THE AUSTRALIAN ENTOMOLOGIST

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Cover: This undescribed species of Bombyliidae of the genus Docidomyia is

from the Goldfields Region of Western Australia. The genus belongs to the

subfamily Tomomyzinae, and has closest relatives in southern Africa and North

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Australian Entomologist 22 (2) August 1995

NOTES ON THE HABITAT, LYING ES AND

BEHAVIOUR OF AUSTROPHLEBIA COSTALIS (THEE ARDY

(ODONATA: AESHNIDAE) IN BRISBAN NE ROREST PARK,

QUEENSLAND P

PETER F. WOODALL

Department of Anatomical Sciences and Centre for Conservation Biology,

University of Queensland, Brisbane, Qld, 4072

Abstract

During surveys in Brisbane Forest Park (1991-1994), Austrophlebia costalis was recorded from

November to February. Most observations were from flowing streams in open or closed forest

but two were next to a dry, rocky creek-bed in eucalypt open forest. Three dawn-to-dusk

surveys in December, January and February indicated that A. costalis was active from 0600 to

1800 h EST with no marked crepuscular peaks in activity but low activity from 0900-1200 h

EST. The mean speed of the hawking flight was 6.2 km/hr and that of fast flight was 17.7

km/hr. The fastest timed speed was 34.1 km/hr.

Introduction

Austrophlebia costalis (Tillyard) is one of the largest Australian dragonflies

(abdomen 70 mm, hind-wing length 60-68 mm [Fraser 1960]) and its size,

chocolate brown body and the dark brown bands running along the costal

length of each wing make it quite distinctive. It has a wide distribution along

the east coast of Australia (Watson et al. 1991) but little is known of its

biology apart from the early observations of Tillyard (1916). However, it has

received some international consideration (Hocking 1953, Corbet 1962)

following the statements by Tillyard (1916, 1917) concerning its flight speed.

This paper provides additional observations on its habitat preference, daily

activity patterns and flight speed.

Study Area and Methods

Incidental observations were made during 12 km walked transects for avian

surveys in Brisbane Forest Park, west of Brisbane, conducted twice a month

from May 1991 to March 1994. The transects cover most of the major

vegetation types of Brisbane Forest Park (Young 1982) including closed

notophyll forest, closed microphyll forest, open eucalypt forest and woodland

and they cross some of the many small streams draining the park, including

those flowing east into Love Creek/Cedar Creek and Enoggera Creek.

More detailed observations were collected during dawn-to-dusk observations at

Love Creek (27?19'S 152?45'E, 600 m asl) where the shallow, rocky stream

flows in open forest with a mid-stratum of bangalow palm Archontophoenix

cunninghamiana. Records of flight speed were made here using a stopwatch

(accurate to 0.01 s) to time flights between prominent landmarks over

stretches of the stream (7-11 m) measured using a rangefinder (accurate to 0.2

m at this distance). This relatively short distance was used because I found it

very difficult to detect fast flying A. costalis at ranges greater than about 20

m in the conditions of low light intensity and it was essential to have a lead-

in to prepare for timing prior to the timed and measured distance.

34 Australian Entomologist 22 (2) August 1995

Results and Discussion

Habitat Selection

Most observations of this species were made at flowing streams in closed

(Greene's Falls) or open forest (Love Creek) surrounded by dense vegetation.

Nymphal exuviae identified as belonging to A. costalis were found at Love

Creek, indicating that it does breed here. However, on two occasions

(17.xii.1991, 28.xi1992) an adult was seen settled on shrubs adjacent to a dry

rocky creek bed in eucalypt open forest, between Boombana and Jolly's

Lookout, lacking palms or any other type of dense vegetation and having an

understorey of grass. Water flows in this section of the creek only for a short

period after heavy rainfall (pers. obs.) but further downstream it joins

Enoggera Creek which has permanent water and is surrounded by a thin strip

of closed forest.

Most previous descriptions of the habitat of this species from the Dorrigo

Tableland, NSW and Mount Tambourine, Qld. (Tillyard 1916) related to

flowing streams surrounded by dense vegetation (? closed forest) but, at

Dorrigo, Tillyard (1916) reported flushing two newly emerged females from

their perching sites "some hundred yards or more above the beginnings of a

small gully". These observations indicate that A. costalis will occasionally

move into more open, drier habitats away from the forest streams but the

reasons for these movements require further investigation.

Daily Activity

Four dawn-to-dusk (0400 - 1900 h EST) surveys were made at Love Creek on

30.xii.1992, 27.1.1993, 12/13.11.1993 and 26.xi.1993. A record of A. costalis

activity was deemed to be a flight either up or down the creek past the

observer. These were collected incidentally to a survey of avian calling

activity but it is considered unlikely that any A. costalis were missed due to

the other monitoring activity. Weather conditions during these days varied

from overcast conditions with showers to bright sunny periods. Even during

the sunny spells, 90-9596 of the creek was in shade due to the canopy of

palms. There was no obvious association between the weather and flight

activity; A. costalis was recorded flying during both drizzle and sunny periods.

No A. costalis were recorded during the November survey but results from the

other three days are presented in Fig. 1. There are records of A. costalis

activity from throughout the day, apart from the first two hours of light and

low activity from 1000 to 1200 h and there is no indication of any marked

crepuscular peak in activity.

The only observations on the activity patterns of A. costalis are given by

Tillyard (1916) who stated "The earliest time of day at which I saw this insect

in flight (apart from occasionally disturbing resting females in the morning)

was about 1 o'clock on a dull stormy day. They seldom fly at all until 4

pm but from that hour to sunset are particularly active". Other

statements, possibly repeating those of Tillyard (1916), are given by Fraser

(1960) "... probably mainly crepuscular"; Houston and Watson (1988) "...

Australian Entomologist 22 (2) August 1995 35

[ZZA dec 1992 HE Jan 1993 UZZ Feb 1993

No. of Records

4- 5- 6- 7- 8- 9- 10- 11- 12- 13- 14- 15- 16- 17- 18-

Eastern Standard Time

Fig. 1. Daily activity pattern for Austrophlebia costalis at Love Creek,

Brisbane Forest Park.

tending to crepuscular activity"; Watson er al. (1991) "... probably mainly

crepuscular". At Love Creek, A. costalis is active for a far greater period of

the day than these statements imply.

Speed of Flight

During observations at Love Creek, two types of flight were observed. One

was a slower hawking flight with a variable number of lateral deviations or

jinks' while the other was a fast direct flight, with no lateral deviations. The

mean speed of the hawking flight was 6.2 km/hr (s.e. 1.0, n = 9) and the

mean speed of the fast flight was 17.7 km/hr (s.e. 2.5, n = 11). The fastest

timed flight was of 34.1 km/hr but one untimed flight was considerably

faster.

The first account of the flying speed of A. costalis was given by Tillyard

(1916) "The distance from my watching-place, .... to the ledge of rock over

which the swift-flying males first appeared was, as near as I could judge,

about eighty yards. From the time they first appeared to the time they passed

me was barely sufficient for me to grip my net and steady myself to strike.

At the most it could only have been two or three seconds, which gives these

insects the incredible speed of about sixty miles an hour". This description

indicates that neither the distance nor the time were measured but it was

repeated by Tillyard (1917) as, "I had the opportunity once of timing it over a

measured stretch of between eighty and ninety yards. The distance was

covered in three seconds; so that this Dragonfly can fly at the rate of nearly

sixty miles per hour!" [c. 100 km/hr]. This value has been cited subsequently

as exceptionally fast but its validity has been questioned by Corbet (1962)

following calculations of Hocking (1953) which indicated a theoretical

maximum air-speed of 57 km/hr and a maximum sustained flight of 38 km/hr

36 Australian Entomologist 22 (2) August 1995

for A. costalis. However, Hocking (1953) acknowledged that factors such as

a following wind, downhill flight or the short distance could account for the

difference. It is also obvious that the original description by Tillyard (1916)

suggests far less precision than his subsequent statement (Tillyard 1917) and a

difference in the estimated time of one or two seconds would cause a major

difference to the final calculations of speed. The fastest speeds timed from

Love Creek were much lower than those of Tillyard (1916, 1917) and more in

line with the calculations of Hocking (1953) but my (untimed) observation

indicated that A. costalis is capable of even faster speeds, at least over short

distances.

General Behaviour

As Tillyard (1916) recorded, the species generally hangs vertically from

streamside vegetation, often on the underside of a tree-fern frond and in this

position with its dark coloration it is particularly difficult to see. Flying

activity of adults has been recorded from November to February in Brisbane

Forest Park, slightly extending the period of November to January given by

Fraser (1960) but Tillyard (1916) also records two specimens taken in

February. Tillyard (1916) records females ovipositing into small submerged

logs or twigs. This was preceded by a "quick to-and-fro" movement of the

abdomen which he interpreted as a sawing of the wood with the projecting

teeth of abdominal segment 10 to open the wood. I twice saw females

probing with the end of their abdomen into a fallen branch or the

roots/rhizomes exposed at the edge of the river bank. These were both out of

water; in the case of the branch it was above a small pool and in the case of

the roots it was within a few cm of the water's edge. I cannot be certain that

actual oviposition took place in either instance.

Acknowledgment

I am grateful to an anonymous referee for useful comments on the MS.

References

CORBET, P.S. 1962. A biology of dragonflies. Pp. 247. H.F. & G. Witherby, London.

FRASER, F. C. 1960. A Handbook of the Dragonflies of Australia. Pp. 67. Royal Zoological

Society of New South Wales, Sydney.

HOCKING, B. 1953. The intrinsic range and speed of flight of insects. Transactions of the

Royal Entomological Society of London 104: 225-345.

HOUSTON, W.W.K. and WATSON, J.A.L. 1988. Odonata. /n Zoological Catalogue of

Australia. Vol. 6. Pp. 33-132. (Vol. Ed. W.W.K. Houston). Australian Government Publishing

Service, Canberra.

TILLYARD, R.J. 1916. Life -histories and descriptions of Australian Aeschninae; with a

description of a new form of Telephlebia by Herbert Campion. Journal of the Linnean Society

(Zoology) 33: 1-83.

TILLYARD, R.J. 1917. The Biology of Dragonflies. Pp. 396. Cambridge University Press,

Cambridge.

WATSON, J.A.L., THEISCHINGER, G., and ABBEY, H.M. 1991. The Australian

Dragonflies. Pp. 278. CSIRO, Canberra.

YOUNG, P. 1982. Vegetation Classification /n: Vegetation Map - Brisbane Forest Park.

Department of Mapping and Surveying, Brisbane.

Australian Entomologist 22 (2) August 1995 37

THE LIFE HISTORY OF ZETONA DELOSPILA

(WATERHOUSE) (LEPIDOPTERA: LYCAENIDAE)

M.F. BRABY*

Department of Zoology, James Cook University of North Queensland, Townsville, Qld. 4811

Abstract

The early stages of Zetona delospila (Waterhouse) are described from northern Queensland.

Larvae feed at night on hemi-parasitic vines, Cassytha filiformis L. in Queensland and C.

glabella R.Br. in coastal Western Australia, which grow over tussocks of Triodia or Spinifex.

The distinctive pupal morphology provides support for the species current placement within

the Candalidini. Egg and larval morphology and the food plants of Z. delospila suggest a close

relationship to the Candalides (Erina) group of Cassytha-feeding species.

Introduction

Zetona delospila (Waterhouse) is an uncommon butterfly confined to the dry

tropical region of northern Australia. It has a wide, sporadic distribution

with most records from the far north-west, where it ranges from Broome and

the Edgar Range south-west of Kimberley to the Ord River at Queens Islet in

Western Australia (Common 1981, Common and Waterhouse 1981, Dunn

and Dunn 1991). In Queensland the species is known from five disjunct

areas: (1) 30 km W of Fairview (Monteith and Hancock 1977); (2)

Porcupine Gorge National Park 70 km NW of Hughenden (Valentine 1981,

Valentine and Johnson 1982); (3) 9 km WSW Torrens Creek and on the

Burra Range 18 km ENE of Torrens Creek about 140 km SW of Charters

Towers (Braby 1994); (4) on the Selwyn Ranges at Mt. Elliot Mine

approximately 140 km SE of Mt. Isa (1 9, 5.iii.1993) and at Selwyn Mine

approximately 160 km SE of Mt. Isa (1 C, 20.iv.1993) (T.A. Woodger,

pers. comm.); (5) Johnson Creek about 70 km NW of Mt. Isa (2 BC’, 2 99,

18.v.1993) (A. Eggleton, pers. comm.).

Adults of this distinctive lycaenid fly close to the ground amongst grass and

low shrubs (Common and Waterhouse 1981), often in open sandy areas or on

sandstone escarpments among clumps of spinifex (Triodia). The early stages

and general biology have not been reported. The purpose of this paper is to

document the life history, summarise available information on its biology

and comment on its taxonomic relationships with related Candalidini. Most

of the work described here is based on observations made on or near the Burra

Range (20?43'S, 145? 10'E) in northern Queensland during 2-3.v.1992 and 13-

14.v.1993.

Early Stages

Food plants. Cassytha filiformis L. in Queensland (Voucher: Braby AQ

625005, Queensland Herbarium Indooroopilly), C. glabella R.Br. in coastal

Western Australia (Lauraceae).

* Present address: CSIRO Division of Entomology, GPO Box 1700, Canberra, A.C.T., 2601

38 Australian Entomologist 22 (2) August 1995

Egg (Fig. 1). Hemispherical, whitish-grey, surface deeply pitted; ridges of

pits widely spaced, pits approximately hexagonal in shape; micropylar area

depressed. Diameter 0.6 mm, height 0.4 mm.

Final instar larva (Fig. 2). Thorax and abdominal segments 1-6 with red

middorsal line and a broad whitish-yellow subdorsal band, edged below by a

dark purple dorsolateral band; lateral surface green with a faint yellow

ventrolateral line. Abdominal segments 7-10 green with pronounced reddish-

purple middorsal line and a faint reddish-purple dorsolateral line; Newcomer's

organ present on segment 7 but weakly developed; raised areas on segment 8

but organs not everted. Prothoracic and anal plates green, somewhat

flattened; segments 9-10 narrower than segments 7-8; abdominal segment 10

distinctly U-shaped posteriorly. Head light brown, hidden beneath prothorax.

Body with numerous light brown setae; spiracles light brown. Length 11-12

mm.

Pupa (Figs. 3, 4). Elongate, pale cream-yellow with scattered small black

dots, especially on head and thorax. Head and abdomen with weakly

developed lateral flanges. Thorax with weakly developed dorsal ridge. A dark

reddish-brown middorsal line on head, thorax and abdominal segments 1-7,

more pronounced on metathorax and abdominal segments 1-2. Abdominal

segments with pink dorsolateral line edged above white; wings with series of

faint black longitudinal lines; two variable black lateral spots or patches on

abdominal segments 1-2. Spiracles black. Attached to silken pad by anal

hooks and central girdle. Length 10 mm, width 3 mm.

Observations

On the sandstone plateau of the Burra Range Z. delospila was limited to open

areas which supported eucalypt woodland and a fairly dense ground cover of

Triodia pungens R.Br. (Spinifex or Porcupine grass) (Fig. 6), upon which

the food plant grew as a hemi-parasitic vine. Within these areas Cassytha

filiformis occurred in relatively discrete dense patches, but elsewhere it did

not appear to be common. Adults collected from the Selwyn Ranges,

western Queensland, also were observed flying over patches of Cassytha that

were parasitic on Triodia pungens (T.A. Woodger, pers. comm.). On the

Burra Range Z. delospila occurred together with the more abundant

Candalides geminus Edwards & Kerr and C. erinus (Fabricius), which was

scarce. The early stages of C. geminus, however, were only located on the

larger vine Cassytha pubescens R.Br.

On the Burra Range eggs of Z. delospila were found singly on the flower

buds and young shoots of the food plant; on 3.v.1992, at 1100 h, a female

was also observed to deposit an egg beneath the stem of the food plant which

was tightly wrapped around a leaf of Triodia. Near Cable Beach, Broome,

Western Australia, females were observed (on 20.v.1981) ovipositing on the

flower buds and new growth of Cassytha glabella which was parasitising

Australian Entomologist 22 (2) August 1995 39

Figs. 1-6. Life history of Zetona delospila: (1) dorsal view of egg; (2) dorsal

view of final instar larva; (3, 4) dorsal and dorsolateral views of pupa; (5) adult

female; (6) habitat at the Burra Range. Scale = 0.3 mm for Fig. 1; 2.0 mm for Figs

2-4.

Spinifex (D.P.A. Sands pers. comm.). On the Burra Range larvae were

collected during the day by shaking clumps of Triodia which supported dense

clumps of the food plant. In captivity the larvae fed only at night; during the

day they possibly hide deep within the Triodia tussocks. Larvae were not

attended by ants. Two pupal shells (both parasitised) were found attached to

the stem of a small shrub festooned with the larval food plant: both pupae

were orientated in an upright position and situated about 10 cm above the

ground. No pupae were found amongst the leaf litter and debris déspite

extensive searching, although D.P.A. Sands (pers. comm.) located an empty

pupal case in a stiff curled leaf near Cable Beach, Western Australia. In

captivity, the duration of the pupal stage varied from 13 to 16 days (n=2 9).

40 Australian Entomologist 22 (2) August 1995

Adults (Fig. 5) were locally abundant on the Burra Range and nearly all

specimens captured on 13-14.v.1993 were in fresh condition. They flew

close to the ground, usually in close proximity to the food plant. Their

flight was generally weak and somewhat resembled that of C. erinus. Adults

were most active around midday, but only during sunny periods. During the

early hours of the morning and late afternoon they were very inactive and

settled mostly on stems of the Triodia tussocks for relatively long periods.

Sun basking was observed during the cooler hours of the morning (from

0900 to 1000 h); butterflies would typically open their wings at 90-135?

towards the sun whilst settled on small plants and other herbage near the

ground. Females were apparently more numerous than males during 13-

14.v.1993 (sex ratio of total captures: Q/9 1:5, n=24). Butterflies were

recorded feeding from a range of flowers: at Porcupine Gorge near Pyramid

Lookout I took a female feeding on flowers of Bursaria incana Lindley

(Pittosporaceae) at 1500 h on 1.v.1992; near Torrens Creek two adults were

taken on flowers of Waltheria indica L. (Sterculiaceae) at 1045 h on

2.v.1992; and on the Burra Range both sexes were feeding commonly on

flowers of Cassytha pubescens in company with Candalides geminus on 2-

3.v.1992.

Discussion

Z. delospila has a rather disjunct distribution and until quite recently very few

specimens were known; the species was considered rare by Common and

Waterhouse (1972). Its patchy occurrence may reflect the patchy distribution

of suitable food plant-grass associations (i.e. Cassytha-Triodia/Spinifex),

particularly since many of these associations are confined to sandstone

escarpments and open sandy areas which tend to be spatially patchy in nature.

Moreover, within these habitats colonies of Z. delospila appear to be very

localised.

In addition, Z. delospila appears to be quite seasonal and this also may partly

account for the paucity of records. The known (combined) flight period is

from March to August with one record from the Northern Territory in

October (Monteith and Hancock 1977). Adults were not flying on the Burra

Range in summer despite extensive searching during 12-13.1i.1994, although

much of the habitat had been recently destroyed by fire. In Queensland, most

sightings/captures have been in late autumn-early winter; in June 1977 on

the Triodia-covered sandstone outcrops near Fairview (Monteith and Hancock

1977), in May 1980-81 at Porcupine Gorge National Park (Valentine 1981,

Valentine and Johnson 1982), in May 1993 on the Johnson Creek north-west

of Mt. Isa (A. Eggleton pers. comm.). Appearance of many fresh specimens

in May 1992/93 at the Burra Range suggests the main emergence occurs in

autumn.

The species delospila currently is placed in the monotypic genus Zetona,

erected by Waterhouse (1938) after a long period of generic confusion. In his

Australian Entomologist 22 (2) August 1995 41

original description of the species, Waterhouse (1903) provisionally placed

delospila under Zizera Moore based on a single female, but noted that it was

quite distinct from any described species that belonged to the 'grass-blues'.

Waterhouse (1938) later suggested that Zetona may be related to Lucia

Swainson, based on the superficial resemblance of the cell spots of the wing

underside, but only five specimens were available then for comparison.

Common and Waterhouse (1972) felt Zetona was still related to Zizeeria

Chapman, Zizu/a Chapman and allied genera, but in a detailed study Eliot

(1973) showed that the wing venation and male genitalia of Zetona were

more closely related to Candalides Hübner sens. lat. and he placed the genus

in the newly erected tribe Candalidini.

Evidence reported here on the early stages of Z. delospila clearly supports

Eliot's view, particularly the shape and form of the pupa which is perhaps the

most distinctive feature of the tribe. The pupae of these species are

characterised by several features: (1) a lateral flange on the head, which may

be strongly indented in the middle; (2) a lateral flange on the abdomen, which

sometimes is upturned; (3) usually with a dorsal ridge on the thorax and

abdomen; and (4) often with a pair of dorsal projections on the thorax. Z.

delospila shares many of these characters, although the dorsal ridge is less

pronounced on the thorax (and absent on the abdomen) and the flanges are not

as strongly developed as in most other members, for example Candalides

cyprotus (Olliff) (Atkins and Heinrich 1987). The pupal shape of Z.

delospila somewhat resembles that of C. acastus (Cox) (see Fisher 1978,

Fig. 81e), which is also elongate with a relatively narrow abdomen.

Structurally, the pupa is perhaps closest to C. geminus which also has the

lateral flanges less pronounced and the dorsal ridge weakly developed

compared with C. hyacinthinus (Semper) and C. erinus (Braby unpubl. data).

The association of the early stages of Z. delospila with Cassytha spp. also

indicates an ecological relationship with other members of Candalides,

particularly those of the Erina Swainson species group (Tite 1963) which

feed as larvae exclusively on Cassytha (e.g. Common and Waterhouse 1981,

Fisher 1978). The early stages of Z. delospila are also very similar in

morphology to these species. For example, the egg closely resembles that of

Candalides geminus (which has the same shape and pit structure but is

considerably larger, about 0.8 mm dia. (Braby unpubl. data)), but differs quite

substantially from those of Nesolycaena albosericea (Miskin) and C.

absimilis (Felder), both of which have triangular shaped pits with raised

blunt projections at the pit junctions (Sands 1971, Braby unpubl. data). The

larva of Z. delospila is rather colourful and similar in pattern to C. geminus.

However, unlike C. geminus, Z. delospila lacks the raised dorsal reddish

spots on abdominal segments 1 to 6 and does not possess the distinctive

broken white dorsolateral line on the thoracic segments and abdeminal

segments | to 6 (Edwards 1980). Larvae of these two species are more

brightly coloured than those of C. hyacinthinus, C. erinus and C. acastus

which are green and less conspicuously striped.

42 Australian Entomologist 22 (2) August 1995

Until a detailed comparative study is undertaken on the Candalidini,

particularly the Erina (Cassytha-feeding) species group, Z. delospila is best

retained in its current (separate) genus for the present.

Acknowledgments

I am very grateful to Tony Eggleton, Don Sands and Terry Woodger for

providing their unpublished observations. Don Sands Kindly commented on

an earlier draft of the manuscript.

References

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eastern Queensland - Part III. Queensland Naturalist 32: 121-129.

COMMON, I.F.B. 1981. Part IV. Insects. Pp. 60-67. In McKenzie, N.L. (ed.). Wildlife of the

Edgar Ranges area, south-west Kimberley, Western Australia. Wildife Research Bulletin of

Western Australia No. 10.

COMMON, I.F.B. and WATERHOUSE, D.F. 1972. Butterflies of Australia. Angus and

Robertson, Sydney.

COMMON, I.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Revised Edition.

Angus and Robertson, Sydney.

DUNN, K.L. and DUNN, L.E. 1991. Review of Australian Butterflies: distribution, life history

and taxonomy. Parts I & III. Privately published by the authors, Melbourne.

EDWARDS, E.D. 1980. The early stages of Adaluma urumelia Tindale and Candalides

geminus Edwards and Kerr (Lepidoptera: Lycaenidae). Australian Entomological Magazine 7:

17-20.

ELIOT, J.N. 1973. The higher classification of the Lycaenidae (Lepidoptera): a tentative

arrangement. Bulletin of the British Museum of Natural History (Entomology) 28: 373-505.

FISHER, R.H. 1978. Butterflies of South Australia. Government Printer, South Australia.

MONTEITH, G.B. and HANCOCK, D.L. 1977. Range extensions and notable records for

butterflies of Cape York Peninsula, Australia. Australian Entomological Magazine 4: 21-38.

SANDS, D.P.A. 1971. The life history and taxonomic relationships of Nesolycaena albosericea

(Miskin) (Lepidoptera: Lycaenidae). Journal of the Australian Entomological Society 10: 290-

292.

TITE, G.E. 1963. A revision of the genus Candalides and allied genera (Lepidoptera:

Lycaenidae). Bulletin of the British Museum of Natural History (Entomology) 14: 197-259.

VALENTINE, P.S. 1981. In search of Zetona. Brolga. Newsletter of the Wildlife Preservation

Society of Queensland, Townsville Branch. 11(5): 3-4.

VALENTINE, P.S. and JOHNSON, S.J. 1982. New records of Lycaenidae and Hesperiidae

(Lepidoptera) from northern Queensland. Australian Entomological Magazine 9: 1-3.

WATERHOUSE, G.A. 1903. Notes on Australian Rhopalocera: Lycaenidae. Part III. -

Revisional. Proceedings of the Linnean Society of New South Wales 28: 132-275.

WATERHOUSE, G.A. 1938. Notes on Australian butterflies in the Australian Museum. No. 1.

Records of the Australian Museum 20: 217-222.

Australian Entomologist 22 (2) August 1995 43

DEVELOPMENT OF THE CAUDAL LAMELLAE IN

AUSTROARGIOLESTES ISABELLAE THEISCHINGER AND

O'FARRELL (ODONATA: MEGAPODAGRIONIDAE)

Ken Murray

41 Stanley St, Croydon Park, N.S.W., 2133

Abstract

The caudal lamellae in larval Austroargiolestes isabellae Theischinger & O'Farrell are

described and illustrated. The appendages are strongly nodate for much of the larval

development.

Introduction

Caudal lamellae, the hypertrophied paraprocts and appendix dorsalis are found

at the end of the abdomen in larval damselflies. They are believed to function

as external gills, aids to swimming and in enhancing displays (Johnson

1991). The development of these organs has been described in general terms

by Tillyard (1917a, b) and MacNeill (1960). There was however, little

information on caudal lamellae in the Megapodagrionidae and the

development of the Australian megapodagrionids remained undescribed.

Tillyard (19172) described and figured the lamellae of the final instars of two

Australian species of Argiolestes. He remarked on the unusual dorso-

ventrally flattened structure of these organs in Australian Megapodagrionidae,

a unique feature in his experience. Although Tillyard never saw the early

stages he concluded that the lamellae showed evidence of this type originating

from a two-jointed form. This supposition is confirmed in the observations

presented here.

Methods

Exuviae of Austroargiolestes isabellae Theischinger & O'Farrell were

collected during the rearing of this species (Murray 1992). Lamellae of

instars 2-8 were studied on exuviae placed in a small Petri dish containing

water. Final stage lamellae were removed from one exuvium and mounted on

slides with Euparal. Drawings were made using a 'Vixen' stereo-microscope.

Results

The first instar (prolarva) was not seen and observations began with second-

instar larvae. The morphology of the second-instar larva is remarkably

uniform throughout the Zygoptera and the caudal lamellae of A. isabellae had

the characteristic single-segmented style-like form, covered with setae (Fig.

1).

At the moult to the third instar the caudal lamellae became strongly nodate,

appearing two-segmented, with a club-shaped basal and a style-like distal

portion (Fig. 2). Both portions were of almost equal length. Widely spaced

small setae marked the lateral carina and fringed the margin of the proximal

44 Australian Entomologist 22 (2) August 1995

dele E EC Id ag, ve of

"X DA

` - -

tl P . r ?] : z r

Meh halal, Les A LL n Gd la aan

Figs 1-9. Development of the caudal lamellae in Austroargiolestes isabellae,

dorsal view. All except Fig. 9 are of the left lamella: (1) single-segmented style-

like appendage of second instar; (2) two-segmented (nodate) appendage of third

instar; (3-6) nodate form in fourth to seventh instars; (7) penultimate instar; (8)

final instar; (9) median lamella of final instar. (Not to scale).

Australian Entomologist 22 (2) August 1995 45

portion. A scattering of setae was present over the surface. In early instars

lamellae were spread in the usual way and later they were held horizontally.

Through subsequent moults the basal portion of the lamellae grew and

became progressively more flattened and leaf-like in appearance. There was a

gradual increase in setae on the lateral carina and about the margins. Both

short and long setae occurred, the latter concentrated around the more rounded

margins of the segments and on the lateral carina, especially through the later

stages of development.

In contrast with the expansion of the basal portion, the style-like distal

portion shortened (in absolute terms) with each moult, reducing in relative

size from nearly half the length of the basal portion in the fourth instar (Fig.

3) to approximately one quarter in the fifth instar (Fig. 4) and thereafter was

minute in comparison with the basal portion (Figs 5, 6). The style was lost

completely by the penultimate instar (Fig. 7), being replaced by a small

tapering tip to the basal portion. This pointed tip in turn was lost at the

moult to the final instar when lamellae were subequal and uniformly rounded

atthe tip. The lateral lamellae were narrow at the base and broader distally

and were asymmetrical about their long axis, with the more convex margin of

each lamella held outermost (Fig. 8). The median lamella was symmetrical

in shape, with lateral margins mostly straight and expanding from a narrow

base to a broader distal end (Fig. 9). In living animals the median lamella

was held at a slight angle to the laterals when viewed laterally.

Discussion

The Megapodagrionidae have a circumtropical distribution but both the higher

taxonomic positioning of and relationships within the family are very

uncertain (Houston and Watson 1988). Australian forms are characterised by

the unusual dorso-ventral flattening and horizontal positioning of the caudal

lamellae of the larvae. Confirmation that A. isabellae develops through a

highly nodate saccoid stage may throw light on the relationships between the

Australian Megapodagrionidae and other problematic taxonomic groups in the

region.

Acknowledgments

I thank Dr R.J. Rowe, Department of Zoology, James Cook University,

Townsville for generous assistance with the manuscript and one anonymous

referee for helpful comments. Thanks also to Max Moulds, Australian

Museum, for help with reference material.

References

HOUSTON, W.W.K. and WATSON, J.A.L. 1988. Odonata. In HOUSTON, W.W.K. (ed.)

Zoological Catalogue of Australia Vol. 6, pp. 33-132.

JOHNSON, D.M. 1991. Behavioural ecology of larval dragonflies and damselflies. Trends in

Ecology and Evolution 6: 8-13.

46 Australian Entomologist 22 (2) August 1995

MACNEILL, N. 1960. A study of the caudal gills of dragonfly larvae of the suborder

Zygoptera. Proceedings of the Royal Irish Academy 61(B)(7): 115-140.

MURRAY, K. 1992. A note on the rearing of Austroargiolestes isabellae Theischinger &

O'Farrell (Odonata: Megapodagrionidae). Australian Entomological Magazine 19: 49-50.

TILLY ARD, R.J. 1917a. On the morphology of the caudal gills of the larvae of zygopterid

dragonflies. Part i (General morphology) and Part ii (Studies of the separate types).

Proceedings of the Linnean Society of New South Wales 42: 31-112.

TILLY ARD, R.J. 1917b. On the morphology of the caudal gills of the larvae of zygopterid

dragonflies. Part iii (Ontogeny) and Part iv (Phylogeny). Proceedings of the Linnean Society of

New South Wales 42: 606-632.

Australian Entomologist 22(2) August 1995 47

A GENITALIC ABERRATION OF

TRAPEZITES PRAXEDES (HESPERIIDAE: TRAPEZITINAE)

FROM NEW SOUTH WALES

A.F. ATKINS! and C.N. SMITHERS?

! The University of Newcastle, Callaghan, N.S.W. 2308

? Australian Museum, College St., Sydney, N.S.W. 2000

Abstract

A male specimen of the Australian endemic skipper Trapezites praxedes (Plótz) collected from

the Barrington Range, central-eastern New South Wales, shows remarkable morphological

differences in the genitalia. The entire structure is 'stretched' to a narrow, elongate form,

although in detail each component is present and unaltered in proportion when compared with

the genitalia of typical male specimens from nearby coastal localities. The comparative male

genitalia are illustrated.

Introduction

The known distribution of Trapezites praxedes (Plótz) (see Sands et al. 1984)

is eastern Victoria, coastal and montane New South Wales (Common and

Waterhouse 1981), a few occurences in southern Queensland and an

unconfirmed, isolated record from central Queensland (Dunn and Dunn 1991).

Trapezites praxedes is currently under taxonomic review by one of us (AA).

The holotype is from the Sydney area (‘Port Jackson') (Waterhouse 1932).

Specimens very similar to the type occur along the coastal and near-coastal

fringe of New South Wales, north to at least Port Macquarie (R. Mayo, pers.

comm.) and in Queensland as far north as Fraser Island (pers. obs.). Another

form, possibly a distinct taxon, is found in montane areas north from the

Barrington district to at least southern Queensland (unpubl. data).

The recent capture of a male similar to the typical form, in the Barrington

Range, is thus significant. The slightly worn specimen (now in the

Australian Museum, Sydney) was collected on 24.iii.1991 at "Tuglo' near Mt

Royal, by A.S. Smithers. Dissection of the genitalia showed, however, that

the structure was unlike that of any known trapezitine species.

Genitalia

All the components of the genitalia are present (Figs 1-3) and can be

compared with those of the typical form (Figs 4-6), but the general structure,

particularly the distal section, appears drawn out or stretched in an almost

‘plastic’ way.

Combined tegumen and uncus shorter than valva, laterally narrow and

posteriorly elongate, curved downward but slightly upward at uncus which is

rounded and blunt, slightly raised lateral flanges; vinculum normal, beak

shaped; valvae somewhat asymmetrical (right valva narrower), shaped as in

that of typical form, but narrow and elongate, tapered at posterior end with

lower section (harpe) extending beyond upper to a pointed sclerotized

48 Australian Entomologist 22(2) August 1995

Figs 1-6. Male genitalia of Trapezites praxedes. (1-3) Aberration from Tuglo,

N.S.W.: (1) lateral view of inside right valva, aedeagus and uncus; (2) ventral view

of uncus tip; (3) lateral view of inside left valva. (4-6) Typical form from

Whitebridge, N.S.W.: (4) lateral view of inside right valva, aedeagus and uncus;

(5) ventral view of uncus tip; (6) lateral view of inside left valva.

— —

Australian Entomologist 22 (2) August 1995

projection, a recurved flattened process on inner edge; aedeagus as in typical

form but tapered posteriorly.

Discussion

Trapezites praxedes varies marginally throughout a range of biomes along the

central coast of New South Wales. The skipper most frequently occurs in

open, mixed woodlands with a heavy heath understory, from sea-level to 500

m, in areas where the larval foodplant Lomandra Labill. (especially L. obliqua

[Thumb.]) is abundant (Atkins 1993). The locality at Mt Royal contains a

mixed variety of forest types, especially wet sclerophyll and rainforest. The

aberrant male specimen was collected at approximately 700 m, beyond the

normal altitude of this form, but may have originated from sclerophyll

woodlands on the lower foothills. To our knowledge, this is the first record

of the typical coastal form from the Barrington area.

Only one other aberration of T. praxedes is known. In December 1967, A.

Sibatani collected an unusual male specimen near Coffs Harbour, northern

New South Wales, in which both upper and lower wing surfaces had broadly

exaggerated maculations (Sibatani 1970). Sibatani stated "Its antennae, head

(palpi missing), body, wing shape and genitalia (dissected) are similar to

those of Trapezites maheta praxedes (Plótz)." It is interesting that both

aberration and a comparative male specimen (collected in December 1968 at

the same locality, Coffs Harbour), illustrated by Sibatani (1970), are in fact

atypical of the coastal form of this species. Conversely, the Mt Royal

aberration, in appearance, closely resembles male coastal specimens from the

Hunter Valley district.

Acknowledgments

We thank Mr Ted Edwards, CSIRO, for alerting us to A. Sibatani's paper and

for discussion. Our thanks also to Mr Michael Braby, Mr Rod Eastwood, Mr

Russel Mayo and Dr Grant Miller for discussions on T. praxedes and to Dr

David McAlpine of the Australian Museum, Sydney, for the loan of

specimens.

References

ATKINS, A. 1993. Biological, behavioural, and distributional records of some butterflies from

New South Wales. Victorian Entomologist 23: 54-56.

COMMON, L.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Revised Edition.

Angus & Robertson, Sydney.

DUNN, K.L. and DUNN, L.E. 1991. Review of Australian Butterflies: distribution, life history

and taxonomy. Privately published, Melbourne.

SANDS, D.P.A., MILLER, C.G., KERR, J.F.R. and ATKINS, A.F. 1984. The specific status of

Trapezites praxedes (Plotz) (Lepidoptera: Hesperiidae): previously considered to be a

subspecies of T. maheta (Hewitson). Australian Entomological Magazine 11: 27-33.

SIBATANI, A. 1970. An aberrant form of Trapezites maheta praxedes (Plotz) (Lepidoptera:

Hesperiidae) from New South Wales. Journal of the Australian Entomological Society 9: 67-

68.

WATERHOUSE, G.A. 1932. Australian Hesperiidae. II. Notes and descriptions of new forms.

Proceedings of the Linnean Society of New South Wales 57: 218-238.

50 Australian Entomologist 22 (2) August 1995

Erratum

Atkins, A. 1994. A new genus Herimosa (Lepidoptera: Hesperiidae:

Trapezitinae) and its relationship to the Proeidosa group of endemic

Australian skippers. Australian Entomologist 21: 143-152.

In paragraph 2, page 148, 'The adult males of three species of Anisynta (A.

sphenosema, A. cynone and A. tillyardi ) and Antipodia have a sex-brand (or

stigma) on the upperside of the forewing' should read 'The adult males of two

species of Anisynta (A. dominula and A. monticola ) and Antipodia have a

sex-brand..'.

Australian Entomologist 22 (2) August 1995 51

RECORDS OF ACULEATE WASPS FROM FLOWERING

SUBTROPICAL RAINFOREST TREES

G. WILLIAMS? and P. ADAM!

! School of Biological Science, University of New South Wales,

P.O. Box I, Kensington, N.S.W., 2033.

2 Entomology Department, The Australian Museum,

6-8 College Street, Sydney, N.S.W., 2000.

Abstract

More than 88 species of aculeate wasps are recorded visiting flowers of 11 species of trees in

lowland subtropical rainforest or rainforest - wet sclerophyll forest ecotone sites in northern

New South Wales. The tiphiid subfamily Thynninae was the most diverse taxon with 31

species collected.

Introduction

Adult aculeate wasps feed on nectar (Naumann 1991, Houston 1984) and, in

Australia, have been recorded visiting flowers of sclerophyllous plants

(Brown 1987, 1989a, b; Armstrong 1979; Houston 1984; Hawkeswood

1981; Allsopp 1992; Webb 1989) and pollinating orchids (Armstrong 1979

and references therein). Apart from a record of the scoliid Campsomeris

tasmaniensis (Saussure) as a pollinator of Macadamia (Proteaceae) (Vithanage

and Ironside 1986) and aculeates as pollinators of Neolitsea dealbata

(Lauraceae), Litsea leefeana (Lauraceae) and Diospyros pentamera (Ebenaceae)

trees in tropical Queensland (House 1985), there are few plant-visiting records

for higher wasps in Australian rainforests.

Methods

Wasps of the families Pompilidae, Scoliidae, Sphecidae, Tiphiidae and

Vespidae were collected by hand-netting as they fed on tree blossoms in

seven lowland subtropical rainforest sites (sensu Adam 1992) and two

rainforest-wet sclerophyll ecotone sites in northeast New South Wales

(Tables 1, 2). A description of the collection sites, their floristic

composition and location is given in Williams (1993) and Williams and

Adam (1991). Wasps were observed and collected during three seasons from

late 1990 until early 1993. Only wasps that alighted on blossoms were

included in the study. Representatives of each species were examined for the

presence of pollen.

Twenty species of mass-flowering rainforest trees, possessing unspecialised

entomophilous floral morphologies (see Williams and Adam 1994), were

included in the study but particular species did not necessarily flower during

each season. In addition, the flowering patterns of individual tree species and

populations were seasonally heterogeneous so that it was not possible to

equally replicate sampling effort. Phenological data on flowering will be

published elsewhere.

Australian Entomologist 22 (2) August 1995

Results and Discussion

The number of wasp species attracted to individual tree species is given in

Table 1 and flower-visiting records in Table 2. More than 50 percent of the

species listed in Table 2 were recorded from only 1 or 2 specimens. Aculeate

wasps were not collected from nine of the 20 trees included in the study:

Cassine australis (Celastraceae) (Manning Point), Caldcluvia paniculosa and

Schizomeria ovata (Cunoniaceae) (Lorien Wildlife Refuge), Elaeocarpus

obovatus (Lansdowne Reserve) and E. reticulatus (Elaeocarpaceae) (Lorien

Wildlife Refuge), Drypetes australasica (Euphorbiaceae) (Manning Point),

Scolopia braunii (Flacourtiaceae), Cyptocarya microneura (Lauraceae) and

Acradenia euodiiformis (Rutaceae) (Lorien Wildlife Refuge).

Table 1. Number of wasp species recorded visiting plants. "*" indicates records

from rainforest-wet sclerophyll forest ecotone sites, all other records from

rainforest only.

Plant Taxa — Site g No. spp.

Alphitonia excelsa Harrington 60

Tristaniopsis laurina Wingham Brush 23

Tristaniopsis laurina Lorien Wildlife Refuge* 13

Euroschinus falcata Saltwater Reserve 10

Waterhousea floribunda Wingham Brush 8

Euroschinus falcata Harrington 6

Acmena smithii Harrington 6

Alectryon coriaceus Manning Point 6

Acmena smithii Manning Point 5

Cuttsia viburnea Lorien Wildlife Refuge 4

Acmena smithii Woko National Park 4

Alphitonia excelsa Kenwood Wildlife Refuge* 4

Diospyros australis Lansdowne Reserve 3

Guioa semiglauca Harrington 3

Waterhousea floribunda Lorien Wildlife Refuge 2

Diospyros australis Lorien Wildlife Refuge l

Abrophyllum ornans Lorien Wildlife Refuge* l

Rhodomyrtus psidioides Lorien Wildlife Refuge* l

Alectryon coriaceus Harrington l

The number of wasp species varied between both plant taxa and collection

sites (Tables 1, 2) and the total number of wasps recorded in Table 2 was not

present at any single site. Tiphiidae was the most diverse family (n= ca 36

spp.) and Alphitonia excelsa (Rhamnaceae) flowers (in rainforest) at

Australian Entomologist 22 (2) August 1995 53

Harrington were visited by the highest number of wasp species (n= >60). A.

excelsa flowers are distinctive in that they produce a fetid odour rather than a

sweet fragrance. At other species of flowering trees (and A. excelsa trees at

Kenwood Wildlife Refuge) there were fewer aculeate wasps and, on these, the

anthophilous insect fauna was generally dominated by mixed pollinator

guilds principally comprising bees, Diptera, Coleoptera and Thysanoptera.

Aculeate wasps occasionally occurred in large numbers on flowering trees

(e.g., Alphitonia excelsa at Harrington) but did not appear to disturb co-

foraging native bees.

All the aculeates examined carried pollen conspecific with that of the flowers

from which they were collected; however, individual pollen loads varied and

frequently contained pollen of more than one species, indicating that

individual wasps may not exhibit foraging fidelity. Data in Table 2 also

indicate a general lack of fidelity. Large wasps may undertake relatively

long-distance interplant flights exceeding 200 m (G.Williams, pers. obs.) and

potentially contribute to out-crossing in subtropical rainforest tree

populations, which typically consist of widely scattered individuals.

Few studies have addressed the contribution of anthophilous wasps to the

breeding systems of rainforest plants. However, several ecological studies

provide data on wasps as flower visitors in non-rainforest plant communities.

Petanidou and Ellis (1993), in a 30 ha Greek 'phrygana' shrubland plot,

collected 21 species of Sphecidae and 15 species of Vespidae. Heithaus

(1979) collected Pompilidae (12 spp.), Scoliidae (3 spp.), Sphecidae (39

spp.), Tiphiidae (10 spp.) and Vespidae (52 spp.) from Costa Rican savannah

and tropical deciduous and oak forests [these are similar to 'dry rainforest'

under Australian closed forest classifications and were regarded as rainforest

by Schimper (1903)], but >50 percent of species were represented by only 1

or 2 individuals. Inoue et al. (1990) recorded Scoliidae (1 sp.), Sphecidae (16

spp.) and Vespidae (11 spp.) from a Japanese temperate deciduous forest and

Kato et al. (1990) recorded Sphecidae (10 spp.) and Vespidae (5 spp.) from

Japanese primary beech forest (presumably in these forests the pollinators are

understorey specialists as the canopies are wind pollinated). Vespidae (5

spp.) and Sphecidae (2 spp.) were collected by Kato et al. (1993) from

flowers in Japanese cool-temperate subalpine forests and meadows.

In comparison with the records of Heithaus (1979), Inoue et al. (1990) and

Kato et al. (1990, 1993), Tiphiidae were well represented in our study region

and comprised approximately 41 percent of the wasp diversity (species

richness: n= >88 spp.). However, this at least in part may be due to the

greater representation of tiphiids in Australia; Tiphiidae constituted «896 of

wasp taxa recorded by Heithaus (1979) from neotropical habitats. Most of

the tiphiids collected by us were Thynninae, which is a diverse and abundant

subfamily in Australia and occurs widely on Australian sclerophyllous plants

(Hawkeswood 1981; Brown 1987, 1989a, b; Armstrong 1979; Keighery

54 Australian Entomologist 22 (2) August 1995

1975). However, Inouye and Pyke (1988) did not record thynnines visiting

the 43 species of alpine flora they observed in Kosciusko National Park,

southern New South Wales. In Australian lowland subtropical rainforest and

adjoining ecotonal forests, Thynninae may be a significant component of the

anthophilous wasp community, which in palaearctic, nearctic and neotropical

forests is dominated by Sphecidae and Vespidae.

Table 2. Aculeate wasps collected from flowers of rainforest trees.

(number in parentheses = number of species).

Plant Species

| - Euroschinus falcata (Anacardiaceae). 2 - Diospyros australis (Ebenaceae).

3 - Abrophyllum ornans (Escalloniaceae). 4 - Cuttsia viburnea (Escalloniaceae).

5-Acmena smithii (Myrtaceae). 6 - Rhodomyrtus psidioides (Myrtaceae).

7 - Tristaniopsis laurina (Myrtaceae). 8 - Waterhousea floribunda (Myrtaceae).

9 - Alphitonia excelsa (Rhamnaceae). 10 - Alectryon coriaceus (Sapindaceae).

11 - Guioa semiglauca (Sapindaceae).

Sites

A - Harrington (32°52'30"S, 152?41'00"E).

B - Manning Point (31°53'30"S, 152?40'00"E).

C - Saltwater Reserve (approx. 12 km SE of Taree) (32°00'30"S, 152°33'45"E).

D - Lansdowne Reserve (0.5 km SE Lansdowne) (31°47'30"S, 152?32'30"E).

E- Lorien Wildlife Refuge (3 km N Lansdowne: 2 sites) (31?45'00"S,

152°32'30"E).

F - Kenwood Wildlife Refuge (4 km NNW Lansdowne) (31°44'45"S, 152°31'30"E).

G - Wingham Brush (Wingham) (31°52'40"S, 152°22'00"E).

H - Woko National Park (approx. 24 km NNW Gloucester) (31°49'00"S,

151°47'00"E).

Month of Collection

N (Nov.), D (Dec.), J (Jan.), F (Feb.), M (Mar.).

Species Plant/Site Month

POMPILIDAE

Chirodamus defensor Smith 8G D

Chirodamus ? raptor Smith 7E D

Chirodamus sp./spp. 7G(3), 8E, 9A(2) NDJ

?Chirodamus sp./spp. IC N

Chryptocheilus bicolor (Fabricius) 7G J

Chryptocheilus sp./spp. 9A(2) JE

?Chryptocheilus sp./spp. 8G, 9A DJ

Phanagenia fasciata Fabricius 1C N

Platyderes collaris (Fabricius) 9A F

Australian Entomologist 22 (2) August 1995

Table 2 (cont.). Aculeate wasps collected from flowers of rainforest trees.

Species Plant/Site Month

SCOLIIDAE

Campsomeris ?tasmaniensis (Saussure) 2E, 9A DF

Campsomeris zonata Smith UE D

Scolia verticollis (Fabricius) TE, 9A DJF

Scolia ?verticollis (Fabricius) 7G J

Scolia sp. nr. verticollis (Fabricius) 7G J

Scolia sp./spp. 1C, 7G, 9A(3) DJF

?Scolia sp./spp. IC D

SPHECIDAE

Acanthosthethus sp./spp. 1A, 9A(2) NJF

Bembicinus sp. 1A, 9A NF

Bembix ?kamulla Evans and Matthews 7E D

Bembix promontorii Lohrman 9A J

Bembix sp./spp. 5B, 9A(2), 10B(2) NDJF

Cerceris ?australis Saussure 7G D

Cerceris minuscula Turner 7G DJ

Cerceris sp./spp. 5H, 7E, 7G DJ

Ectemnius reginellus Leclercq 9A F

?Ectemnius sp. 8G(2), 9A NDF

Larra sp. 9A F

Pison sp. 9A F

Sceliphron laetum (Smith) 7G, 9A DF

Sericophorus sp. 9A J

Sphex ephippium Smith 9A F

Sphex fumipennis Smith 9A F

Sphex globosus Smith 9F F

Sphex ?luctuosus Smith 7E D

Sphodrotes sp./spp. 1A, 9A(2), 9F NJFM

?Sphodrotes sp./spp. 9A F

Spilomena sp./spp. 4E(2), 8E ND

Tachysphex sp. 1A, 1C, 9A(2), 9F NDJF

?Tachysphex sp./spp. 9A(2), 10B(2) DJFM

Williamsita sp./spp. 9A J

TIPHIIDAE

Anthoboscinae

Anthobosca australasiae Guerin 8G D

Anthobosca ?laevifrons (Smith) IC D

Anthobosca signata Smith 7E, 9A DJF

Anthobosca ?signata Smith 7E J

Diamminae

Diamma bicolor Westwood 9A J

Thynninae

Acanthothynnus ater Brown 9A F

Acanthothynnus ?ater Brown 9A E

Agriomyia maculata Guerin 7G J

Agriomyia manifesta Turner 5H N

56 Australian Entomologist 22 (2) August 1995

Table 2 (cont.). Aculeate wasps collected from flowers of rainforest trees.

Species

TIPHIIDAE (cont.)

Thynninae (cont.)

Agriomyia variegata Klug

Dimorphothynnus dimidiatus (Smith)

Epactiothynnus tasmaniensis (Saussure)

Epactiothynnus ?tasmaniensis (Saussure)

?Epactiothynnus sp.

Eirone ?parca (Turner)

Eirone sp. nr. parca (Turner)

Eirone schizorhina

Eirone sp. nov.

Hemithynnus apterus (Oliver)

Hemithynnus rufiventris (Guerin)

Lestricothynnus fravenfeldianus (Saussure)

Rhagigaster ?denticulatus (Turner)

Rhagigaster sp. nr. kiandrensis Guerin

Rhagigaster ?mutatus Turner

Rhagigaster ?unicolor Guerin

Thynnoturneria sanguinolentus (Turner)

Thynnoturneria sp. nr. umbripennis (Smith)

?Thynnoturneria sp.

Tmesothynnus dispersus (Turner)

Tmesothynnus iridipennis (Smith)

Zaspilothynnus sp. nr. campanularis (Smith)

Zeleboria contigua (Turner)

Zeleboria ?contigua (Turner)

Zeleboria sp. nr. contigua (Turner)

Zeleboria xanthorrhoei (Smith)

?Zeleboria sp.

VESPIDAE

Polistinae

Polistes humilis (Fabricius)

Polistes ?humilis (Fabricius)

Polistes tepidus (Fabricius)

Ropalidia sp./spp.

Eumeninae

Abispa splendida (Guerin)

Abispa ?splendida (Guerin)

Bidentodynerus bicolor (Saussure)

Deuterodiscoelius ephippium Saussure

?Epiodynerus sp./spp.

Leptomenoides sp./spp.

Paralastor sp./spp.

Pseudabispa confusa van der Vecht

Plant/Site Month

8G D

5A, 5B, 9A, 11A NF

9A FM

5A N

9A F

7G J

5H, 7E, 7G, 8G NDJ

7G D

7g J

5B N

7E F

7E D

9F F

SA, 9A, 11A NDJF

9A F

7G, 8G D

7G DJ

7G, 9A DJF

9A J

IC, 9A NF

7G D

9A F

5A N

5B, 9A, 11A NDF

IC N

5A, 5B, 9A NF

9A JF

IC, 6E, 9A, 10B N-M

4E, 9A DF

9A F

4E D

TE, 9A DM

7G J

7G D

2D D

2D(2), 7E, 9A DJ

9A J

1A(3), 1C, 3E, SA, 5H,

9A(3), 10A, 10B, NDJF

7G, 9A J

Australian Entomologist 22 (2) August 1995 57

Acknowledgments

Dr Graham Brown (Museum and Art Gallery of the Northern Territory,

Darwin) and Dr Ian Naumann and Ms Josephine Cardale (CSIRO, Canberra)

are thanked for assistance with identifications. Dr Brown is additionally

thanked for comments on our manuscript. The NSW National Parks and

Wildlife Service are thanked for permission to collect in Woko National

Park. Mr and Mrs R. Moylan and Miss L. Moylan (Lansdowne) kindly

permitted studies on Kenwood Wildlife Refuge. One of us (GW) thanks the

Australian Museum, Sydney and the Australian Entomological Society for

grants in aid of research.

References

ADAM, P. 1992. Australian Rainforests. Oxford Monographs on Biogeography, (eds)

George, W., Hallam, A. and Whitmore, T.C. Oxford University Press, Oxford: 293 pp.

ALLSOPP, P.G. 1992. Volatile compounds as attractants for Campsomeris tasmaniensis

(Saussure) (Hymenoptera: Scoliidae). Australian Entomological Magazine 19: 107-110.

ARMSTRONG, J.A. 1979. Biotic pollination mechanisms in the Australian flora - a review.

New Zealand Journal of Botany 17: 467-508.

BROWN, G.R. 1987. Revision of the Australian genus Acanthothynnus Turner (Hymenoptera:

Tiphiidae). Journal of the Australian Entomological Society 26: 181-188.

BROWN, G.R. 1989a. Revision of the Australian genus Doratithynnus Turner (Hymenoptera:

Tiphiidae). Journal of the Australian Entomological Society 28: 1-17.

BROWN, G.R. 1989b. The Australian genus Encopothynnus Turner (Hymenoptera:

Tiphiidae). Journal of the Australian Entomological Society 28: 255-266.

HAWKESWOOD, T.J. 1981. Notes on the pollination of Nuytsia floribunda (Labill.) R.Br.

(Loranthaceae) and some literature reviewed. The Western Australian Naturalist 15: 17-21.

HEITHAUS, E.R. 1979. Community structure of neotropical flower visiting bees and wasps:

diversity and phenology. Ecology 60: 190-202.

HOUSE, S.H. 1985. Relationships between breeding and spatial pattern in some dioecious

tropical rainforest trees. Ph.D thesis, Australian National University, Canberra: 350 pp.

HOUSTON, T.F. 1984. Bionomics of a pollen-collecting wasp, Paragia tricolor

(Hymenoptera: Vespidae: Masarinae), in Western Australia. Records of the Western

Australian Museum 11: 141-151.

INOUE, T., KATO, M., KAKUTANI, T., SUKA, T. and ITINO, T. 1990. Insect-flower

relationship in the temperate deciduous forest of Kibune, Kyoto: an overview of the flowering

phenology and the seasonal pattern of insect visits. Contributions from the Biological

Laboratory, Kyoto University 27: 377-463.

INOUYE, D.W. and PYKE, G.H. 1988. Pollination ecology in the Snowy Mountains of

Australia: comparisons with montane Colorado, USA. Australian Journal of Ecology 13: 191-

210.

58 Australian Entomologist 22 (2) August 1995

KATO, M., KAKUTANI, T., INOUE, T. and ITINO, T. 1990. Insect-flower relationship in

the primary beech forest of Ashu, Kyoto: an overview of the flowering phenology and

seasonal pattern of insect visits. Contributions from the Biological Laboratory, Kyoto

University 27: 309-375.

KATO, M., MATSUMOTO, M. and KATO, T. 1993. Flowering phenology and anthophilous

insect community in the cool-temperate subalpine forests and meadows at Mt. Kushigata in the

central part of Japan. Contributions from the Biological Laboratory, Kyoto University 28: 119-

172.

KEIGHERY, G.J. 1975. Parallel evolution of floral structures in Darwinia (Myrtaceae) and

Pimelea (Thymeleaceae). The Western Australian Naturalist 13: 17-19.

NAUMANN, I.D. 1991. Hymenoptera (wasps, bees, ants, sawflies). Pp. 916-1000 in CSIRO

(ed.), The Insects of Australia. Melbourne University Press: Carlron.

PETANIDOU, T. and ELLIS, W.N. 1993. Pollinating fauna of a phryganic ecosystem:

composition and diversity. Biodiversity Letters 1: 9-22.

SCHIMPER, A.F.W. 1903. Plant Geography upon a Physiological Basis. (English translation).

Clarendon Press, Oxford: 837 pp.

VITHANAGE, V. and IRONSIDE, D.A. 1986. The insect pollinators of Macadamia and

their relative importance. Journal of the Australian Institute of Agricultural Science 52: 155-

160.

WEBB, G.A. 1989. Insects as potential pollinators of Micromyrtus ciliata (Sm.) Druce,

Myrtaceae. Victorian Naturalist 106: 148-151.

WILLIAMS, G.A. 1993. Hidden Rainforests: subtropical rainforests and their invertebrate

biodiversity. New South Wales University Press & The Australian Museum, Sydney: 188 pp.

WILLIAMS, G.A. and ADAM, P. 1991. Rainforest remnants on headlands in the Manning

Valley: their composition and conservation significance. Wetlands (Australia) 11: 21-30.

WILLIAMS, G.A. and ADAM, P. 1994. A review of rainforest pollination and plant-

pollinator interactions, with particular reference to Australian subtropical rainforests.

Australian Zoologist 29: 177-212.

Australian Entomologist 22 (2) August 1995 59

ESTIMATION OF LARVAL INSTARS OF

HYPSIPYLA ROBUSTA MOORE (LEPIDOPTERA:

PYRALIDAE) BY LARVAL FRASS WIDTHS

J. MO and M.T. TANTON

Department of Forestry, The Australian National University, Canberra, ACT, 0200

Abstract

Frass widths are used to estimate the larval instars of Hypsipyla robusta Moore, a shoot borer

of Australian red cedar Toona australis (F. Muell. Harms. Based on the relationship betwen

larval frass widths and larval head capsule widthe, the inter-instar boundaries of larval frass

widths are calculated. The average frass widths of individual larvae are then compared with

the boundaries to determine the larval instars. Ninety-three percent of laboratory larvae were

assigned to their correct instars by frass widths and 87% of field larvae were estimated as in

the same instars by frass widths as that determined by head capsule widths.

Introduction

The Cedar tip moth Hypsipyla robusta Moore is a serious pest of a number

of Meliaceae species, including Australian Red Cedar Toona australis (F.

Muell.) Harms.* (Beeson 1919). The larvae feed inside various tissues of the

host plant, especially the growing shoots and pupate inside larval tunnels

(Beeson 1919). Due to their cryptic nature, the development stages of larvae

cannot be determined directly. As an indirect approach, this paper explores

the possibility of using larval frass widths (FW) to estimate larval head

capsule widths (HCW) and therefore larval instars. Such a technique may be

used by forresters in fine-tuning the timing of control measures against this

pest.

Frass of H. robusta larvae is found as conspicuous clumps at the openings to

the larval tunnels (Roberts 1968). The inside of the tunnels, however,

contains little frass (personal observation), suggesting that fresh frass is

constantly being pushed out. It is therefore possible to relate the exterior

frass to the current development stages of the larvae.

Methods

Larvae were obtained from a laboratory stock originating from mature larvae

collected in a red cedar plantation in Macksville, NSW and maintained on the

artificial diet of Couilloud and Guiol (1980). To enhance feeding, a small

amount of macerated fresh young red cedar shoots was incorporated into the

diet. Larvae from the original site were incorporated into the stock at least

twice a year.

One hundred newly-hatched larvae were reared separately in glass vials (50x12

mm) until pupation. The instar of a larva was determined from the number

of head capsules it shed. Ist to 3rd instar larvae were fed with the terminal

parts of young shoots whilst the older larvae were supplied with cuttings

*The name Toona ciliata M. Roem may soon replace Toona australis (F. Muell.) Harms.

60 Australian Entomologist 22 (2) August 1995

from the stouter parts of young shoots, in accordance with their natural

feeding habits (personal observation). Food was replaced every 1-3 days,

depending on the consumption rate and freshness of the tissue. Frass was

removed daily from the glass vials. Rearing was in a room with temperature

at 26+1°C and light period at 14L:10D. Humidity was not controlled, the

room maintained humid by a vaporiser (KAZ Model 76).

At least 20 larvae at every instar were measured for HCW and FW, to the

nearest 1/40 mm, under a stereo microscope fitted with an eyepiece scale.

Due to the frequent rupture of head capsules in the last moult, the HCW's of

the last instar larvae were replaced with the corresponding larval head widths

just before pupation. For each larva measured for HCW, 20 air-dried frass

pellets produced by that larva were measured and the mean FW calculated.

A separate set of data involving 30 larvae was collected in a red cedar

plantation at Macksville, NSW, to test the effectiveness of FW in estimating

larval instars in the field. The frass was transfered from the infested shoots to

glass vials and then the shoots were dissected for larvae. The frass and the

associated larvae were taken back to the laboratory and the larvae were further

reared to obtain the head capsules for HCW measurements.

Results

Larvae moulted either 5 (3296) or 6 (6896) times before pupation, as noted

previously by Atuahene and Souto (1983). A recent study by the authors

showed 5- and 6-instar forms in larvae of both sexes (82% and 75% of 6-

instar forms in males and females respectively), hence the variation in the

number of larval instars is not likely to be sex dependant.

Larvae of the 5- and 6-instar forms showed similar HCW ranges in the Ist to

5th instars. Hence data were pooled and the joint mean and ranges are given

in Table 1. Total separation was achieved by HCW for the first 4 instars,

whereas the 5th and 6th instar larvae showed some overlapping in their HCW

ranges. Further examination of the data showed that amongst the 25 5th

instar larvae measured, only one had its HCW range fall within that of the

6th instar larvae. Thus HCW can still be considered a reliable predictor of

larval ages. The inter-instar boundaries in HCW for any two non-overlapping

instars were arbitrarily determined as the average of the maximum HCW of

the former instar and the minimal HCW of the following instar, with that for

the 5th and 6th instars as the minimum HCW of the 6th instar (Fig. 1).

Overlapping in FW started in the 4th instar and the relative within-instar

variations (expressed as SE/mean) were consistently higher than that in HCW

(Table 1). However, FW showed apparent positive correlation with HCW

(Fig. 1) and the correlation was significant (t=51.37, df=133, p«0.001). The

relationship was well fitted by linear regression (Fig. 1). Assuming the

regression equation correctly described the true realtionship between HCW and

FW, the inter-instar boundaries of FW obtained by supplanting the HCW

Australian Entomologist 22 (2) August 1995 61

o ist instar [s] 3rd instar ^ 5th instar

o 2nd instar u 4th instar A 6th instar

Inter-instar boundaries in HCW

Average frass width (FW) (mm)

Inter-instar boundaries in FW

0 0:5 1 15 2 2.5 3

Head capsule width (HCW) (mm)

Fig. 1. The relationship between average frass width (FW) and larval head capsule width

(HCW) in Hypsipyla robusta, with their inter-instar boundaries shown as dotted lines. Data

from larvae reared on host plant material. (See text for details).

boundaries into the equation (Fig. 1) should perform equally well in

delimiting larval instars. In effect, the percentages of correct estimations of

larval instars by comparing individual FW's with the FW boundaries were

100% for the first 3 instars, 95% for the 4th instar, 7696 for the 5th instar

and 90% for the 6th instar. Overall, 93% of the measured larvae were

assigned to their correct instars by their FW's. Most of the misclassifications

occurred in the 5th and 6th instar, which is probably due to the overlapping

of the HCW ranges of these two instars.

With field data, 26 larvae (8790) were assigned to the same instars by both

HCW and FW. Two larvae that were assigned to the 5th instar by HCW

were estimated as 6th instar by FW and 2 larvae that were determined as 5th

instar by HCW were estimated as 4th instar by FW.

Discussion

The above analysis demonstrates that FW is a useful predictor of larval

instars of H. robusta, especially for the first 4 instars. The degree of

predictiveness is comparable with that of HCW. Since FW data are more

easily accessed than HCW data, the FW approach appears promising. When

applied to field situations, care should be taken to measure only those

62 Australian Entomologist 22 (2) August 1995

Table 1. Larval head capsule widths (HCW) and frass widths (FW) in

Hypsipyla robusta.

head capsule width (HCW) — — — frasswideh(FW) —— —

larval —

instar mean+SE(n) range mean+SE(n) range

(SE/mean) (SE/mean)

Ist — 0.290.01 (20) 0.28-0.30 0.050.00 (20) 0.05-0.06

(0.03) (0.05)

2nd 0.49+0.04 (20) 0.45-0.53 0.10+0.02 (20) 0.08-0.15

(0.08) (0.20)

3rd 0.85+0.08 (20) 0.70-0.95 0.25+0.03 (20) 0.16-0.29

(0.09) (0.12)

4th 1.24+0.13 (20) 1.05-1.40 0.39+0.06 (20) 0.30-0.48

(0.10) (0.15)

5th 1.78+0.15 (25) 1.53-2.20 0.59+0.11 (25) 0.46-0.93

(0.08) (0.19)

6th 2.25+0.15 (30) 2.00-2.25 0.77+0.10 (30) 0.60-1.02

(0.07) (0.13)

frass pellets of apparently larger sizes to minimise the possibility of

accidentally including frass pellets produced at earlier developmental stages.

The number of frass pellets required varies with instars. Under the

assumption of normal distribution of FW, a minimal number of 16 frass

pellets is recommended to keep the relative sampling error below 10%.

Finally, the inter-instar boundaries of FW given here are based on larvae

reared in an artificial environment. Although they were validated by one set

of field data, further validation and possibly modification may be needed

before widespread application of the method.

Acknowlegdment

We wish to thank Dr F.L. Bygrave (Division of Biochemistry and Molecular

Biology, ANU) for providing the field study site.

References

ATUAHENE, S.K.N. and SOUTO, D. 1983. The rearing and biology of the mahogany shoot

borer Hypsipyla robusta Moore (Lepidoptera: Pyralidae) on an artificial medium. Insect

Science and Application 4: 319-325.

BEESON, C.F.C. 1919. The life history of the Toon shoot and fruit borer, Hypsipyla robusta

Moore (Lepidoptera: Pyralidae: Phycitinae) with suggestions for its control. Indian Forest

Records 7(7): 1-216.

COUILLOUD, R. and GUIOL, F. 1980. The laboratory rearing of Hypsipyla robusta Moore.

Revue Bois et Foréts des Tropiques n? 194: 182-186.

ROBERTS, H. 1968. An outline of the biology of Hypsipyla robusta Moore, the shoot borer of

Meliaceae of Nigeria, together with brief comments on two stem borers and one other

lepidopteran fruit borer also found in Nigerian Meliaceae. Commonwealth Forestry Review

47: 225-232.

Australian Entomologist 22 (2) August 1995 63

A NEW DISTRIBUTION RECORD FOR

THECLINESTHES SULPITIUS (MISKIN) (LEPIDOPTERA:

LYCAENIDAE) IN THE NORTHERN TERRITORY AND

NOTES ON THE LIFE HISTORY

C.E. MEYER! and D.N. WILSON?

! 10 Anne Clark Ave, Nicholls, ACT 2913

2 PO Box 554, Palmerston, NT, 0831

Abstract

New distribution records for Theclinesthes sulpitius (Miskin) in the Northern Territory are

given and host plants identified.

Introduction

The salt pan blue Theclinesthes sulpitius (Miskin) is known to occur at

Andoom Creek near Weipa, the Claudie River and from Cooktown south

through coastal Queensland and New South Wales to Lake Wellington and

Manns Beach, Victoria and at Berri in South Australia (Common and

Waterhouse 1981, Dunn and Dunn 1991). There is no previous record of this

butterfly occurring in the Northern Territory.

Discussion

Adults were first collected by the authors in December 1991 and February

1992 on Shoal Bay Peninsula near Darwin, where the butterfly was found

flying around the salt bush Halosarcia indica (Willd.) (Family

Chenopodiaceae). Subsequent collecting trips to the Cox Peninsula region,

Elizabeth River and Leanyer swamps, near Darwin, located further colonies of

the butterfly and another host plant, Tecticornia australasica (Moq.) (Family

Chenopodiaceae).

Larvae were collected from the Shoal Bay Peninsula and Cox Peninsula sites

and successfully reared to adults. The appearance of the early stages of this

butterfly in the Northern Territory agrees with the description given in

Common and Waterhouse (1981). Larvae on both host plants are extremely

well camouflaged and difficult to locate. Larvae collected from the Cox

Peninsula site were tended by numerous small black ants.

The four colonies found to date suggest that populations of the butterfly

probably occur right across the coastal regions of the Northern Territory into

the gulf country of Queensland.

Acknowledgments

We wish to thank Ian Cowie of the Northern Territory Herbarium,

Palmerston for assistance in identifying the food plants and Rod Eastwood for

his enthusiastic support during the February 1992 collecting trip.

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DUNN, K.L. and DUNN, L.E. 1991. Review of Australian Butterflies: Distribution, Life History

and Taxonomy. Part IIl: Family Lycaenidae. Pp 336-512. Privately published by the authors,

Melbourne.

64 Australian Entomologist 22 (2) August 1995

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THE AUSTRALIAN

Entomologist

Volume 22, Part 2, 31 August 1995

CONTENTS

ATKINS, A.F. and SMITHERS, C.N.

A genitalic aberration of Trapezites praxedes (Hesperiidae: Trapezitinae) from

New South Wales.

BRABY, M.F.

The life history of Zetona delospila (Waterhouse) (Lepidoptera: Lycaenidae).

MEYER, C.E. and WILSON, D.N.

A new distribution record for Theclinesthes sulpitius (Miskin) (Lepidoptera:

Lycaenidae) in the Northern Territory and notes on the life history.

MO, J. and TANTON, M.T.

Estimation of larval instars of Hypsipyla robusta Moore (Lepidoptera: Pyralidae) by

larval frass widths.

MURRAY, K.

Development of the caudal lamellae in Austroargiolestes isabellae Theischinger

and O'Farrell (Odonata: Megapodaerionidae).

WILLIAMS, G. and ADAM, P.

Records of aculeate wasps from flowering subtropical rainforest trees.

WOODALL, P.F.

Notes on the habitat, flying speed and behaviour of Austrophlebia costalis

(Tillyard) (Odonata: Aeshnidae) in Brisbane Forest Park, Queensland.

RECENT LITERATURE

An accumulative bibliography of Australian Entomology

ENTOMOLOGICAL NOTICES Inside back cover.

ISSN 0311 1881