 1907 by Silvestri,[10] based on
material from near Syracuse, New York.[12]

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.[17]

References
Andrzej Szeptycki (2007). "Catalogue of the World Protura" (PDF).
"Proturans / Coneheads". North Carolina State University College of Agriculture
and Life Sciences. Archived from the original on May 15, 2008. Retrieved July 30,
2008.
"Order Protura - Coneheads". Iowa State University Department of Entomology.
Retrieved July 30, 2008.
Charles S. Henry (2005). "Insect phylogeny". University of Connecticut. Archived
from the original on 2006-09-05.
Galli, Loris; Shrubovych, Julia; Bu, Yun; Zinni, Matteo (2018). "Genera of the
Protura of the World: diagnosis, distribution, and key". ZooKeys (772): 1–45.
doi:10.3897/zookeys.772.24410.
Ryuichiro Machida (2006). "Evidence from embryology for reconstructing the
relationships of hexapod basal clades" (PDF). Archived from the original (PDF) on
2007-07-20.
Charles E Cook, Qiaoyun Yue & Michael Akam (2005). "Mitochondrial genomes suggest
that hexapods and crustaceans are mutually paraphyletic". Proceedings of the Royal
Society B. 272 (1569): 1295–1304. doi:10.1098/rspb.2004.3042. PMC 1564108. PMID
16024395.
P. J. Gullan & P. S. Cranston (1994). The insects: an outline of entomology.
Chapman and Hall. ISBN 978-0-412-49360-7.
G Pass & NU Szucsich (2011). "100 years of research on the Protura: many secrets
still retained" (PDF).
Christopher Tipping (2004). "Proturans". University of Florida.
"Protura". CSIRO.
John R. Meyer (March 5, 2005). "Protura". North Carolina State University.
Archived from the original on May 15, 2008.
Beutel, Rolf G.; Friedrich, Frank; Ge, Si-Qin; Yang, Xing-Ke (2014). Insect
Morphology and Phylogeny. De Gruyter. p. 184. ISBN 978-3-11-026263-6.
"Gordon's Protura Page". November 11, 2005.
"Protura". Tree of Life Web Project. January 1, 2002.
J. Krauß & W. Funke (1999). "Extraordinary high density of Protura in a windfall
area of young spruce plants". Pedobiologia. 43: 44–46.
Behan-Pelletier, V.M. (1993). "Diversity of soil arthropods in Canada: systematic
and ecological problems". In G.E. Ball; H.V. Danks (eds.). Systematics and
Entomology: Diversity, Distribution, Adaptation and Application. Memoirs of the
Entomological Society of Canada. 165. Entomological Society of Canada. pp. 11–50.
External links
icon Arthropods portal
Data related to Protura at Wikispecies
Media related to Protura at Wikimedia Commons
Proturans on the University of Florida / Institute of Food and Agricultural
Sciences Featured Creatures website
vte
Extant Arthropoda classes by subphylum
Taxon identifiers
Wikidata: Q271631Wikispecies: ProturaADW: ProturaBOLD: 734357BugGuide: 258495EoL:
7116EPPO: 1PROTOFauna Europaea: 12099Fauna Europaea (new): ab8cfaf8-4e20-49a8-a353-
332e1460f3f2GBIF: 1185iNaturalist: 83242IRMNG: 11348ITIS: 99210NBN:
NBNSYS0000160810NCBI: 29999NZOR: 692cd8de-3e6a-40ee-ad23-469ec92762ffPlazi:
03AF87D2-FFA7-FFD8-72C1-FFBE3AAFFDDB
Authority control Edit this at Wikidata
Categories: ProturaArthropod ordersTaxa described in 1907Taxa named by Filippo
Silvestri
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