Higher Taxon Termitoidae
Higher Taxon Termitoidae
Termites, White Ants
Compiler and date details
31 December 1996 - J.A.L. Watson, L.R. Miller & H.M. Abbey, CSIRO Entomology, Canberra, Australian Capital Territory, Australia
Isoptera, termites or, coloquially, white ants, are a small order of social insects allied to cockroaches. The most appropriate placement of the order among other groups of blattoid–orthopteroid insects is the subject of much debate and outside the scope of this work. It can suffice that 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).
The number of families recognised is the subject of debate and suborders and superfamilies are not in general use in the higher classification of Isoptera (cf. Grassé 1986). 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 have been placed in alphabetical order in this work 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) have 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).
Of the ca 350 species now recognised, only a little more than 260 have validated available names. This work deals only with these 260 or so species. It is, therefore, a document that will require much revision as further groups of termites are given adequate taxonomic treatment. Although the literature on Australian termites is extensive, available biological information on many species has remained unpublished. This work, therefore, includes much unpublished information, almost all gathered from the accession cards for termites held in the ANIC. These accession cards have also provided further details of type localities. We do not acknowledge the sources of these data in the entries for individual species, for it would be too cumbersome to do so.
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 folowed 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 is dominated by contributions from what is now the 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.
The dominant biological features of the Isoptera 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 work classify nests in these terms.
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). 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 work for the diet of termites. However, we give no separate entries on the diets of kalotermitids and termopsids, which nest in their food.
Watson & Abbey (1993) document the distributions of the Australian termites. 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 the general descriptors of distribution given in this work, such records are included in the appropriate marine zone; the names of the islands or island groups are spelled out after the semicolon following the general descriptors.
Associations with particular habitats
Australian termites show few clear associations with the standard categories of habitat used in the Catalogue 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 work after the universal descriptor "social".
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 work, 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.
As is normal practice in this work, the entries for primary types that we have not examined are marked with an asterisk. We have, however, left unmarked those entries for types that our colleague Frank Gay has examined during his recent revisions of Australian termites.
Snyder (1949, 1956–1968), Ernst & Araujo (1986) and, from 1979, Termite Abstracts have reviewed the world literature on termites.
There is also a substantial literature on Australian termites, covering their taxonomy, biology and management. We concentrate here on the taxonomic and biological aspects. The major work on Australian termites is that of Hill (1942), which gave a wealth of biological information in a taxonomic context. Although out of date, Hill's monograph has provided a framework on which all later taxonomic work has depended. This later work includes revisions of Amitermes (Gay 1968); the genera of the Termes complex (Gay 1971 and Miller 1991); the Australian harvester termites Drepanotermes (Watson & Perry 1981) and the drywood termites Cryptotermes (Gay & Watson 1982). There are, in addition, many smaller papers on a range of genera, particularly on Kalotermitidae; these are mentioned in the relevant introductions to families.
The general and biological literature is more diverse. The first major publication was Ratcliffe et al. (1952), a book rendered obsolete by the many later advances in our knowledge of termite biology in Australia. A replacement for it is much needed. Other contributions include books by Lee & Wood (1971) (termites and soils), Watson (1988) (termites in the Canberra region), Watson et al. (1985) (castes), Hadlington (1987) (biology and management of termites), Creffield (1991) (biology, management), and Watson & Abbey (1993) (distributions); and chapters and papers by Calaby & Gay (1959) (ecology and zoogeography), French (1986) (economic importance), Gay (1969) (introduced species of termites), Gay & Calaby (1970) (termites of the Australian region), Gay (1970) and Watson & Gay (1991) (summaries of knowledge of Australian termites), and Perry et al. (1985) (south-western Australian termites).
The senior author, Dr J.A.L. (Tony) Watson, who established the framework of this work, died before the manuscript was complete. It is fitting that his last publication on termites should be a summary of our knowledge of the fauna with which he was so familiar. We thank Dr Keith Houston for preparing the skeletal database files from a checklist into which we compiled the remaining data, and for editorial advice. The work was assisted by a grant from the Australian Biological Resources Study to J.A.L.W.
The information on the Australian Faunal Directory site for the Isoptera is derived from the Zoological Catalogue of Australia database compiled on the Platypus software program. It incorporates changes made to the work published on 2 September 1998 as (Watson, J.A.L., Miller, L.R. & Abbey, H.M., 1998)
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.
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