Higher Taxon Termitoidae

Isoptera

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Higher Taxon Termitoidae

Termites, White Ants

Compiler and date details

February 2017 - A, Wells, CSIRO (Intrioduction modified to reflect changes in classification)

31 December 1996 - J.A.L. Watson, L.R. Miller & H.M. Abbey, CSIRO Entomology, Canberra, Australian Capital Territory, Australia

Isoptera.

Introduction

[Modified after Watson et al. 1998; classification modified to reflect Inward et al. 2007].
'Termitoidae', termites or, coloquially, white ants, are a small group of social insects, a branch of specialised cockroaches. The most appropriate placement of the order among other groups of blattoid–orthopteroid insects was, until recently (Inward et al. 2007), the subject of much debate. The fossil record of the Isoptera is relatively short, dating back to the lower Cretaceous. Although these first known termites belonged to the extant family Hodotermitidae, it seems likely that other recent families also originated in the late Mesozoic. Mastotermitidae were widespread in Tertiary times and other recent families are present in Oligocene amber (Riek 1970; Kukalová-Peck 1991).

Five families are represented in Australia: Mastotermitidae, Termopsidae, Kalotermitidae, Rhinotermitidae and Termitidae. The first four are commonly (but inappropriately) grouped as the 'lower' termites, whereas the Termitidae are often dubbed the 'higher' termites. These five families are placed in alphabetical order in this database for the convenience of self indexing.

When the first edition of The Insects of Australia was published in 1970, 182 termite species were known from Australia and it was believed that our knowledge of the fauna was almost complete. In the second edition, Watson & Gay (1991) increased the known fauna to 348 species, and recognised that the fauna might be far larger than this. The change in perspective has been due to several factors, some of which Watson & Gay (1983) documented: the survey of new areas; the use of new sampling techniques (in particular, the skimming technique for collecting soil-dwelling termites, and the chain saw and chisel for extracting kalotermitids from dry wood in standing trees); the recognition of biological differences as markers of specific difference (cf. Watson & Perry 1981); the relevance of gut structure in termite taxonomy (cf. Miller 1984a, 1984b, 1991); and the taxonomic importance of the chemistry of cuticular hydrocarbons in termites, involving correlations with biological characteristics and with multivariate statistical analyses of measurements (cf. Watson et al. 1989).

Two hundred and sixty-seven species are curently recognised. Although the literature on Australian termites is quite extensive, available biological information on many species has remained unpublished. Watson et al. (1998) included much unpublished information, almost all gathered from the accession cards for termites held in the ANIC. These accession cards also provided further details of type localities; the sources of these data in the entries for individual species were not acknowledged.

HISTORICAL NOTES

Watson & Gay (1983) summarised the history of taxonomic research on Australian 'Isoptera'. An initial phase, to 1927, was dominated first by European taxonomists who had no first-hand knowledge of Australia. This was followed by the far more substantial contributions of Froggatt (1897–1898); the reports of European expeditions of Michaelsen & Hartmeyer (Silvestri 1909) and Mjöberg (1920); and by those parts of G.F. Hill's immense contributions that preceded his joining the then Council for Scientific and Industrial Research. The second phase was dominated by contributions from what was then CSIRO Entomology, and includes Hill's later publications plus those of his successors, particularly F.J. Gay, J.A.L. Watson and L.R. Miller.

The most important repository of primary types of Australian termites is the Australian National Insect Collection, Canberra (ANIC). Many of Froggatt's and Hill's primary types are to be found there, as are all the primary types of Australian species described by Gay, Watson and Miller. Most of Hill's other primary types are in the American Museum of Natural History, New York, having been sold to A.E. Emerson (cf. Watson & Gay 1983), or in the Museum of Victoria, Melbourne. The other major repository is the Naturhistoriska Riksmuseet in Stockholm, where Mjöberg's types are lodged.

BIOLOGY

The dominant biological features of the Termitoidae are firstly, that all termites are eusocial, and secondly, that all derive their metabolic energy from cellulose, a polysaccharide that is a major component of woody plants.

Social structure, castes and nests

Termites live in family groups or colonies, ranging in size from a few hundred to a few million individuals. Division of labour is the rule, accompanied by morphological and behavioural specialisation into castes. The reproductive castes include the primary king and queen, derived from the pair of winged adults (alates) that established the colony; secondary or replacement reproductives (neotenics); and nymphal stages that develop into alates and also commonly give rise to neotenics. The bulk of the colony, however, consists of sterile individuals, of two castes, soldiers and workers. In most termites the sterile castes develop from apterous larvae; these give rise to workers which, in turn, can develop into soldiers through an intermediate presoldier instar. Not only do workers vary in size according to their instar, but they may also show sexual dimorphism and, as a result, be able to give rise to two (or more) sizes of soldier, generally termed major or minor soldiers.

The nests of termites are diverse in structure. Some species build compact nests, whereas others inhabit diffuse systems of chambers and galleries. Compact nests may be in the form of mounds, or arboreal, or situated in hollowed, living or dead tree trunks or stumps, in mounds of other species of termite, or subterranean. However, most subterranean nests are of diffuse structure, as are most nests in the mounds of other termites, and the nests of kalotermitids and termopsids, which live in the wood on which they feed and do not require ground contact. The ecological descriptors provided in this database classify nests in these terms.

Diet

Termites eat a wide range of wood and other cellulosic materials, and play an important role in recycling them (La Fage & Nutting 1978). Termites are, however, very selective feeders, individual species showing marked preferences. Wood is the predominant part of their diet. Few termites damage sound (undecayed) heartwood, whether in the wild (in living or dead trees) or in buildings. Only 20 or so species cause major economic losses, primarily by damaging timber in service and synthetic materials, and by hollowing-out trees (Watson & Gay 1991; Peters et al. 2016). Many more species eat sapwood, or wood affected by fungus, i.e. weathered wood, in which the decay is relatively minor, or rotted wood, the composition and strength of which has been severely degraded. Many kalotermitids nest in and eat dead scar tissue on living trees, in addition to wood in stumps, fallen branches and logs. Many of these species, known as drywood termites, primarily infest dry wood, and some can invade timber in buildings. Other kalotermitids, and termopsids, are commonly associated with damp timber, and are known as dampwood termites.

Many other termites feed on plant detritus: leaves, twigs, fragments of grasses and forbs, fallen flowers and other plant litter.

Some termites favour dry grasses. Many ensheath the grass stems but others forage in the open, and chew the stems into chaff which they carry back through their foraging holes into the central regions of the nest, where it is stored. Such species are termed harvesters; many also gather and chaff a wide range of plant detritus.

Termite nests and workings contain cellulosic materials that other termites utilise. These include the stores that harvesters maintain but also, more importantly, the 'carton' and allied materials from which nests and workings are built. 'Carton' is composed of partly–digested woody material that is incorporated, as faeces, with a component of soil, the proportions varying from one species of termite to another. It has recently been recognised that some Australian termites ingest soil, and subsist on the humus it contains (Miller 1984b, 1991). Some of these soil feeders also inhabit the nests of other termites, probably utilising the faecal material in the nest structure. Termites also use dung as a source of cellulose (Ferrar & Watson 1970). Dung from large domestic herbivores, particularly cattle and horses, is the kind most commonly attacked, but dung pellets, as produced by sheep and macropods, are also eaten, as are emu droppings.

These categories underpin the descriptors used in this database for the diet of termites. However, no separate entries are given on the diets of kalotermitids and termopsids, which nest in their food.

Distribution

Watson & Abbey (1993) documented the distributions of the Australian termites in their 'Atlas'. For each species contained in the Atlas, this important reference, together with the relevant page number, is listed in the general references. Many species of Australian termite have been found on islands that are not closely contiguous with the mainland. In Watson et al. (1998) the distributions were given as rather broad drainage basins. These were then converted as accurately as possible to IBRA Regions, but have not been updated so need to be viewed as simply indicative.

Associations with particular habitats

Australian termites show few clear associations with the standard categories of habitat used in the database or with other habitat categories in common use. A few species appear to be confined to rain forests; some others to spinifex grasslands; a few species extend into deep-cracking, self-mulching soils, although they are not confined to them. These associations are entered in this database after the universal descriptor "social".

TYPES

Termite taxonomists designate either primary reproductives (particularly alates) or soldiers as primary types. Specimens of other castes are referred to as morphotypes—in effect, they are allotypes, and may or may not be paratypes or paralectotypes. It is also necessary to distinguish between two kinds of paratypic or paralectotypic material. The series from which the primary type was selected has greater taxonomic significance than paratypes or paralectotypes collected from other colonies, which are comparable in importance with paratypes of non-social insects.

In the ANIC, the primary type and morphotypes from the same colony are kept separately from the main collection, and the primary types are numbered as part of the ANIC Type Register. In this database, the number of each primary type in the ANIC is entered as "ANIC No. …". Only paratypes and paralectotypes collected from the same colony as the primary type are documented in this work. In the case of material in the ANIC, the morphotypes are mentioned first, followed by other parts of the series from which the primary type was selected. This series is referred to as the type colony, and the reference is followed by its accession number (if any) in the ANIC Termite Collection (as "series No. 10 …"), then by the castes represented, in the order primary reproductives, neotenics, nymphs, presoldiers, soldiers and workers and eggs.

Prior to 1942, Hill rarely designated holotypes, nor did his predecessors. However, in his Monograph Hill (1942) cited individual primary types of most of the species he recognised, and gave the name of the institution in which they were lodged. He referred to most as holotypes, but some as hololectotypes, usage based on the labels placed in the type vial. All constitute lectotype designations under Art. 74(a) of the 1985 Code.

General References

Calaby, J.H. & Gay, F.J. 1959. Aspects of the distribution and ecology of Australian termites. pp. 211-223 in Keast, A., Crocker, R.L. & Christian, C.S. (eds). Biogeography and Ecology in Australia. The Hague : Dr. W. Junk.

Creffield, J.W. 1991. Wood Destroying Insects. Wood borers and termites. East. Melbourne : Commonwealth Scientific and Industrial Research Organisation vi 44 pp.

Ernst, E. & Araujo, R.L. 1986. A Bibliography of Termite Literature 1966–1978. Chichester : John Wiley & Sons vi 903 pp.

Ferrar, P. & Watson, J.A.L. 1970. Termites (Isoptera) associated with dung in Australia. Journal of the Australian Entomological Society 9: 100-102

French, J.R.J. 1986. Termites and their economic importance in Australia. pp. 103-129 in Vinson, S.B. (ed.). Economic Impact and Control of Social Insects. New York : Praeger.

Froggatt, W.W. 1897. Australian Termitidae. Part II. Proceedings of the Linnean Society of New South Wales 21: 510-552 pls xxxv-xxxvi [date on journal 1896, actual date of publication 31 May, 1897 Musgrave, A. 1932. Bibliography of Australian Entomology 1775–1930 with Biographical Notes on Authors and Collectors. Sydney : Royal Zoological Society of New South Wales viii 380 pp. [103]]

Froggatt, W.W. 1897. White ants, with some account of their habits and depredations. Agricultural Gazette of New South Wales 8: 297-302

Froggatt, W.W. 1898. Australian Termitidae. Part III. Proceedings of the Linnean Society of New South Wales 22: 721-758 pls xxxiv-xxxv [date on journal 1897, actual date of publication 4 June, 1898 Musgrave, A. 1932. Bibliography of Australian Entomology 1775–1930 with Biographical Notes on Authors and Collectors. Sydney : Royal Zoological Society of New South Wales viii 380 pp. [103]]

Gay, F.J. 1968. A contribution to the systematics of the genus Amitermes (Isoptera: Termitidae) in Australia. Australian Journal of Zoology 16: 405-457

Gay, F.J. 1969. Species introduced by man. pp. 459-494 in Krishna, K. & Weesner, F.M. (eds). Biology of Termites. New York : Academic Press Vol. 1.

Gay, F.J. 1970. Isoptera (Termites). pp. 275-293 in CSIRO (ed.). The Insects of Australia. A textbook for students and research workers. Carlton : Melbourne University Press 1029 pp.

Gay, F.J. 1971. The Termitinae (Isoptera) of temperate Australia. Australian Journal of Zoology Supplementary Series 3: 1-36

Gay, F.J. & Calaby, J.H. 1970. Termites of the Australian Region. pp. 393-448 in Krishna, K. & Weesner, F.M. (eds). Biology of Termites. New York : Academic Press Vol. 2.

Gay, F.J. & Watson, J.A.L. 1982. The genus Cryptotermes in Australia (Isoptera: Kalotermitidae). Australian Journal of Zoology Supplementary Series 88: 1-64

Grassé, P.-P. 1986. Termitologia. Anatomie, Physiologie, Biologie, Systématique des termites. Paris : Masson Vol. 3 xi 715 pp.

Hadlington, P. 1987. Australian Termites and Other Common Timber Pests. Kensington : New South Wales University Press vii 126 pp.

Hill, G.F. 1942. Termites (Isoptera) from the Australian Region. Melbourne : Council for Scientific and Industrial Research 479 pp. 24 pls.

Inward, D., Beccaloni , G & Eggleton, P. 2007. Death of an order: phylogenetic study confirms that termites are eusocial cockroaches. Biology Letters 3: 331–335

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.

La Fage, J.P. & Nutting, W.L. 1978. Nutrient dynamics of termites. pp. 165-232 in Brian, M.V. (ed.). Production Ecology of Ants and Termites. Cambridge : Cambridge University Press.