Australian Biological Resources Study

Australian Faunal Directory




Regional Maps



Compiler and date details

31 December 1993 - W.W.K. Houston, Australian Biological Resources Study, Canberra, Australian Capital Territory, Australia


Protura are very small (<2 mm in length), elongate, delicate and usually unpigmented hexapods. They are known from all zoogeographic regions. Primitively wingless, they lack eyes, antennae and cerci, and the fore legs are modified for sensory purposes. Their mouth-parts are adapted for sucking and they have been observed feeding on both mycorhizoid and free soil hyphae (Sturm 1959; Nosek 1973).

Protura may be quite common, but are rarely seen. They are cryptic and occur in damp situations, such as in soil, moss, peat, leaf litter, and under bark and decaying wood. They are associated with high levels of organic matter and are more commonly recorded from forest habitats. Raw (1956) reported that they are a significant part of the arthropod fauna of agricultural soils. They can be separated from litter or soil by the use of Tullgren funnels or by flotation (e.g. Southwood 1966; Upton 1991).

Juveniles resemble adults and development is anamorphic. Bernard (1976) provided observations on eggs and embryology and showed that the prelarva is the first postembryonic stage. Aldaba (1985) described some prelarvae from Spain and Bernard & Tuxen (1987) provided a key to the families of proturan juveniles and described each stage.

The order was first recognised by Silvestri (1907), when he described Acerentomon doderoi and related it to the insects. Berlese (1909), in a monograph on the order, considered that Protura are closely related to the myriapods and called them Myrientomata. The class and order Protura is now placed, as the sister group to the Collembola, in the Ellipura (=Parainsecta) which is recognised as the sister group to the Insecta (Kristensen 1991; Kukalová-Peck 1991).

Imadaté (1991), based on the views of Tuxen (1963, 1964) and Imadaté (1966), divided the Protura into two superfamilies, the Eosentomoidea and the Acerentomoidea, each with two families. This classification is followed in the Catalogue.

Nosek (1973) erected a new suborder, Sinentomoidea, for the aberrant Chinese Proturan Sinentomon erythranum Yin. Tuxen (1977b), however, discussed the phylogenetic position of Sinentomon and considered that it belongs in the Protentomidae.

Yin (1983, 1984) considered the Acerentomoidea, which she divided into eight families, were the primitive group and the Eosentomoidea the more specialised group. Although Yin's classification is not used by Imadaté (1991), Imadaté states that it is supported by recent ultrastructure studies on proturan spermatozoa.

Yin (1992) and Yin & Xué (1993) proposed a new classification of the proturan families, again not followed by Imadaté (1994) as yet another classification was in preparation by Yin (Imadaté pers. comm.).

Phylogenetic trends in the Protura were discussed by Tuxen (1963) and information on Protura was consolidated by Tuxen (1964) in his monograph on the world fauna. In this work, he revised the phylogeny and also gave keys to the species described up to 1961. Manton discussed the systematics of Protura in her studies on mandibular mechanisms (1964) and on locomotory mechanisms (1972). Francois (et al. 1992) examined the cephalic anatomy of S. erythranum and compared them with other Protura and Apterygota.

Descriptive and other accounts of the proturan fauna of the world are mainly by country or region. Some of these areas, and works which describe their fauna and/or give a useful lead into the relevant literature, include: catalogue of the world fauna (Paclt 1955); monograph on the world fauna (Tuxen 1964); keys and diagnoses to world genera (Nosek 1978); Hawaii (Zimmerman 1948); Portugal (Cunha 1952); Japan (Imadaté & Yosii 1959; Imadaté 1964, 1974, 1994); Uganda (Condé 1961a); SE Asia (Imadaté 1965); Australia (Tuxen 1967); Europe (Nosek 1973); Bismarck Archipelago and Solomon Islands (Tuxen & Imadaté 1975a); Brazil (Tuxen 1976); Angola and Nigeria (Tuxen 1977a); Kermadec Islands (Ramsey & Tuxen 1978); Madeira (Tuxen 1982); New Zealand (Tuxen 1985); N America—Eosentomon (Bernard 1985, 1990); Korea (Lee & Rim 1988); Poland (Szeptycki & Weiner 1990); and China (Yin 1992).

The presence of Protura in Australia was first reported by Dakin & Fordham (1926) but the samples were lost and their identification was not confirmed. Six years later, Womersley (1932) described six new species. He described another species in 1936. In a review of the Australian fauna, Womersley (1939) described a subspecies (later raised to species level), identified an Australian species—later accorded separate species status by Bonet (1942)—as a North American species, and gave a key to the families and subfamilies of Protura. The major worker on the Australian fauna, however, was Tuxen, who reviewed the fauna and described a further 16 species (Tuxen 1967). Tuxen & Imadaté (1975b) described a further two species. Tuxen (1967) and Nosek (1973) discussed the distinct zoogeographical position of the Australian Protura.

Several species, probably introduced into Australia, were described from material collected outside Australia. They are Acerentulus confinis (Berlese 1908), Gracilentulus gracilis (Berlese 1908) and Protentomon perpusillum (Berlese 1909) described from Italy; and Berberentulus capensis (Womersley 1931) described from South Africa. Prabhoo (1960) inadvertently published details of Baculentulus breviunguis (Prabhoo), a species known from India, that was subsequently described by Condé (1961b).

The proturan fauna of Australia is poorly known. Three families, 10 genera and 32 described species are recorded from Australia; at least four of the species are probably introduced. There has been little recent work on the fauna and extensive collecting, ecological work, and taxonomic revision is needed. Species in existing collections await description and undoubtedly further collecting will increase significantly the total number of species recorded from Australia.


Preparation of this database, a section of the Zoological Catalogue of Australia, was undertaken as part of the author's work within the Zoological Catalogue Section of the Australian Biological Resources Study. I would like to thank Professor G. Imadaté and Professor E. C. Bernard for comments on the manuscript; Ms P. Greenslade for assistance with bibliographic and distribution data; the ANIC for access to types; and the librarians at the CSIRO Black Mountain Library for their help with locating many references.

Database Notes

The information on the Australian Faunal Directory site for the Protura is derived from the Zoological Catalogue of Australia database compiled on the Platypus software program. It incorporates changes made to the work published on 21 November 1994 as (Houston, W.W.K., 1994)

Limital Area

Distribution data in the Directory is by political and geographic region descriptors and serves as a guide to the distribution of a taxon. For details of a taxon's distribution, the reader should consult the cited references (if any) at genus and species levels.

Australia is defined as including Lord Howe Is., Norfolk Is., Cocos (Keeling) Ils, Christmas Is., Ashmore and Cartier Ils, Macquarie Is., Australian Antarctic Territory, Heard and McDonald Ils, and the waters associated with these land areas of Australian political responsibility. Political areas include the adjacent waters.

Terrestrial geographical terms are based on the drainage systems of continental Australia, while marine terms are self explanatory except as follows: the boundary between the coastal and oceanic zones is the 200 m contour; the Arafura Sea extends from Cape York to 124 DEG E; and the boundary between the Tasman and Coral Seas is considered to be the latitude of Fraser Island, also regarded as the southern terminus of the Great Barrier Reef.

Distribution records, if any, outside of these areas are listed as extralimital. The distribution descriptors for each species are collated to genus level. Users are advised that extralimital distribution for some taxa may not be complete.


General References

Aldaba, J. 1985. Contribución al conocimiento de las prelarvas de los Proturos Ibéricos. Actas do Congresso Iberico de Entomologia 2: 177-183

Berlese, A. 1908. Nuovi Acerentomidi. Redia. Giornale di entomologia, Firenze 5: 16-19 pl. I

Berlese, A. 1909. Monografia dei Myrientomata. Parte I. Sistematica e morfologia esterna. Redia. Giornale di entomologia, Firenze 6: 1-182 [Pt I pp. 1-59 29 May 1909; Pt II pp. 60-182 14 Aug. 1909]

Bernard, E.C. 1976. Observations on the eggs of Eosentomon australicum (Protura: Eosentomidae). Transactions of the American Microscopical Society 95: 129-130

Bernard, E.C. 1985. Two new species of Protura (Insecta) from North America. Proceedings of the Biological Society of Washington 98: 72-80

Bernard, E.C. 1990. New species, clarifications, and changes in status with Eosentomon Berlese (Hexapoda: Protura: Eosentomidae) from the United States. Proceedings of the Biological Society of Washington 103: 861-890

Bernard, E.C. & Tuxen, S.L. 1987. Class and Order Protura. pp. 47-54 in Stehr, F.W. (ed.). Immature Insects. Dubuque, Iowa : Kendall/Hunt.

Bonet, F. 1942. Sobre algunos Prouros de Mexico (Nota preliminar). Ciencia (Mexico) 3: 14-17

Condé, B. 1961a. II Protures. British Museum (Natural History) Ruwenzori Expedition 1952 2(11): 69-80 [Jan. 1961]

Condé, B. 1961b. Un protoure nouveau a distribution Indo-Madécasse (Acerentulus breviunguis n. sp.). Bulletin du Muséum d'Histoire Naturelle. Paris 2 33: 318-321

Cunha, A.X da 1952. Quelques protures inédits de la fane portugaise. Mem. Estud. Mus. Zool. Univ. Coimbra No. 212. 15 pp. 7 figs

Dakin, W.J. & Fordham, M.G.C. 1926. Birth of Peripatus in England. Nature (London) 117(2955): 858 [Letter]

Francois, J., Dallai, R. & Yin, W.Y. 1992. Cephalic anatomy of Sinetomon erythranum Yin (Protura: Sinetomidae). International Journal of Insect Morphology and Embryology 21(3): 199-213

Houston, W.W.K. 1994. Protura. 1-18, 157-164 (Appendix & Index) in Houston, W.W.K. (ed.). Zoological Catalogue of Australia. Protura, Collembola, Diplura. Melbourne : CSIRO Australia Vol. 22 188 pp.

Imadaté, G. 1964. Taxonomic arrangement of Japanese Protura (I). Bulletin of the National Science Museum, Tokyo 7: 37-81

Imadaté, G. 1965. Proturans-fauna of Southeast Asia. pp. 195-302 in Kira, T. & Iwata, K. (eds). Nature and Life in Southeast Asia. Kyoto : Fauna and Flora Research Society.

Imadaté, G. 1966. Taxonomic arrangement of Japanese Protura (IV). Bulletin of the National Science Museum, Tokyo 7: 37-81

Imadaté, G. 1974. Protura (Insecta). Fauna Japonica Tokyo : Keigaku Publishing Co. Ltd 351 pp.

Imadaté, G. 1991. Protura. pp. 265-268 in CSIRO (ed.). The Insects of Australia. A textbook for students and research workers. Melbourne : Melbourne University Press Vol. 1 xiii 542 pp.

Imadaté, G. 1994. Contributions towards a revision of the Proturan fauna of Japan (IX). Collecting data of acerentomid and sinetomid species in the Japanese Islands. Bulletin of the Department of General Education, Tokyo Medical and Dental University 24: 45-70

Imadaté, G. & Yosii, R. 1959. A synopsis of the Japanese species of Protura. Contributions from the Biological Laboratory Kyoto University 6: 1-43

Kristensen, N.P. 1991. Phylogeny of extant hexapods. pp. 125-140 in CSIRO (ed.). The Insects of Australia. A textbook for students and research workers. Melbourne : Melbourne University Press Vol. 1 xiii 542 pp.

Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. pp. 141-179 in CSIRO (ed.). The Insects of Australia. A textbook for students and research workers. Melbourne : Melbourne University Press Vol. 1 xiii 542 pp.

Lee, B.H. & Rim, M.G. 1988. Acerentomid proturans (Insecta), with two new species and two new records for Korea. Korean Journal of Systematic Zoology 4(1): 1-11

Manton, S.M. 1964. Mandibular mechanisms and the evolution of arthropods. Philosophical Transactions of the Royal Society of London B 247: 1-183

Manton, S.M. 1972. The evolution of arthropodan locomotory mechanisms. Part 10. Locomotory habits, morphology and evolution of the hexapod classes. Journal of the Linnean Society of London, Zoology 51: 203-400

Nosek, J. 1973. The European Protura. Their taxonomy, ecology and distribution with keys for determination. Geneva : Museum d'Histoire Naturelle 345 pp.

Nosek, J. 1978. Key and diagnoses of Proturan genera of the world. Annotationes Zoologicae et Botanicae, Bratislava 122: 1-59

Paclt, J. 1955. Protura. Genera Insectorum 211: 1-123

Prabhoo, N.R. 1960. Acerentulus breviunguis Condé (Protura: Acerentomidae) from India. Current Science. Bangalore 29: 273-274

Ramsay, G.W. & Tuxen, S.L. 1978. Protura (Insecta) from the Kermadec islands, and a redescription of Acerentulus nemoralis from Argentina. New Zealand Journal of Zoology 5: 601-606

Raw, F. 1956. The abundance and distribution of Protura in grassland. Journal of Animal Ecology 25: 15-21

Silvestri, F. 1907. Descrizione di un novo genere d'insetti apterigoti rappresentante di un novo ordine. Bollettino del Laboratorio di Zoologia Generale e Agraria della Facoltà Agraria in Portici 1: 296-311

Southwood, T.R.E. 1978. Ecological methods with particular reference to the study of insect populations. London : Chapman & Hall xxiv 524 pp.

Stumpp, J. 1990. Zur okologie einheimischer Proturen (Arthropoda: Insecta) in Fichtenforsten. Zoologische Beiträge. Berlin N.F. 33(3): 345-432

Sturm, H. 1959. Die Nahrung der Proturen. Beobachtungen an Acerentomon doeroi Silv. und Eosentomon transitorum Berl. Naturwissenschaften 46: 90-91

Szeptycki, A. 1988. New genera and species of Protura from Altai Mts. Acta Zoologica Cracoviensia 31(7): 297-362

Szeptycki, A. & Weiner, W.M. 1990. 1. Protura—Peirwogonki. pp. 15-16 in Razowski, J. (ed.). Checklist of animals in Poland. Wyd. : PAN Vol. 1 Pt xxxii.

Tuxen, S.L. 1963. Phylogentical trends in the Protura as shown by relationship between recent genera. Zeitschrift für Zoologische Systematik und Evolutionsforschung 1: 277-310

Tuxen, S.L. 1964. The Protura. A revision of the species of the world with keys for determination. Actualités Sci. Industr. No. 1311 Problèmes d'écologie. Paris : Hermann 360 pp.

Tuxen, S.L. 1967. Australian Protura, their phylogeny and zoogeography. Zeitschrift für Zoologische Systematik und Evolutionsforschung 5: 1-53

Tuxen, S.L. 1970. The systematic postition of entognathous apterygotes. Anales de la Escuela Nacional de Ciencias Biologicas, Mexico 17(1968): 65-79 [30 Mar. 1970]

Tuxen, S.L. 1976. The Protura (Insecta) of Brazil, especially Amazonas. Amazoniana 5(4): 417-463

Tuxen, S.L. 1977a. Protura (Insecta) from Angola and Nigeria with a key to subsaharan Protura. Publicaçoes Culturais da Companhia de Diamantes de Angola 89: 177-193

Tuxen, S.L. 1977b. The systematical position of Simentomon (Insecta: Protura). Bulletin of the National Science Museum Series A Zool. 3: 25-36

Tuxen, S.l. 1982. The Protura (Insecta) of Madeira. Bocagiana 65: 1-20

Tuxen, S.L. 1985. Fauna of New Zealand Number 9. Protura (Insecta). Wellington : Department of Scientific and Industrial Research 80 pp.

Tuxen, S.L. & Imadaté, G. 1975a. The Protura of the Bismarck Archipelago and Solomon Islands. Bulletin of the British Museum (Natural History) Entomology 31: 331-375

Tuxen, S.L. & Imadaté, G. 1975b. Corrections to S.L. Tuxen's "Australian Protura" (1967). Journal of the Australian Entomological Society 14: 193-195

Womersley, H. 1931. A South African species of Protura. Annals of the South African Museum 30: 89-91

Womersley, H. 1932. A preliminary account of the Protura of Australia. Proceedings of the Linnean Society of New South Wales 57: 69-75

Womersley, H. 1936. A new species of Protura from Australia. Entomologist's Monthly Magazine 72: 65-66

Womersley, H. 1939. Protura. pp. 279-289 in Womersley, H. (ed.). Primitive Insects of South Australia. Silverfish, springtails and their allies. Adelaide : Frank Trigg, Government Printer.

Yin, W.Y. 1983. Grouping the known genera of Protura under eight families with keys for determination. Contributions from the Shanghai Institute of Entomology 3: 151-163 [in Chinese]

Yin, W.Y. 1984. A new idea on phylogeny of Protura with approach to its origin and systematic position. Scientia Sinica B 27(2): 149-160

Yin, W.Y. 1992. Protura. pp. 395-414 in Yin, W.Y. (ed.). Subtropical soil animals of China. Beijing : Science Press. [in Chinese]

Yin, W.Y. & Xué, L. 1993. Comparative spermatology of Protura and its significance on Proturan systematics. Scientia Sinica B 36(5): 575-589

Zimmerman, E.C. 1948. Order Protura Silvestri, 1907. pp 42–43 in, Insects of Hawaii. A manual of the Insects of the Hawaiian Island, including and enumeration of the species and notes on their origin, distribution, hosts, parasites, etc. Honolulu : University of Hawaii Press Vol. 2 Apterygota to Thysanoptera.


History of changes

Note that this list may be incomplete for dates prior to September 2013.
Published As part of group Action Date Action Type Compiler(s)
07-Aug-2012 07-Aug-2012 MOVED
07-Aug-2012 07-Aug-2012 MOVED
12-Feb-2010 (import)