Tettigoniidae of Australia Volume 2: Austrosaginae, Zaprochilinae and Phasmodinae

Introduction

This second volume in the series covers the two endemic subfamilies, the Zaprochilinae and Phasmodinae, both of which are bizarre in appearance and equally unique in food preferences, and the similarly endemic Austrosaginae, subfam. nov., which consists of Australian representatives of a Palaearctic and Ethiopian group of primitive katydids. The first two of these three subfamilies are closely related to one another, whereas, the third is not related at all. But their inclusion in this second volume of the series is not at all surprising. All three of these subfamilies and the Tettigoniinae, the topic of Volume 1 of this series, can be collected together in the springtime. The phasmodines and zaprochilines are adult at that time whereas the others are found mostly in the nymphal stages in spring. If rearing and culturing of live material is the object, then the four subfamilies are important because they are simultaneously encountered in the field in the same habitats.

Katydids of the Zaprochilinae and Phasmodinae are strikingly phasmatoid in appearance. In the former, the type genus, Zaprochilus Caudell, has tessellated hind wings as do many stick insects but more surprisingly they are coloured pinkish or purple at the base, just as they are in most phasmatids. The tegmina are raised at an angle from the body at rest (Fig. 24) and rolled phasmatid-like. This is a form of crypsis exhibited during the day when the insect is inactive conveying the appearance of a twig. While the head is strongly prognathous in all known zaprochiline species, it is somewhat less so in the phasmodines. But in the latter, the phasmatoid appearance is even more pronounced (Fig. 16) in the elongation of the body and spindly appearance of the legs. Interestingly, the elongation of the thorax is the result of the exaggeration of different thoracic segments in the respective groups. In the Phasmodinae, the prothorax is disproportionately extended relative to the other segments, while in the phasmatids it is the mesothorax that is elongate. The hind femur of phasmodine katydids is slender and not expanded basally as is normal in Saltatoria, but the katydids have not lost their ability and inclination to jump when threatened. They do so with considerable agility and effect even though seemingly lacking the muscles of an expanded femoral pagina which would allow them to do so.

Recent understanding of the biology of species of both subfamilies has resulted in an abundance of specimens for study where just a few years ago they were very rare in collections. Most species are common and many can be abundant during most years when favourable ecological conditions exist. Their presence in spring is associated with the abundance of flowers, especially monocotyledons.

Both the Zaprochilinae and the Phasmodinae utilise flowers as a food source. They do so in very different ways. The phasmodines feed on the entire flower; they consume the petals, anthers, sepals and stigma. A most ironical situation exists in Kings Park, Perth from time to time when Phasmodes ranatriformis Westwood, hitherto, one of the world’s rarest katydids in collections, could be considered a ‘pest’ because it destroys blossoms of the equally unusual Kangaroo Paws, Anigozanthos, spp. The zaprochilines, with one possible exception, do not appear to cause any damage to the flowers on which they feed. They carefully manipulate the anthers to release pollen which they consume and sip large amounts of nectar. With the possible exception of Windbalea viride, gen. et sp. nov., I found no trace of injury to any of the wide variety of plants that they visit. On the other hand, it seems quite likely that they may be involved in pollination. Their hairy bodies, often pollen-covered, indicate a potential for the transmission of pollen. These katydids are not host specific but are opportunists. Kawanaphila, gen. nov. species seem to be most abundant in Lepidosperma spp. habitats and have been observed feeding on the reproductive parts of these plants, but they also have been seen on adjacent plants that are entirely unrelated to Lepidosperma. With the exception of at last two species in that genus, they occur on or very close to the ground. In contrast, Zaprochilus, Windbalea and Anthophiloptera Rentz and Clyne species may be found in low bushes or high in trees.

Zaprochiline species occur widely in Australia (Maps 2–6) in all quarters except the northwest. The greatest abundance of species is in the southwest corner of the continent in the sand plain heath community. Kawanaphila, gen. nov. and Zaprochilus have very broad ranges with species being found on both sides of the southern portion of the continent. Anthophiloptera, while confined to the eastern seaboard, has an unusually extensive range for its single species (Map 3). This is somewhat parallel to the distributional pattern of Windbalea viride, a similarly monotypic genus but occupying a broad range of the southern coast west of Esperance, Western Australia. Both the Phasmodinae and Zaprochiline have specialised mouthparts and alimentary tracts adapted for the floral-feeding specialisation.

The Austrosaginae are rather typical-looking katydids. All known species are micropterous, brachypterous or mesopterous and most occur in late spring and throughout the summer. They blend a mixture of predation with frugivory. Some species are very spiny (Fig. 197), a characteristic of predaceous katydids, others are less so, and many are heavy-bodied. The latter, although opportunistically predaceous, are adapted for seed and fruit-eating, the incredibly large heads (see Fig. 199) replete with muscle to facilitate breaking through tough seed-coats. More information about this subfamily is given later in the text.

Format

I have followed a similar format to that in the first volume of this series, Rentz (1985). The reader should consult that work for terminology, anatomical parameters for measurements and conventions for the citation of label data. Descriptions, at least in Kawanaphila, are brief due to the uniformity of species in all but a few critical characters, especially those of the genitalia.

For the first time I have utilised the scanning electron microscope to study and illustrate the sclerotised portion of the concealed male genitalia, appropriately termed the titillators. There is an inherent problem with this technique in the inability of the machine to differentiate the membranous from sclerotised portions of the anatomy. The technique is proferred for the genera Hemisaga and its relative Austrosaga, gen. nov. (Figs 95–97). I have adopted the technique common with chromosomal preparations in cutting the taxonomically important portions from the photographs and affixing them to a plate. In addition to the time-saving aspects of using SEM’s, microanalysis reveals a texture to these structures that has not been revealed in previous illustrations. The fact that the titillators are generally asymmetrical is also not widely appreciated.

Collecting and Rearing

As was discovered with the tettigoniine katydids, specialised collecting is more productive than more generalised techniques. Previously, I found that collecting in early spring yielded an abundance of tettigoniine and austrosagine nymphs that could be reared to maturity in the laboratory. Fortuitously, it was discovered that adult phasmodines and zaprochilines were present at the same time so that adults of one group and nymphs of another could be collected simultaneously. Had I not gone into the field in late winter and early spring, the phasmodines and zaprochilines might easily have been overlooked.

Katydids of both the Phasmodinae and Zaprochilinae can be collected with equal success night and day. Because of the stick or twig-like appearance of these groups they can be detected during the day only by disturbing them. At night they are easily seen in the light of a torch while they are feeding. Sweeping or beating is generally very productive if the appropriate host plants are checked. For Zaprochilus species one should examine flowering grass trees Xanthorrhoea spp., isolated flowering acacias and eucalypts and Kunzea spp. With Kawanaphila and Phasmodes species, monocotyledonous plants seem to be preferred by most species. In the southwest of the continent, heath areas with grasses and reed-like plants such as Lepidosperma spp. can usually harbor large numbers of individuals. As with Zaprochilus species, flowering grass trees can be an important food source for Kawanaphila species.

Phasmodine katydids are wingless and produce no high frequency sounds. Zaprochilines stridulate with unusually high frequencies, most of which cannot be heard by the human ear. However, they can be heard using the ‘bat detector’. Even on relatively cold nights (10°C), in rainy conditions, Kawanaphila species are active and singing.

Rearing zaprochilines and phasmodines is not difficult if the proper foods are presented. The former can be induced to feed on pollen granules obtained from healthfood stores. I add a small amount of honey to their water container and have been able to maintain nymphs and adults in the laboratory for many weeks. One problem that is not easy to overcome, is the concomitant development of large populations of cereal mites (Anoetidae and Acaridae, det. R. B. Halliday) that can buildup when large numbers of specimens are kept. These mites seem to be present in small numbers in most environments and feed and thrive on cereal products. The conditions of humidity and food availability in the glass jars used to rear katydids seem to be ideal for the mites. There is no way to rid the cultures of these creatures other than by regularly changing of the containers and food. If left to develop in cultures, the mites will eventually kill the katydids, attaching themselves to the recently-moulted nymphs and feeding on their body fluids.

Collecting adult austrosagines can be frustrating at first but once their habits are determined, they can be obtained with little difficulty. They are similar in a number of respects to the tettigoniines. In appearance, they can easily be mistaken for tettigoniines, especially when juvenile. They are frequently collected with juveniles of the Tettigoniinae in the same habitats. They can be reared to maturity in the laboratory but frequently have difficulty in moulting with legs and other appendages often becoming distorted after moulting. Whether this is the result of inadequate diet, artificial lighting or improper relative humidity, I have been unable to determine. Adults live for long periods in the laboratory provided with the diet suggested in Volume 1. Adults are active both night and day. During the day, in the field, males assume perches in tall bushes or small trees where they broadcast their calls. At this time they are extremely wary and have very keen eyesight, probably to detect potential predators, and they are difficult to approach. Considerable time must be spent doing so. Females are seldom encountered at this time. After dark stridulation continues, often well past midnight, but the same individuals are less wary and less reactive to disturbance of their habitat. They are much easier to collect but females are only seldom encountered and are best obtained by rearing nymphs both sexes of which are abundant in the short grasses and low shrubs in spring. All species with which I am familiar stridulate with frequencies easily audible to the human ear.

Preservation of Specimens

Because of the soft-bodied nature of most zaprochiline species, they are best preserved first in Pampel’s fixative then in 70% ethyl alcohol (Volume 1, p. 8). This preserves the soft genitalia without distortion which are critical in species determination. Even fully-winged Zaprochilus species should be preserved this way although the majority of specimens seen in this study have been pinned and they are quite useful and can be relaxed and the tip of the male’s abdomen removed and softened for study. Kawanaphila species that are dry-mounted shrivel and distort and are virtually useless for study. Anthophiloptera, Windbalea and Phasmodes species are more sclerotised and may be pinned, after gutting and dusting with preserving powder, without destruction to their soft body parts.

Austrosagine katydids offer problems in colour preservation. All are soft-bodied and can be preserved in fixative and then kept in alcohol. But their colours are often very useful in species determination and the greens and yellows fade in fluid. Because of the large size of most individuals, it is often thought desirable to preserve them on pins. But green austrosagines have a component in the exoskeleton chemistry that inhibits complete preservation of the green colours with techniques that are successful with other katydids. This is true with the typical sagines and with a number of genera of the Listroscelidinae. Only partial success can be obtained by the gutting and stuffing technique. Documentation of the true colours and intricate colour patterns is best accomplished through colour photography.

As will be shown below, the eggs of tettigoniids offer useful characters for the taxonomist. They require special preservation if they are to be studied properly. One important consideration is that naturally laid eggs be obtained. Eggs removed from the abdomens of preserved females often lack the critical sculpturing that is species distinctive. Females can be induced to lay in the laboratory, often with difficulty, in sand or moist cotton. Once eggs have been obtained, the most useful taxonomic study material is that examined with the scanning electron microscope. Eggs that are to be examined with the SEM must first be pricked with a micropin to allow penetration of the following solutions. If the eggs are not first punctured with a minute pin, they will shrivel and distort. The egg is then placed in deionised water with mild detergent and then sonicated for a few seconds to remove fine bits of dirt. Then it is placed in a solution of ‘Super skipper’ (a penetrant) overnight, then into ‘Carl’s solution’ (a fixative) for a 6 hours. Once this is accomplished, the egg is dehydrated through a series of 30, 50, 70, 80, 85, 90, 95, 100% ethanol for about 5 minutes each. Then it is placed in a mixture of amyl acetate and 100% ethanol and finally into pure amyl acetate. The egg is then critical-point dried, gold coated and is ready for examination. This technique has much wider uses, see Grodowitz et al., 1982, who provided details on the composition of the requisite solutions. Although this technique is laborious and time-consuming, it is the only method I know that will provide the results needed to examine the external surface of eggs properly.

Song

One of the results of impending age is the inability to hear high frequency sounds and therefore, a simple, inexpensive electronic device called a ‘Mini Bat Detector’ has helped to solve the problem by enabling one to hear in the field sounds in the 10–160 kHz range and to virtually pin-point their source. The ultra-sound is picked up by a capacitance microphone which has a frequency response extending to 160 kHz, and the signals are fed to a very low-noise amplifier mounted in a small transistor radio body. Many species not otherwise encountered by collectors can be discovered using this device.

Analysis of the calling song of the katydids included in this volume was accomplished much along the lines described in Volume 1, p. 14. However, as technology advances, and it is at an ever increasing rate, the technique has varied slightly. Recordings are made as noted there, but as discussed in Rentz (1987) the songs were digitized at 40 kHz and 8-bit resolution by a microcomputer using a revolutionary memory bank of 256 K bytes. The digitized signal is then output on a digital plotter to produce the amplitude with the time plot in the figures. Once the song is computerized, any portion of it can be expanded accurately and precisely. This is what has been done in each of the oscillograms presented in this volume. The rectangle to the above right of each oscillogram indicates which portion of the trace above is presented below. The vertical bars within this rectangle delimit the portion of the song that is presented.

Material Examined

Although this study is based largely on specimens in the Australian National Insect Collection, Canberra, I have examined all of the relevant material in all of the major collections of Australia and overseas. All types have been examined.

Museums and collections from which specimens are cited in the Specimens Examined sections are abbreviated according to the following code. Material of some species collected by the author will be deposited in other collections in Australia as well as overseas.

Morphological Characters Studied

Austrosagines are morphologically similar to tettigoniines and most of the comments regarding external morphology made for those katydids, Volume 1, p. 11 are referable to the Austrosaginae. The following comments apply mostly to the Phasmodinae and Zaprochilinae.

As was true with the Tettigoniinae, the genitalic structure of both sexes, both the membranous and sclerotised portions, provide the most valuable characters for species recognition. The stridulatory file is less useful in the Zaprochilinae because of the relatively small number of teeth and the overlap in number and length among species. Analysis of recorded song is a very distinctive character but, unfortunately, songs of only a few species are available at this time. Colour and pattern that are so important in the tettigoniines and austrosagines are less important among zaprochilines where there is great variation. The specialised nature of the feeding habits of both zaprochilines and phasmodines is reflected in the specialisation of the mouthparts, especially the mandibles and the alimentary canal. Similarly, the mandibles of the austrosagines are diagnostic. These are most distinctive at the generic level. The most important structures are noted below.

Head

The structure of the head of Phasmodes is strongly phasmatoid, that is, horizontal or prognathous with the anterior margin rounded (Fig. 6A). In Zaprochilus, Anthophiloptera, Windbalea, and Kawanaphila it is prognathous but more elongate and cordate (Fig. 41).

(a) Mandibles

I could find no distinctive characters on the labium or maxilla of any of the species but the mandibles are quite distinctive. The difference in feeding habitats between Phasmodes and the zaprochiline species is reflected in the structure of the mandibles. The mandible of Phasmodes spp. is robust and massive and bears 3 large, stout incisors anteriorly (Fig. 1A). There is a single large incisor and an ill-defined grinding area. Posteriorly, on the internal ventral margin there are 3 setae; externally as well as three large teeth, there is a row of setae above the grinding area and 2 or 3 irregular rows posteriorly at the base.

Fig. 1. (Opposite). Mandible of Phasmodes ranatriformis, female, Kings Park, Perth, W.A. A, entire mandible, 66x; B, molar region, 400x; C, portion of antenna illustrating its unspecialised nature documenting only a few olfactory organs, 300x; see text.

Fig. 2. Mandibles of some zaprochiline species. Zaprochilus australis, Cooloola Nat. Park, Qld.: A, entire mandible, 60x; B, molar grinding area, note particles of pollen adhering to the teeth, 260x; C, scale pattern at base of mandible, seen to the top right of A, 2400x; D, Kawanaphila nartee, sp. nov.: Tutanning Nat. Res., W. A., tip of mandible with ‘nectar groove’ and tooth, 260x; E, entire mandible, 94x; molar grinding area, note sharpness of teeth compared with that of Z. australis, 480x.

The zaprochiline genera have a more slender mandible (Figs. 2–4) with the principal incisor extended forward. This incisor is very sharp in Zaprochilus and Kawanaphila and more blunt in the other genera. In each of the genera, on the internal surface, there is a groove (Fig. 2A, 4A, 4B) on the inside of the first incisor which ends in an arrangement of setae. I presume the groove serves as a channel for conducting nectar from the flower and the setae serve as a sieve. The arrangement and number of setae seems generically distinctive and is probably correlated with foodplant preferences. In Zaprochilus there are 3 or 4 rows of irregularly arranged stout setae. In Anthophiloptera there are more rows of more irregularly arranged but shorter setae and in Kawanaphila there is a single row of regular or irregularly arranged stout setae (Fig. 2A). The number of principal teeth on the distal end of the mandible is also generically distinct. In Zaprochilus there are 3 teeth (Fig. 2A), in Kawanaphila there is a single blunt tooth and in Anthophiloptera and Windbalea there are no teeth (Fig. 4A,B) at all. The molar area of Zaprochilus consists of many peg-like projections, in Windbalea there is a grinding area of minute teeth and in Anthophiloptera (Fig. 4B) it is nearly smooth; Kawanaphila (Fig. 3B), on the other hand, bears many slender, sharp projections. It presumed that this structure is used to grind or pierce hard pollen grains (see Fig. 3B which has pollen still attached). At the base of the mandible on the internal margin in Kawanaphila and Zaprochilus there is an area of imbricate scales (Fig. 2C) each with 5–7 acute, triangulate projections of undetermined function. These are absent in Anthophiloptera and Windbalea.

Fig. 3. Mandible of typical Kawanaphila, gen. nov. species. A, K. goolwa, sp. nov., note elongate incisor with groove supposedly for intake of nectar, 120x. B, K. iyouta, sp. nov., molar grinding area, apparently with pollen still attached, 480x.

Fig. 4. Mandibles of two zaprochilines. A, a, Anthophiloptera dryas, A, entire mandible, 36x; a, enlargement of grinding region on lower central margin of A, 94x; B, b, Windbalea viride, gen. et sp. nov., B, entire mandible, 40x; b, enlargement of grinding region on lower central margin of B, 120x.

(b) Antennae

I have observed a peculiarly distinctive feeding behaviour in 2 species of Phasmodes. Prospective food flowers are examined with the antennae. The flower is carefully probed by the apical one-fifth of one or both antennae for several seconds. The flower is antennated nervously in the same manner as is usual for some wasps when probing. If the flower is acceptable, the katydid begins feeding, if not, it moves to another flower without making contact with any of the mouthparts. Scanning electron micrographs were made at several points along the length of the antenna for distinctive sensory receptors. The antenna is clothed in dense, stout setae over its entire length. The setae are more densely packed in the distal portion of the flagellum. Pit chemoreceptors occur, apparently randomly, along the full length of the flagellum. I could find no additional morphological evidence to suggest a modification for the behaviour I observed. No similar behaviour was observed in any of the zaprochiline genera.

Thorax

(a) Foregut and proventriculus

With such specialised food preferences it is not unexpected to find some modification of the foregut and proventriculus in the Phasmodinae and Zaprochilinae.

In Phasmodes the foregut is an extremely long, tubular structure (Fig. 5A, a). The gastric caeca consist of consist of 2 short, apically truncate lobes each of which is shallowly divided. The proventricular area consists of 6 rows of smaller sclerotised cushions separated by a toothless band, the neck, followed by a zone of 3 to 5 structures as normal for the Tettigoniidae. Each row consists of 7 to 9 weakly sclerotised, pillow-like, tuberculate pads.

In Zaprochilus species the foregut (Fig. 5B, b) is also elongate and tubular. The gastric caeca consist of 2 low, rounded pads. The proventricular area bears 6 rows of from 3 to 5 digitiform cushions, each bearing elongate setae. The neck region is distinct and posterior to this there are 6 rows of weakly sclerotised, spiniform appendages.

The foregut in Kawanaphila species (Fig. 5C, c) is similarly elongate and extremely fragile. The gastric caeca are 3-lobed, each lobe a different length, the internal structure of the proventricular region is unique in lacking rows, ridges or teeth. The alimentary canals of Anthophiloptera and Windbalea have not been examined. There may be a correlation between the development of the molar grinding region of the mandible and the development of the proventriculus. It appears that Kawanaphila species have the most intricate molar grinding apparatus but with a poorly developed proventricular area. Zaprochilus species bear a moderately developed molar grinding apparatus and a moderately developed proventriculus. If this is a sequence, then one might predict that with Windbalea and Anthophiloptera lacking a molar grinding apparatus, they might be expected to have a better developed proventricular area.

(b) Pronotum and sternum

Only Anthophiloptera and Windbalea have pronota extending cephalad or caudad to any degree. The caudal margin of the pronotum of both Anthophiloptera and Windbalea extends slightly, roof-like over the base of the tegmina. The remainder of the pronotum is decidedly sellate and produced forward forming a long neck probably to fascilitate probing deep into flowers. The pronotum of Phasmodes (Fig. 6C) is convex dorsally. There is no median carina; the species possess irregularly crenulate lateral carinae. The cephalic margin of the pronotal disk ranges from evenly rounded in the median portion in Kawanaphila and Windbalea to acute in Anthophiloptera and Zaprochilus and concave or emarginate in Phasmodes. The caudal margin of the disk ranges from truncate in Kawanaphila to obtuse in Zaprochilus, Anthophiloptera, and Windbalea to emarginate in Phasmodes. The lateral lobes of the pronotum are not well developed in any known phasmodine or zaprochiline species.

Fig. 5. Foregut and proventriculus of some phasmodine and zaprochiline species. A, a, Phasmodes ranatriformis Westw., A, foregut and proventriculus, a, dissected proventriculus; B, b, Zaprochilus australis Bruilé, B, foregut and proventriculus, b dissected proventriculus; C, c, Kawanaphila nartee gen. et sp, nov., C, foregut and proventriculus, c, dissected proventriculus.

Fig. 6. Diagnostic characters of Phasmodes ranatriformis. A, lateral view head, female; B, ventral view thorax, female, C, dorsal view pronotum, female; D, lateral view hind tarsus, femaie, E, dorsal view apex of abdomen, female; F, ventral view subgenital plate, male; G, ventral view subgenital plate, female.

The thoracic sternites are simple and unmodified in all species of Phasmodinae and Zaprochilinae. Phasmodes (Fig. 6B) is typical for both subfamilies. The thoracic auditory opening is small, open and ovoid in all zaprochiline species and absent in the Phasmodinae.

(c) Tegmina and wings

Tegmina and wings are absent from the Phasmodinae. Anthophiloptera and Windbalea are macropterous, Zaprochilus is macropterous in all but one species which is known from a single micropterous female. Kawanaphila is brachypterous in the male but apterous in the female. The outline and shape of the tegmen (Figs. 63D) seems characteristic for each genus. The indentation of the anterior margin of the tegmen of Kawanaphila species is characteristic and unique. Males of this genus have small wings, the apices of which protrude slightly beyond the tegmina. They are apically positioned in such a way that they appear ‘rolled’ or tubular. The stridulatory files of males (Figs. 54, 55; Table 5) are notable because of the paucity of teeth. They share this in common with the unrelated Microtettigoniinae. The length of the file, number of teeth and general curvature are often species distinctive but there is considerable overlap in this character.

(d) Legs

The legs of all known species in the Phasmodinae and Zaprochilinae are largely unspecialised. All are remarkable for their slenderness. Even the hind femur is cylindrical and relatively little laterally compressed for saltatory orthopterans. However, all species, even the elongate Phasmodes species are saltatorial in their movements, especially when disturbed. There is no coxal spine in any species and the femora and genicular lobes of all species are unarmed.

The fore femur of Phasmodes is strongly compressed in the proximal half just as it is in many phasmatoids. This accommodates the head when the forelegs are extended forward. Zaprochilus and Kawanaphila only show feeble compressing of the fore femur and Anthophiloptera and Windbalea bear no modification in this structure at all. The legs of all zaprochilines are densely setaceous. In Phasmodes the fore and middle legs are mostly bare with a few stout setae positioned along the dorsal margin. The hind legs bear a more dense arrangement of setae most of which are very long.

The tibial auditory tympanum is present but faint in females of Phasmodes species, absent in males. Both sexes of all species of Zaprochilinae bear an ovoid, open auditory tympanum on both sides of the leg. The dorsal surface of the fore and middle tibiae in unarmed in all species of both subfamilies; the ventral surface of the fore tibia bears several spines in the median portion in Anthophiloptera and Windbalea as well as a pair of apical spines. In Kawanaphila only 2 apical spines are present. Zaprochilus varies from 0–2 spines near the apex of the fore and middle tibia. The ventral margin of the middle tibia bears spines only in Kawanaphila and a series of spines and apical spines in Anthophiloptera. In Zaprochilus there is a pair of opposing spines, one on each side in the apical one-quarter, as well as 0–2 spines on either side near the apex; in Phasmodes the ventral margin of the middle tibia is unarmed. The ventral surface of the hind tibia is armed only with a pair of apical spurs in Phasmodes and Kawanaphila, although those of the former are extremely minute; Anthophiloptera and Windbalea have a number of large spines on both the internal and external margins in the apical one-third. Zaprochilus may have 1–2 spines on either side near the apex. The tarsi of all species are simple, unmodified, depressed. There is a small pair of pulvilli at the base of the first segment. The second last segment is strongly laterally flanged in all zaprochiline species and much more slender in Phasmodes species.

Abdomen

The abdomen in all species of both the Phasmodinae and the Zaprochilinae is basically unmodified except at the distal end. The surface is smooth and hairless and bears no carinae; laterally the tergites of Zaprochilus species are angulate. There is no sclerotisation, other than shagreening, in the concealed male genitalia.

Males of Zaprochilus species have the ninth and tenth tergites produced hoodlike over the tip of the abdomen. In Phasmodes, Anthophiloptera, and Windbalea only the tenth tergite is modified. In Kawanaphila the tenth tergite bends ventrally, its lateral angles forming a hook-like appendage which is frequently emphasised by colour. Paraprocts are developed hook-like in Zaprochilus, Anthophiloptera, Windbalea and in one species of Kawanaphila. The supra anal plate is elongate and longitudinally sulcate or greatly reduced and hidden. The cerci of Phasmodes species are well sclerotised and serve a clasping function while those of all zaprochiline species are styliform and unmodified.

Females of both subfamilies bear highly diagnostic characters in the form of carinae or folds on the eighth, ninth and tenth tergites. There is little distinctive about the genitalia on the dorsal surface of the abdomen but the shape of the subgenital plate is highly distinctive, especially in Zaprochilus. The ovipositor is styliform in Phasmodes, lanceolate in Kawanaphila and tending towards falcate in Zaprochilus, Anthophiloptera and Windbalea.

Colour

Colour is a distinctive character in the Zaprochilinae and Phasmodinae but mostly at the generic level. Phasmodes species are generally greenish with a small percentage of yellow or brown forms present from time to time. Zaprochilus species are usually grey with the base of the wing purple or pink. Kawanaphila species are generally grey but in some species brown, tan or green forms and combinations thereof, occur within the same population. Only one species K. pillara gen. et sp. nov. seems to be distinctive in colour and this is due to the unique banding pattern of the legs. Anthophiloptera is distinctively greenish grey, supposedly resembling an Angophora leaf. Windbalea is uniformly light green with the base of the antenna purple and the male stridulatory area dark brown.

Eggs

Recent investigation of the eggs of Australian tettigoniids reveal that they bear distinctive and useful taxonomic characters. This has been known for some time for the European species (see, for example, Hartley, 1990). For a number of reasons I have not had the opportunity to examine the eggs of all species included herein but have had the chance to study a number and they are as follows. In the Zaprochilinae, the eggs of Zaprochilus australis, Z. ninae, sp. nov., Kawanaphila nartee, ungarunya, gen. et spp. nov.; Anthophiloptera dryas Rentz and Clyne and Windbalea viridis, gen. et sp. nov. Those of Zaprochilus, Kawanaphila and Anthophiloptera are similar in that they each bear a cap at one and and the micropyles form a ring at the opposing end. Windbalea in unexpectedly different in its simplicity (Fig. 75A). The egg is very slender and without a cap and with only a uniform reticulate pattern on the surface and with micropyles (Fig. 75B) nearer one end. Even more surprising is the similarity of the egg Windbalea viride gen. et sp. nov. to that of Phasmodes ranatriformis (Fig. 18A). In the latter the egg is elongate, blunt at both ends and with a rather uniform lattice-like pattern over the entire surface. An area of micropyles is present near the narrower end of the egg (Fig. 18B). Eggs of a number of species of austrosagines have been examined.

Habitats

The taxa covered in this volume occur mostly in arid bushland and heath habitats. Only Anthophiloptera and some species of Zaprochilus occur in forest situations with the former probably being found in rainforests over a portion of its range. This results in most species in this volume being found in the southern portion of the continent, especially in southwestern Western Australia. With most efforts at habitat conservation being directed to eastern rainforests, or at least most of the publicity being given to those habitats, it is important that the plight of heathlands, mallee, and semi-arid bushlands be brought to the fore. They are undergoing more widespread destruction due to agriculture than any of the others. Clearing of heathlands for wheat and cattle grazing continues unrelentlessly. Recent efforts by the Save the Bush campaign is attempting to salvage some of the remnants of the biota of these areas but it may be too late for a number of species, including some katydids (see Appendix I).

Although the zaprochilines and phasmodines are not related to the austrosagines, they are linked because they often occur in the same habitats, although they are present at different stages in their life-history development. They are also present with a number of the tettigoniine species discussed in Volume 1 of this series. Some of the habitat photographs presented there are useful in illustrating habitats of species in this volume. Vol 1, Fig. 27 NE of Peak Charles illustrates a site where one might expect to find at least one species of the tiny zaprochiline Kawanaphila mirla, gen. et sp. nov. These nocturnal katydids could be found on the grasses and sedges (especially Lepidosperma spp.) at the bases of the shrubs in the background as adults at the same time as nymphs of the austrosagines Hemisaga denticulata (White) and H. venator, sp. nov. As they mature, these austrosagines will ascend the shrubbery more or less permanently. H. denticulata will assume sites slightly higher than those of H. venator such as in the shorter eucalypts with the H. venator sheltering in the Melaleuca sp. in the background. Females will return to the ground to oviposit. Pachysaga ocrocercus, sp. nov. could be found as