Order DIPLURA

Campodea tillyardi

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Order DIPLURA

Compiler and date details

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

Introduction

The Diplura is a cosmopolitan order of narrow-bodied, primitively wingless, entognathous hexapods. It contains some 800 species in nine families. Species vary in size and form from symphylan-like campodeids ‹5 mm in length, to the dermapteran-like japigid Atlasjapyx atlas Chou & Huang in the subfamily Gigasjapyginae (Chou 1984) and which reaches almost 60 mm in length (Chou & Huang 1986).

Diplura have entognathous mouthparts, moniliform antennae, a pair of cerci and, usually, abdominal styli. Compound eyes and ocelli are absent. The cerci are variously developed and range in form from long, filiform and many-segmented to forceps-like and used for grasping prey. Body colour is generally white to yellowish but the posterior segments of some species, such as those with forceps-like cerci, are heavily sclerotised and pigmented.

Taxonomy
The phylogenetic relationships of the Diplura have not been resolved. Kristensen (1991) suggested two entognathan classes: the class Ellipura (= Parainsecta) comprising the orders Protura and Collembola and the class, and order, Diplura. Kukalová-Peck (1991), however, includes the Diplura as entognathans within the Insecta.

Diplura are reported to be represented in the fossil record in the Upper Carboniferous by a species of the genus Testajapyx. Bitsch (1994), however, in a review of the morphological groundplan of the Hexapoda, particularly the Apterygota, considers that this genus should not be attributed to Diplura, but its precise affinities remain to be established.

Paclt (1957) reviewed the world fauna and made major rearrangements of species. His classification, which is almost certainly artificial, has not been adopted entirely by other workers. Further works on the classification of the Diplura are mainly either regional or deal with only sections of the group. They include a monograph on the Campodeidae (Condé 1955a); the Campodeidae of South Africa (Condé 1955b); Campodeidae from caves of New Caledonia (Condé 1980); the Campodeidae of Papua New Guinea (Condé 1982); the Campodeidae of Greece (Condé 1984); the Japygidae of New Zealand (Pagés 1952a); the Parajapyginae of Angola (Pagés 1952b); the Japygidae of South Africa (Pagés 1955); the families and subfamilies of Diplura (Pagés 1959); the Japygoidea of Greece (Pagés 1979); the Japygoidea of North and Central America and the West Indies (Reddell 1983); and the Japygoidea of South America (Reddell 1985).

The Australian fauna is poorly known. There has been little recent work on the fauna and there is clearly a need for extensive collecting and ecological work, and for taxonomic revision. The Diplura section of the Zoological Catalogue of Australia (Houston 1994) was the first comprehensive account of the taxonomic, distributional and biological knowledge of the dipluran fauna of Australia for over 50 years. It provided a baseline and pointed to the many gaps in our knowledge of the fauna. It should also have raised the awareness of the scientific community to the need for a comprehensive taxonomic revision of the Australian Diplura.

Currently five families, Projapygidae, Campodeidae, Japygidae, Heterojapygidae and Parajapygidae are represented. Condé & Pagés (1991) believe that species of the families Anajapygidae and Procampodeidae will be discovered in Australia, but not Dinjapygidae and Evalljapygidae which are known only from the New World. Nine genera and 28 described species have been recorded from Australia but species in existing collections await description and there is little doubt that further collecting will increase significantly the total number of species.

The first native species described from Australia was Japyx longiseta by Silvestri (1908). He subsequently described a further 10 species and one subspecies from Australia (Silvestri 1911, 1928, 1930, 1931, 1947). Other workers were Tillyard (1924), who described one species; Womersley (1934, 1935, 1937, 1939, 1945), who described 14 species and two subspecies; Pagés (1952a), who, in his review of the world fauna was the most recent worker to list all the described species in Australia, renamed one of Womersley's subspecies. A European species, Campodea fragilis, which has been introduced into Australia, was described by Meinert (1865). Condé & Pagés (1991) gave a key to those families found or likely to be found in Australia and summarised the status of our knowledge of the fauna.

Biology
Diplura occur mainly in moist environments but they are also known from arid regions. They may be found in the soil, under rocks, in leaf litter, under bark of dead or dying trees and in rotten wood. Cavernicolous species are also known. They may be separated from litter or soil by the use of Berlese funnels, Winkler-Moczarski apparatus or by flotation (e.g. Southwood 1966, Upton 1991).

Van der Drift (1951), in a study of a beech forest in temperate regions in Europe, found the highest densities of campodeids (›100 per square metre) at the base of old litter in the humus/mineral soil layer. In tropical rainforest, campodeids were found to be more common in the litter layer while japygids were more abundant in the soil (Manton 1972). Price & Benham (1977) found japygids usually below a depth of 30 cm and the depth at which most specimens were collected was over 100 cm. Failure to sample deeply enough could account for the apparent rarity of Diplura.

Diplura are generally carnivorous and feed on soil micro-organisms but some species are herbivorous and are thought to feed on plant roots and organic detritus. Their economic significance as pests requires investigation. Zimmerman (1948) reported that Parajapyx isabellae (Grassi), which is known from many parts of the world but not Australia, fed on plant roots and caused damage to the root cortex of sugar cane. Rusek (1982) reported that Octostigma herbivora Rusek (a new genus and species in Projapygoidea, for which he created the new family Octostigmatidae) was collected from injured roots of peanuts and melons in Tonga. The gut of this species was filled with a mass of plant remains, including clearly visible cells and parenchymatous tissue.

Postembryonic development of Dipljapyx humberti (Grassi) was studied in detail by Gyger (1960) and Pagés (1967). Smith (1961) discussed the post embryonic development of Diplura and gave observations on the development of species of Parajapyginae and Evalljapyginae. Townsend (1970) figured an egg cluster of Heterojapyx novaezeelandie and gave brief notes about the adults and their associated cluster.

The external abdominal morphology of all known types of Diplura was studied in considerable detail by Pagés (1989). The skeletal anatomy of Heterojapyx was described by Snodgrass (1952) and the head structure and endognathy in Campodea was investigated by Manton (1964). Martin (1969) studied the anatomy of the cerci of Projapygidae and showed how the secretions from the cercal glands assisted in capturing prey.

Observations on the habits and habitats of Diplura were included in comparative studies of hexapod locomotory mechanisms by Manton (1972). Pagés (1967), in a detailed study of Dipljapyx humberti, provided data on the biology, ecology and behaviour of Japygiodea, while Reddell (1983) briefly summarised the general behaviour of the Japygoidea, with emphasis on the North American fauna. Both works are useful sources of references on the Diplura.

Acknowledgements

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. The author would like to thank Dr B. Condé of the University of Nancy and Dr J. Pagés, Dijon, France, for comments on the manuscript; Graeme Smith, of Bayer Australia Ltd, for locating type material and providing additional information deposited at the IEA; 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 Diplura 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.