Australian Dragonflies: A Guide to the Identification, Distributions and Habitats of Australian Odonata

1 Introduction

Dragonflies are conspicuous insects. Many are large; they fly strongly; most are brightly coloured. As a result, they have been collected extensively. Their larvae are less familiar. ‘Mud-eyes’, as some are called, are drab, and almost all live in fresh waters, out of sight. They are, perhaps, best known as bait for freshwater fish.

The dragonflies constitute a very distinct order of insects, the Odonata. In Australia, two suborders are represented: damselflies (Zygoptera), generally very slender insects, the fore- and hindwings similar in shape and venation and commonly held closed above the body at rest (Figs 46–63), the larvae with external gills on the end of the abdomen (Figs 4A–C, E); and dragonflies proper (Anisoptera), stouter, stronger-flying insects, the fore- and hindwings more or less dissimilar in shape and venation and commonly held spread at rest (Figs 64–101), the larvae with internal, rectal gills (see Chapter 2). Living representatives of the third suborder (Anisozygoptera) are confined to Japan and the Himalayas. The term ‘dragonfly’ is commonly applied to the entire order.

Australia has a diverse fauna of dragonflies. At least 302 species occur here, one of which is not dealt with in this book; it is a species of Synthemis known only from a larva apparently collected in Albury, in southern New South Wales (Theischinger and Watson 1986). The remaining 301 species are at present grouped into 102 genera and 17 families, 11 of Zygoptera and six of Anisoptera.

Both larvae and adults are predators and, as a result, are liable to concentrate any toxic pollutants that their prey (commonly other freshwater insects) accumulate. Thus the fact that dragonflies depend on fresh water and other organisms that live in it means that their wellbeing is inevitably linked to that of aquatic habitats. As Corbet (in Walker and Corbet 1975) has commented, it is important that fresh waters remain able to ‘sustain a diverse and vigorous dragonfly fauna – a reliable, and delightful, indicator of a healthy environment’.

Fresh water is scarce in Australia and pressure for its exploitation can become overwhelming. In order to minimise the effects on the faunas that depend on it, those who frame decisions on the way that water is exploited, and the ways in which surrounding habitats are managed, need information on the organisms involved. What species occur there? Where else do they occur? In the case of dragonflies, are they present as adults, or as larvae? How restricted are they to specific habitats? How might alterations to those habitats affect their capacity to sustain the faunas? Can we argue for restoration of a habitat based on the loss of part of its fauna, or is it then too late? Answers to all these questions depend on detailed knowledge of the fauna, in this case the dragonflies.

The Australian species of dragonfly are quite well known as adults, but few larvae have been described. Unfortunately, most of the data are scattered. Fraser (1960) provided a key to the adults of Australasian dragonflies but his work is now very much out of date. Watson (1962) gave keys to the adults and larvae of Odonata from southwestern Australia, and Allbrook (1979) did the same for Tasmania, but these two regions contain only a very small fraction of the Australian dragonfly fauna (Watson 1974). Many groups of Australian Odonata have been revised during the last fifteen years, principally by Theischinger and Watson. A replacement for Fraser (1960) is much needed, even if only as a tool that could pave the way for more extensive accounts of our dragonflies.

This book is designed to provide that replacement. It is basically an illustrated key to the adults of Australian Odonata, with notes on their distribution and biology, as Fraser (1960) was. It complements the catalogue of Australian Odonata (Houston and Watson 1988), which gives detailed references to taxonomy and biology.

The introductory, general chapters deal with the biology and affinities of Australian dragonflies; the fauna itself, with descriptions of new genera and species; the distributions of species; and explanatory comments on the structure of dragonflies and the use of the keys. The keys follow.

2 The Biology of Dragonflies

The biology of dragonflies is at the core of most questions that we ask about them – their distribution, their evolution and affinities, their place in ecological systems, their conservation, their morphology and anatomy. In this account of it, our emphasis is on the Australian fauna. However, to put this into perspective, we comment first on the biology of Odonata as a whole.

General Biology

Tillyard (1917) provided one of the earliest accounts of dragonfly biology, a book still of great value here because it deals primarily with Australian examples. Tillyard emphasised functional morphology. Corbet (1980, 1983), on the other hand, wrote more of the lives of dragonflies – their life histories, ecology, and behaviour – a perspective which underpins this chapter and is essential to environmental studies of dragonflies.

All Odonata have a common life pattern.

The eggs are laid in fresh water or, less commonly, near it. Females with a complete ovipositor (all damselflies and aeshnid, neopetaliid and petalurid dragonflies; Figs 1A, B) lay them in the submerged tissues of aquatic plants, sometimes superficially into decayed wood or slime covering rocks, or into twigs and stems above or beside water. The females (or pairs) may submerge themselves as the eggs are laid. Females with a reduced ovipositor (other families of dragonflies; Fig. 1C) generally lay them into the water; the eggs are extruded from the gonopore in large numbers and the female repeatedly dips the tip of her abdomen into the water, washing off the eggs. Some females have a long but incomplete ovipositor (Fig. 1D), apparently enabling them to lay eggs into the muddy floor of bogs or shallows. A few have developed the trick of flicking eggs into small, inaccessible bodies of fresh water, such as water pockets in hollow tree trunks, a habit not yet detected in Australia.

A, B, Petalura gigantea, ventral and left lateral views; C, Antipodogomphus hodgkini, ventral view; D, Odontogomphus longipositor, ventral view.

Dorsal views: A, Austrolestes aleison (from Watson 1962); B, Austroaeschna anacantha (T.W.); C, Synthemis leachii (from Watson 1967a). Lateral views (from Watson 1962): D. Austrogomphus collaris; E, Petalura hesperia; F, Synthemis cyanitincta.

The eggs hatch to the first of, commonly, 10–12 larval stages. With very rare exceptions, the larvae are aquatic. All are predators, most feeding on smaller aquatic invertebrates, especially insects, and less commonly on tadpoles and fish. They catch their prey with a highly specialised, elongate lower lip (labium). Normally folded under the head, it can be extended very rapidly and precisely to impale the prey with the pair of movable hooks, one on each palp at its tip, further heavy spines often serving to make escape even more difficult (Figs 2A–C). The movement of the prey seems to be the most important stimulus inducing the strike. When folded, the labium lies flat in all larval damselflies and in aeshnid, gomphid and neopetaliid dragonflies (Fig. 2D). The palps partly cover the lower part of the face in petalurid dragonflies (Fig.2E), and cover the face up to the eyes in corduliid and libellulid larvae (Fig. 2F).

The gills differ greatly between larval damselflies and dragonflies proper. In all larval damselflies known from Australia, the gills take the form of three leaf-, sac- or spine-like appendages at the end of the abdomen (Figs 4A–C, E). In some larvae the gills are clearly made up of two segments (nodate) (Fig. 3A); in others there is a weakly-marked line of division (subnodate) (Fig. 3B); but in many larvae the gill is not subdivided (Fig. 3C). The gills also aid in swimming. In the dragonflies proper, the gills consist of rows of plates that project into the enlarged rectum, which is richly supplied with breathing tubes (tracheae) (Fig. 3D), and there are no external gills (Figs 40, F–I). ‘Breathing’ is achieved by pumping water into and out of the rectum. Vigorous ejection of water also produces jet propulsion, the active form of swimming.

Median gill of mature larva: A, Eurysticta coolawanyah; B, Pseudagrion microcephalum; C, Nososticta pilbara. (E. Rankin.) D, gill basket of Hemicordulia tau (T.W.)

Larval lives are relatively long, lasting several months to several years. Late in the last larval stage, signs of metamorphosis become apparent: the swelling of the wing-cases to accommodate the adult wings that are developing in them; the enlargement of the pigmented area representing the developing eyes of the adult; and the retraction of tissue from the long larval labium to build the short labium of the adult. The larval gills become ineffective and the larva crawls to the surface to breathe air through the spiracles immediately behind the head (the mesothoracic spiracles). Finally it swims or crawls to a suitable support – an emergent stem or rock, or a tree trunk away from water, or, particularly in the case of gomphid dragonflies, the sloping bank – where the adult splits the larval skin, emerges from it, then expands its abdomen and wings. All adult Odonata emerge from supports of this kind.

A, Austrosticta soror; B, Austroargiolestes icteromelas; C, Synlestes weyersii tillyardi; D, Petalura hesperia E, Diphlebia euphaeoides; F, Notoaeschna sagittata; G, Austrogomphus guerini; H, Diplacodes bipunctata; I, Synthemis macrostigma occidentalis. (Rearranged and magnifications changed from Watson and O'Farrell 1991.)

Emergence marks the beginning of the adult stage, which can be regarded as comprising two phases: the pre-reproductive and the reproductive periods. Together, they usually amount to a total of a few weeks to 2–3 months (excluding species that, for example, overwinter as inactive adults).

Two types of flight behaviour should be noted here. Some dragonflies are ‘perchers’; the adults spend much of their time perched on the ground or on prominent stems or twigs, from which they fly to intercept prey or, in the case of mature males, to combat competing males of their own species or to seduce females. Others are ‘fliers’; they spend all their active time on the wing. Damselflies are perchers, as are our gornphid, petalurid and most libellulid dragonflies. The fliers include dragonflies of the families Aeshnidae, Neopetaliidae and most Corduliidae, with a few Libellulidae. Intermediates are known, principally among the libellulids.

The pre-reproductive period is the period of sexual maturation and usually lasts from several days to a few weeks (three or less). During this time, the newly-emerged (teneral) adults fly from their emergence sites, their wings slightly opalescent (Fig. 82), then start to feed. The colour of the body usually changes, often very markedly, involving wing colour, the brightness of pigmentation (for example, pale yellowish brown to bright red or blue), and the development of white to grey-blue pruinescence (Figs 47, 58, 96), often more marked in males than in females (Figs 87, 89). Indeed, the sexes often differ strikingly in coloration (Figs 49, 50; 93, 94). These changes in colour may continue into the reproductive period.

Like the larvae, the adults are predators. Most feed on other, smaller insects which they capture in flight, and consume in flight or after they have perched. Predation does not appear to be selective, and the prey may include smaller species of Odonata (Figs 74, 97). A few Odonata gather sedentary insects or other organisms from plant stems (gleaning) (Fig. 48).

The pre-reproductive period is commonly the period during which adults disperse. Some (in particular, those confined to streams) disperse little from their emergence sites but others (most spectacularly, those species whose larvae can utilise temporary waters) can fly far, often in flocks. The prime example is Pantala flavescens, a libellulid which occurs around the world in warm temperate and tropical latitudes and commonly breeds in shallow, still waters. Its adults have been collected far from fresh water, including captures on isolated oceanic islets that lack surface waters, and on shipboard, far from land.

The reproductive period begins when the adults, having matured sexually, start to show reproductive behaviour (although mating has been observed in flocks of vagrant species). In the males this is commonly manifested by a return to fresh waters of the kind from which they emerged. Females usually return later, once they have developed a batch of eggs. Immediately they return, the mature males set upon them. Once they have mated and laid their eggs, they leave the water and remain away until another batch of eggs has matured.

Reproductively active males commonly establish territories (areas of water surface, on flowing or still waters) which they defend against males of their own species. In the case of perchers, the male protects his territory from a base, periodically going on patrol flights, whereas fliers patrol their territory as long as they are on it.

Once a male intercepts a female at the breeding site, a complex but apparently universal pattern of behaviour follows. After a preliminary skirmish, which establishes whether or not the female is receptive, the male grasps her head or prothorax with his anal appendages, forming the tandem position (Fig. 5A), in which the pair flies. The female then bends her abdomen so as to apply its end to the secondary genitalia on the second abdominal segment of the male (the wheel position) (Figs 5B, 83, 98); the male had earlier transferred sperm from the apex of his abdomen to the secondary genitalia. The pair usually settles. Following copulation, the female returns to the breeding site – alone or with the male, sometimes still in tandem – and lays her eggs. If the male accompanies her, he may protect her against the attentions of rival males.

Coenagrion lyelli: A, tandem position; B, wheel position. (Rearranged from Watson and O'Farrell 1991.)

Biology of Australian Dragonflies

Watson (1981, 1982a) reviewed our biological knowledge of Australian dragonflies, and briefer summaries appeared in O'Farrell (1970) and Watson and O'Farrell (1991). Watson (1962, 1963, 1969a), Allbrook (1979) and Hawking (1986) included observations on the biology of Western Australian, Tasmanian and Victorian dragonflies, many of which occur in other states, and Rowe (1987) commented extensively on New Zealand odonates, some of which also occur in Australia. Houston and Watson (1988) catalogued the described Australian fauna, including notes on the distribution and ecology of each species. Watson (1974, 1977a) summarised the data on distributions.

Some useful perspectives have emerged from these reviews, involving biological peculiarities that Australian dragonflies show. Of major relevance are the relationships between larval habitats and the distributions and behaviour of the adults; the problems posed by drought; the large number of species confined to permanent streams and rivers; and the apparent origin of many Australian Odonata from the dragonflies that inhabited Gondwana, the land mass that began to fragment some 135 million years ago and gave rise to the southern continents and the Indian subcontinent.

Three generalisations need to be borne in mind. First, today's dragonflies are primarily tropical and warm temperate insects so that, in general, the rich faunas will be found near the equator. Second, many of our stream-breeding dragonflies are of southern origin, so that the groups to which they belong are very well represented in the south. However, the ranges of these southern groups extend north along the Great Divide, many reaching northern Queensland, where they mingle with more purely northern dragonflies (see Figs 6A, B). Third, female dragonflies commonly lay more than one batch of eggs, totalling many hundreds, but only two of these eggs need give rise to successfully breeding adults for population levels to remain stable. This is particularly relevant in the case of dragonflies breeding opportunistically, as in fresh waters subject to drying or salinity.

Larval Habitats

The larvae of many Australian dragonflies live only in permanent streams and rivers, or in the permanent pools that remain in them during the dry season. Others inhabit still waters – ponds and lakes – many also occurring in flowing water. Larvae of a few species are found in temporary waters, usually still: we deal with their biology in the next section.

The behaviour and distribution of adults are correlated, at least in broad terms, with the larval habitats (cf. Watson 1981; Watson et al. 1982). Obligatorily stream-dwelling larvae generally give rise to adults that remain close to the streams from which they emerged. At the other extreme, species that breed in still, particularly shallow, waters commonly have adults that fly far from water, apparently during the pre-reproductive phase. This is a general characteristic of dragonflies that can utilise temporary ponds. Species with predominantly riverine or lake-dwelling larvae apparently show variable, intermediate dispersal. This pattern of behaviour makes sense in terms of the availability of larval habitats in Australia.

Thus there are far more species of Odonata in the well-watered areas of coastal northern, eastern, south-eastern and south-western Australia, where flowing as well as still waters are abundant, than there are in the inland. Most inland species are widely distributed, some ranging throughout mainland Australia and a few extending to Tasmania, or New Zealand, or beyond (see Chapter 5). They are, in addition, taxonomically diverse: at least two families of damselflies (Coenagrionidae and Lestidae) and three of dragonflies proper (Aeshnidae, Corduliidae and Libellulidae) are represented in the inland fauna.

Dragonfly Larvae and Drought

Australian dragonflies whose larvae can be found in habitats that periodically lack fresh water include not only those utilising temporary ponds, but also species breeding in other environments. It is useful to comment on five such habitats: temporary, shallow pools; saline habitats; streams and swamps liable to seasonal drying; water-filled holes in tree trunks (phytotelmata): and rainforest floors.

A, a southern group, the corduliid subfamily Synthemistinae; B, a northern group, the libellulid genus Orthetrum, excluding the widespread species O. caledonicum. (Based on specimens in the Australian National Insect Collection.)

(l) The mechanisms that enable our dragonflies to utilise temporary ponds are of at least three kinds.

The first is rapid larval growth. The shortest larval lives occur in species that develop in shallow, warm water, and the rate of larval development rises steeply with increasing temperature. Some odonates that use