THE AUSTRALIAN ENTOMOLOGIST

The Australian Entomologist (formerly Australian Entomological Magazine) is a

non-profit journal published in four parts annually by the Entomological Society

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Editor: Dr D.L. Hancock

Dept of Primary Industries

Assistant Editors Dr G.B. Monteith

Queensland Museum

Mr G. Daniels

University of Queensland

Business Manager Dr A.P. Mackey

Queensland University of Technology

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Cover: This illustration shows a worker ant of the Iridomyrmex purpureus group.

The ants of this complex are better known as Australia's ubiquitous meat ants.

They form large colonies which live in broad flattened nests with multiple

entrances and with the surface decorated with small pebbles. Illustration by Geoff

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Australian Entomologist 23 (2) September 1996 33

FLOWER VISITATION BY THE BUMBLE BEE BOMBUS

TERRESTRIS (L.) (HYMENOPTERA: APIDAE) IN TASMANIA

T.D. SEMMENS

Department of Primary Industry and Fisheries, GPO Box 192B, Hobart, Tasmania 7001

Abstract

To date the bumble bee Bombus terrestris (L.), first detected in Tasmania in February 1992, is

known to have visited 156 introduced and 14 native species of plants there.

Introduction

The bumble bee Bombus terrestris (L.), first detected in Australia in a garden

in Battery Point, Hobart in February 1992 (Semmens et al. 1993), has spread

throughout most of Hobart, including the Eastern Shore and Collinsvale,

down into the Huon Valley, at Franklin and Cygnet, into the Channel area

south of Hobart; also to the west of Hobart to New Norfolk and Mt Field and

to the north of Hobart to Kempton. It appears to be thriving in southern

Tasmanian conditions. This species is one of 4 introduced into New Zealand

from the United Kingdom in 1885-1906 for pollination purposes, particularly

of red clover; it may have spread to Australia from New Zealand.

Flower visitation

The flower visiting rate of Bombus terrestris is much higher than for other

bees (Donovan and Macfarlane 1984). In Tasmania at least 170 plant species

have been visited - 156 introduced and 14 native species (Table 1). This

agrees with findings in New Zealand, where Bombus terrestris was reported

to have visited at least 400 introduced and 19 native species (Donovan and

Macfarlane 1984).

Table 1. Plant species visited by the bumble bee Bombus terrestris in Tasmania to

July 1996.

Native Species

Acacia pravissima (Ovens Wattle) Eucryphia lucida (Leatherwood)

Banksia marginata (Banksia) Exocarpos cupressiformis (Native Cherry)

Callistemon pallidus (Bottlebrush) Grevillea sp. (Grevillea)

Callistemon viridiflorus (Green Bottlebrush) Melaleuca armillaris (Bracelet Honey Myrtle))

Correa nummularifolia (Native Fuchsia) Melaleuca squamea (Swamp Honey Myrtle)

Epacris languinosa (White Epacris) Oxylobium ellipticum (Golden Rosemary)

Eucalyptus ficifolia (Red Flowering Gum) Westringia sp. (Native Rosemary)

Introduced Species

Abelia schumannii (Abelia) Aquilegia sp. (Columbine, Granny's Bonnet)

Acanthus sp. (Bear's Breeches, Oyster Plant) Arbutus unedo (Irish Strawberry)

Acer psuedoplatanus (Sycamore) Arctotis sp. (African Daisy)

Acontium napellus (Monkshood) Asparagus officinalis (Asparagus)

Actinidia chinensis (Kiwi Fruit) Aubrieta sp. (Aubrieta)

Agapanthus orientalis (Agapanthus) Bellis sp. (Daisy)

Ajuga sp. (Ajuga) Beta vulgaris (Swiss Chard)

Alcea rosea (Hollyhock) Borago officinalis (Borage)

Allium schoenoprasum (Chives) Brassica oleraceae (Broccoli)

Alstroemeria sp. (Peruvian Lily) Brassica sp. (Tatsoi)

Antirrhinum majus (Snapdragon) Buddleia sp. (Buddleia)

Australian Entomologist 23 (2) September 1996

Introduced Species (cont.)

Cakile maritima (Sea Rocket)

Ceanothus sp. (Ceanothus)

Centaurea cyanus (Cornflower)

Centranthus ruber (Valerian, Kiss-me-quick)

Cerastium tomentosum (Snow-in-summer)

Chaenomeles sp. (Japonica)

Citrus aurantium (Orange)

Citrus limon (Lemon)

Citrus paradisi (Grapefruit)

Citrus reticulata (Mandarin)

Clematis sp. (Clematis)

Coleonema sp. (Golden Diosma)

Consolida sp. (Larkspur)

Convolvulus sp. (Convolvulus)

Conyza sp. (Erigeron)

Coprosma sp. (Coprosma)

Corylus avellana (Hazel)

Cosmos sp. (Cosmos)

Cotoneaster horizontalis (Cotoneaster)

Crocus sp. (Crocus)

Cucurbita sp. (Pumpkin)

Cytisus palmensis (Tree Lucerne)

Daboecia cantabrica alba (Irish Heath)

Dahlia sp. (Dahlia)

Delphinium sp. (Delphinium)

Dianthus sp. (Carnation)

Digitalis purpurea (Foxglove)

Dimorphotheca sp. (African Daisy)

Echium fastuosum (Pride of Maideira)

Erica sp. (Erica)

Escallonia sp. (Escallonia)

Fortunella sp. (Cumquat)

Freesia refracta (Freesia)

Fuchsia sp. (Fuchsia)

Fumaria muralis (Fumitory)

Gaillardia sp. (Blanket Flower)

Genus unknown (Broom)

Genus unknown (Cactus)

Genus unknown (Orchid)

Geranium sp. (Geranium)

Gladiolus sp. (Gladiolus)

Hebe sp. (Hebe)

Hedera helix (Ivy)

Hedychium sp. (Ginger Lily)

Helianthus annuus (Sunflower)

Hydrangea macrophylla (Hydrangea)

Hypericum perforatum (St Johns Wort)

Impatiens glandulifera (Balsam)

Jacaranda mimosifolia (Jacarada)

Jasmine sp. (Jasmine)

Kerria japonica (Kerria)

Laburnum sp. (Laburnum)

Lampranthus sp. (Pigface)

Lathyrus odoratus (Sweet Pea)

Lavandula alba (English Lavender)

Lavandula allardi (French Lavender)

Lavandula dentata (French Lavender)

Lavandula vera English Lavender)

Lavatera trimetris (Mallow)

Limonium sp. (Statice)

Lobelia sp. (Lobelia)

Lobularia maritana (Sweet Alice)

Lonicera japonica (Honeysuckle)

Lophomyrtus sp. (Lophomyrtus)

Lupinus sp. (Lupin)

Lycopersicon esculentum (Tomato)

Malus domestica (Apple)

Malus ioensis (Flowering Apple)

Matthiola sp. (Stock)

Meconopsis betonicifolia (Blue Poppy)

Melissa officinalis (Balm, Lemon Balm)

Mentha piperita var citrata (Mint)

Monarda sp. (Bergamot)

Myosotis sp. (Forget-me-not)

Myrtus communis (European Myrtle)

Myrtus ugni (Chilean Guava)

Nepeta cataria (Catmint)

Nothofagus sp. (Beech)

Origanum vulgare (Oregano)

Paeonia sp. (Peony Rose)

Papaver nudicaule (Icelandic Poppy)

Papaver orientale (Oriental Poppy)

Passiflora spp. (Passionfruit)

Pelargonium sp. (Pelargonium)

Penstemon sp. (Penstemon)

Petunia sp. (Petunia)

Phaseolus coccineus (Scarlet Runner Bean)

Philadelphus mexicanus (Mock Orange)

Phragmites karka (Bamboo)

Pittosporum sp. (Pittosporum)

Plantago lanceolata (Lamb Tongue, Plantain)

Polemonium caeruleum (Jacob's Ladder)

Polygonatum sp. (Solomon's Seal)

Protea aurea (Cream Protea)

Prunus persica (Peach)

Prunus sp. (Cherry)

Pseudofumaria alba (Corydalis)

Pyracantha sp. (Pyracantha)

Rhododendron spp. (Rhododendron, Azalea)

Ribes nigrum (Blackcurrant)

Ribes sanguineum (Flowering Currant)

Rosa spp. (Roses)

Rosmarinus officinalis (Rosemary)

Rubus fruticosus (Blackberry)

Rubus idaeus (Raspberry)

Rubus ursinus (Boysenberry)

Salvia officinalis (Sage)

Salvia sp. (Bog Sage)

Scabiosa columbaria (Butterfly Blue)

Schizanthus pinnatus (Poor Man's Orchid)

Senecio cruentus (Cineraria)

Solanum tuberosum (Potato - "Pinkeye")

Solidago canadensis (Goldenrod)

Symphyandra hoffmannii

Symphytum sp. (Comfrey)

Syringa sp. (Lilac)

Tagetes sp. (French Marigold)

Taraxacum officinale (Dandelion)

Tecoma sp. (Tecoma)

Tibouchina sp. (Lasiandra)

Australian Entomologist 23 (2) September 1996 35

Introduced Species (cont.)

Trifolium repens (White Clover) Vicia faba (Broad Bean)

Trifolium sp. (Clover) Viola sp. (Pansy)

Tropaeolium sp. (Nasturtium) Viola wittrockiana (Pansy)

Tulipa sp. (Tulip) Virgilea sp. (Virgilea)

Ulex europaes (Gorse) Watsonia sp. (Watsonia)

Veronica sp. (Veronica) Weigela sp. (Weigela)

Vicea sp. (Vetch) Wistaria sp. (Wisteria)

Acknowledgment

I thank Dennis Morris of the Tasmanian Herbarium for assistance in

identification of the plants.

References

DONOVAN, B.J. and MACFARLANE, R.P. 1984. Bees and Pollination. Pp 247-70, in: Scott,

R.R. (ed.), New Zealand Pests and Beneficial Insects. Caxton Press: Christchurch.

SEMMENS, T.D., TURNER, E. and BUTTERMORE, R. 1993. Bombus terrestris (L.)

(Hymenoptera: Apidae) now established in Tasmania. Journal of the Australian Entomological

Society 32: 346.

36 Australian Entomologist 23 (2) September 1996

NOTES ON THE FOODPLANT OF

DEUDORIX EPIRUS AGIMAR FRUHSTORFER (LEPIDOPTERA:

LYCAENIDAE)

1M. De BAAR and 2S.J. JOHNSON

1 Queensland Forest Research Institute, PO Box 631, Indooroopilly, Qld 4068

2Qonoonba Veterinary Laboratory, PO Box 1085, Townsville, Qld 4810

Abstract

This note records a name change and provides extra information for the foodplant of Deudorix

epirus agimar Fruhstorfer, documented by De Baar and Johnson (1980).

Discussion

The foodplant for this lycaenid was identified as Harpullia angustifolia

Radlk. (Sapindaceae) by the Queensland Herbarium and recorded as such by

De Baar and Johnson (1980). This herbarium sample was then reassigned to

Harpullia ramiflora Radlk (Reynolds 1981), previously only known from

New Guinea. Leenhouts and Vente (1982) have listed H. angustifolia as a

synonym of H. ramiflora and occurring in the Philippines, Moluccas

(Halmaheira), New Guinea and Iron Range (northern Queensland). The

name change for H. augustifolia is mentioned by Dunn and Dunn (1991) as

De Baar pers. comm. (1990) without further explanation.

No further foodplant records have been gathered for Deudorix epirus agimar

since De Baar and Johnson (1980), when two larvae were collected in

rainforest along the upper Claudie Riv. near Tozer's Gap, northern

Queensland. The fruit of H. ramiflora is eye catching as the capsules are red

and contain two seeds covered in yellow aril. The trees are a small straggling

species within the rainforest canopy.

References

DE BAAR, M. and JOHNSON, S.J. 1980. Rhopalocera hosts and notes from Iron Range trip

June, July 1978. News Bulletin of the Entomological Society of Queensland 8(4): 45.

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

and taxonomy, part 3: Family Lycaenidae. Privately published by authors, Melbourne. Pp 336-

512.

LEENHOUTS, P.W. and VENTE, M. 1982. A taxonomic revision of Harpullia (Sapindaceae).

Blumea 28(1): 1-51.

REYNOLDS, S.T. 1981. Notes on Sapindaceae in Australia, 1. Austrobaileya 1(4): 388-419.

Australian Entomologist 23 (2) September 1996 37

THE LIFE HISTORY OF

PHALACROGNATHUS MUELLERI (MACLEAY)

(COLEOPTERA: LUCANIDAE)

1G.A. WOOD, ?J. HASENPUSCH and ?R.I. STOREY

1 P.O. Box 122, Atherton, Qld 4883

2 P.O. Box 26, Innisfail Qld 4860

3 Dept. of Primary Industries, P.O. Box 1054, Mareeba, Qld 4880

Abstract

The life history of Phalacrognathus muelleri (Macleay) is described and aspects of its biology

discussed. The species is restricted to the wet tropics of northern Queensland where it breeds in

rotting wood in rainforest. Larvae have been extracted from the wood of 27 tree species in 13

families. All larvae found were in wood attacked by white rot fungi. The final instar larva is

described. Larva, pupa, and parasites are figured.

Introduction

Phalacrognathus muelleri (Macleay) (Coleoptera: Lucanidae) is a well

known but poorly documented species. It is variously known as the golden,

rainbow, magnificent, Mueller's and king stag beetle, and is the largest

Australian member of its family. It is illustrated as the centrepiece to colour

plates in both the major textbooks on Australian insects, viz. Tillyard, 1926

and CSIRO, 1970. Since 1973 it has been the official symbol of the

Entomological Society of Queensland. The history of the discovery of P.

muelleri is an interesting story and we have included it here in some detail.

The authors wish to thank Dr Geoff Monteith (Queensland Museum,

Brisbane) for providing the information in the following two paragraphs.

P. muelleri was initially described as a new species of Lamprima in 1885 by

Sir William Macleay in Sydney on the basis of a single female from “ North

Australia” sent to him from Victoria by Charles French Snr (Macleay,

1885a). French was later Government Entomologist in Victoria but at that

time he was assistant to the redoubtable Victorian Colonial Botanist, Baron

Ferdinand von Mueller. Macleay noted that French had asked him to name

the beetle after von Mueller, a request “. . . to which it gives me great

pleasure to comply . . .”. Macleay speculated that the species “may well

form the type of a new genus . . . but in the absence of a male specimen . . . it

would be premature . . .”. This prompted French, perhaps with a twinge of

conscience, to immediately send Macleay the male specimen he had held

back previously, and in the same year Macleay published a second paper

(Macleay 1885b) erecting the new genus Phalacrognathus. Macleay

described the male as “I think the most beautiful insect I have ever seen, not

surpassed in brilliancy of metallic lustre by the most gorgeous of the

Buprestidae”.

Macleay exhibited the female specimen at the Linnean Society meeting in

Sydney on 29 April 1885 where it “excited much attention” (Anon. 1885a)

and he exhibited the male at the same venue on 30 September (Anon.

1885b). The origin of those specimens which French sent Macleay remains a

38 Australian Entomologist 23 (2) September 1996

mystery. At that general time von Mueller was employing botanical

collectors to work in north Queensland. One of these was William Sayer,

who was also a cousin of Charles French. Another collector in north

Queensland at that period was Walter Froggatt who, according to Musgrave

(1932), sent insect specimens to both French and von Mueller. It is very

likely that French’s Phalacrognathus came from either Sayer or Froggatt.

Contemporary entomologists were also curious about the origin of those

exciting specimens. At the meeting of the Linnean Society of New South

Wales in 1892 Mr A.S. Olliff enquired”. . . as to the exact habitat of the

splendid lucanid beetle . . . described by Sir William Macleay . . . about

which the only information then available was that they came from North

Australia?" Mr F.A. Skuse, then entomologist at the Australian Museum,

replied that specimens had recently been received by the Museum from

*Russell Scrub, Boar Pocket, near Cairns, Queensland" ( Anon. 1892). This

locality was a popular staging post at that time on the newly opened pack

trail from Cairns to the Herberton tinfields via the Mulgrave River and

presumably this is how the collector of those specimens would have accessed

the area. Much is now submerged under the Tinaroo Dam at the north end of

the Atherton Tableland (Toohey, 1994). The species was redescribed under

the synonymic name of Phalacrognathus westwoodi by Shipp (1893) from a

single male in the Oxford University Museum. This specimen had been

purchased by Professor Westwood from the dealer Boucard in 1889 and

came from the same uninformative locality as had French's specimens:

“North Australia".

The few papers published on the biology of this species are largely the result

of casual observation (Brooks 1970, Hancock 1970, Dodd 1971, Brunet

1981, Quick 1984). This paper provides detailed information on the biology

and natural history of this the most beautiful of Australian beetles.

Observations were made over a period of 14 years on a large number of

specimens of all stages, taken in the wild or bred in captivity.

Distribution and Abundance

Phalacrognathus muelleri is confined to the rainforests and adjacent wet

sclerophyll forests of coastal north-eastern Queensland between Helenvale

(15°43'S 145°14'E) near Cooktown and Pine Creek (19°15'S 146?28'E) at the

southern end of the Paluma Range. It is referred to variously as very rare

(Britton 1970) or extremely rare (Hawkeswood 1987). As pointed out by

Wood and Hasenpusch (1990), this conclusion is likely a result of the species

only occasionally coming to light, the most frequently used method of insect

survey and collection in rainforests. Our study found P. muelleri to be

common in virtually any rainforest block within its known distribution,

although seemingly more abundant on the ranges and tablelands than at sea

level. As the total area of rainforest within its distribution is an estimated

Australian Entomologist 23 (2) September 1996

Figs 1-8. Third instar larva of Phalacrognathus muelleri. (1). Head capsule

(mandibles removed); (2). epipharynx, ventral view; (3). left mandible, dorsal view;

(4). antenna, inner surface; (5).apex of hind leg; (6). pars stridens on mesocoxa;

(7).plectrum on metatrochanter; (8). raster on ventral side of abdominal sternite 10 .

(Scale lines = 1 mm ).

40 Australian Entomologist 23 (2) September 1996

7910 km? (Bell et al 1987), P. muelleri can certainly not be described as

rare.

Breeding Sites

Phalacrognathus muelleri breeds in rotting wood in both fallen and standing

living or dead trees. Larvae may be found in suitable decayed wood with a

minimum volume of approximately 10 litres (0.01 cubic metres). Although

P. muelleri larvae prefer moderately moist conditions they have also been

found in both dry and saturated substrates; waste wood left by logging

operations is commonly used. Waste wood tends to be available for a

relatively short time when compared to unlogged forests where food supply

is continuous. In standing trees where there is an abundance of substrate,

such as dead portions of a large, living tree, some individuals may complete

their life-cycle without leaving the confines of the trunk. Adults feed on the

same material as the larvae and supplement their diet with plant sap and

fruits. Trees up to 1.5 m in diameter and containing up to 35 cubic metres of

wood have been found as suitable. An indicator of this may be a fallen

branch riddled with larvae. Such trees can support generations of beetles.

Life History

Oviposition and Egg

Up to 50 eggs are laid by each female, either in wood pulp produced by the

female as she excavates a tunnel with her mandibles and tibial spines or in

pulp produced by older larvae. Substrate the consistency of balsa is suitable

for oviposition. Although each egg is deposited singly, females have been

observed to lay up to 30 eggs in close proximity. Captive females laid

readily in whole pieces of substrate of suitable size but did not oviposit in

artificially produced pulp from the same timber. Newly laid eggs are 2.9-3.7

mm long and 2.2-2.8 mm wide. The surface is smooth with distinct, fine,

hexagonal microsculpture, and they are slightly longitudinally flattened in

shape. Eggs take 10-14 days to hatch, each expanding to become almost

spherical and approximately doubling in volume as they mature. Colour

changes from translucent white to dark cream. The larva is visible within the

egg just before emergence. The egg splits lengthwise on hatching but a seam

or ridge is not visible beforehand.

Larval Description and Behaviour

Of the 18 genera of Lucanidae currently recognised in the Australian fauna

(Moore and Cassis 1992), the larval stage has been described for very few.

Alderson (1975a, 1975b) compared and partially described species of

Lamprima Latreille, Lissapterus Deyrolle, Lissotes Westwood, Syndesus

W.S. Macleay and Ceratognathus Westwood. Lawrence (1981) discussed

larval Lucanidae in general and although he examined larvae of ten

Australian genera (including Phalacrognathus), he did not include formal

descriptions but did provide a partial larval key to genera and higher

Australian Entomologist 23 (2) September 1996 41

Figs 9-11. (9). Parasites and ? hyperparasite of P. muelleri larvae (from left to right):

Amphibolia ignorata Paramonov (Tachinidae); Liacos insularis (Smith) (Scoliidae);

Mordella elongatula (Macleay) (Mordellidae); (10). Larva of Liacos insularis (Smith)

attached to final instar P. muelleri larva; (11). Pupa of male P. muelleri showing

lateral spines on apical abdominal segment.

42 Australian Entomologist 23 (2) September 1996

categories. It is beyond the scope of this paper to discuss the larval

relationships of P. muelleri. However, the following description of the final

instar larva and illustrations of some characters used in the classification of

lucanid larvae are provided:

Head (Fig. 1). Maximum width 7.2 - 12.0 mm. Colour light brown with

posterior 2/3 of clypeus, antennae and labrum darker brown, anterior frontal

angles and mandibles black. Frontal setae (pairs): 1 anterior angle, 1

anterior, 1 exterior, 2 posterior. Clypeus slightly convex with lateral seta and

a short lateral tubercule. Labrum obtusely pointed apically. Left mandible

(Fig. 3) with 3 apical teeth, right mandible with two, both mandibles with a

shallow groove on outer side from base for about half the length. On the

epipharynx (Fig. 2), haptomerum with 5 sensilla in a straight line,

chaetoparia with 6-11 scattered setae towards outer margin, phobia

semicircular, irregular, mostly made up of short blunt spines, epitorma

narrow, about 1/3 length of phobia, pternotormae short, obtuse, two well

sclerotized lateral and one medial nesia in the haplolachus. Oncylus of

hypopharynx strong, triangular, apex acute. Antennae (Fig. 4) 4-segmented,

first segment short almost completely fused to the second long, glabrous

segment, third segment shorter with inner, irregular lateral and oval,

subapical sensory patches, fourth segment short with two setae midway

along inner margin, and one seta on subacute apex.

Thorax . With a pair of dorsolateral sclerotized plates just behind head and a

pair of ventral sclerotized plates in front of procoxa, meso- and metapleural

lobes sclerotized above legs. Tibiotarsus, femur, and trochanter of each leg

with ventral and lateral surfaces strongly granulate at base of each seta.

Tarsungulus (Fig. 5) short, blunt, with an apical and a subapical seta. Pars

stridens (Fig. 6) an area of confused microgranules, with a row of about 50-

60 distinct larger granules along outer edge, increasing in size towards apical

end. Plectrum (Fig. 7) consists of confused granules at apical end, grading to

a single series of about 30 granular rows finally becoming oval, striated

plates nearest coxa.

Spiracles C-shaped, thoracic largest, abdominal spiracles 4 and 5 smallest.

Abdomen. Dorsal surface of all abdominal segments with fields of short to

long posteriorly directed setae, setae longest near the posterior margin of

each segment. Raster (Fig. 8) present on ventral surface of segment 10 in the

form of a V-shaped band of spinules, interrupted at apex. Upper anal lobe

transverse, triangular, bare, crescent shaped on outside of lateral anal lobes.

Larvae live singly, but often in close proximity, within the decaying wood

and may travel several metres through it as they develop. Excavated pulp is

packed behind as the larva moves forward. Pieces of substrate are isolated

by the mandibles and then grasped by the mandibles and legs and passed

backwards. This action is achieved by arching the body upwards in the

Australian Entomologist 23 (2) September 1996 43

Table 1. Host Tree Species for P. muelleri larvae

Samples

Apocynaceae Alstonia scholaris (L.) R. Br. v

Cunoniaceae Ceratopetalum succirubrum C.T. White 12

Cunoniaceae Caldcluvia australiensis (Schltr.) Hoogl 7

Elaeocarpaceae | Sloanea australiensis (Beth.) F. Muell. 2

Elaeocarpaceae |Sloanea sp. 2

Flindersiaceae — | Flindersia pimenteliana F. Muell. 3

Flindersiaceae — | Flindersia pubescens F.M. Bail. 1

Lauraceae Beilschmiedia bancroftii (F.M. Bail.) C.T. White 4

Lauraceae Beilschmiedia spp. 3

Lauraceae Cryptocarya mackinnoniana F. Muell. 1

Lauraceae Endiandra monothyra B. Hyland ssp. monothyra 1

Meliaceae Dysoxylum sp. 1

Meliaceae Synoum muelleri C. DC. *

Mimosaceae Acacia melanoxylon R. Br. 1

Monimiaceae Daphnandra repandula F. Muell. 1

Monimiaceae Doryphora aromatica (F.M. Bail.) L.S. Smith 1

Moraceae Ficus - 2 spp. *

Moraceae Ficus sp. 1

Myrtaceae Eucalyptus torelliana F. Muell. *

Myrtaceae Xanthostemon whitei Gugerli 1

Proteaceae Cardwellia sublimis F. Muell. 5

Proteaceae 3 Proteaceae spp. 3

Rutaceae Halfordia kendack (Montr.) Guill. 1

Sapotaceae Palaquium (Lucuma) galactoxyla (F. Muell.) H.J. Lam M

* Hancock (1970.)

Table 2. Fungi associated with P. muelleri larval development

No. of Samples

Ganodermataceae | Ganoderma applanatum (Pers.) Patouillard [|4

Polyporaceae Nigrofomes melanoporus (Mont.) Murr. 1

Polyporaceae Phellinus nr. glaucescens (Petch) Ryvarden. |10

Polyporaceae Phellinus robustus (P.Karst) Baird. & Galz. |1

Polyporaceae Phellinus - 3 spp. 24

Polyporaceae Pycnoporus sp. 2

centre, which draws the head under the body. Further movement is achieved

by peristaltic movement in the folds of the surface of the thoracic segments.

As the larva straightens, material is pressed into the rear wall of the cell.

These arching and pressing movements are also used in the production of

44 Australian Entomologist 23 (2) September 1996

oval ecdysal cells before moults. These cells are lined with faeces. Arching

into a "C" shape the larva deposits a faecal pellet between the legs. The

pellet is broken up with the aid of the mandibles and the larva then uses the

flat anterior surface of the head to compress the faecal material on to the cell

wall. Each ecdysal cell takes about 24 hours to construct. A few hours after

completing the cell the larva moults, eating the cast skin before breaking out

of the cell several hours later.

Each larva passes through three instars. From egg to adult takes a minimum

of 12 months, with the larval stage a minimum of 11 months. Hancock

(1970) reported the pupal stage lasting 4-5 weeks and observed length of

time in this study was 3-5 weeks. Larvae that develop in 12 months produce

small adults. Larger specimens take 2 to 3 years to develop. Specimens in

poor quality substrate may take 4 years to develop and are usually small.

Larval Host Plants

Like most members of the family Lucanidae, larvae of P. muelleri feed on

decaying wood of various species of trees. Authors names of tree species

mentioned here are given in Table 1. Two of the earliest records (Hancock

1970, Britton, 1970) of a suitable species were red cedar (Toona australis).

Monteith (pers. comm.) felt that both these records were based on the

opinion of the late George Brooks, an avid beetle collector based many years

in Cairns, that a ‘red-timbered log’ from which he had taken P. muelleri

specimens on several occasions was this species. Brooks (1970), about the

same time, however, obtained an identification for the log in question which

turned out to be Synoum muelleri C.DC. (Meliaceae). The record of Synoum

muelleri cited by Brooks (1970), repeated by Hancock (1970), was later

suggested to be a misidentification (J.G. Brooks, pers. comm. to D.L.

Hancock) and requires confirmation. Other host records cited by Hancock

(1970), apart from Toona australis, were supplied by R. Parrott (D.L.

Hancock, pers. comm). Toona did not produce any specimens of P. muelleri

in the current study, so should be removed from the host list. Hancock

(1970) recorded four other species as hosts, namely Palaquium (as Lucuma)

galactoxyla, Cerbera manghas, and Ficus spp. (two species). It is likely that

Cerbera manghas (Milky Pine) is in reality Alstonia scholaris as the former

is primarily a sea level species, the latter being found on the Atherton

Tablelands proper (Monteith, pers. comm.). Table 1 gives the tree species

recorded in this study. The records of Hancock (1970) are also included

except for the removal of Toona and the substitution of Alstonia for Cerbera.

A large percentage of logs containing P. muelleri larvae have decayed to a

point where species identification is not possible. Undoubtedly other tree

species are also suitable. Timber samples from logs infested by P. muelleri

larvae were identified by officers of the Queensland Forest Service. Leaf

specimens from living trees were identified by Bernie Hyland of CSIRO,

Atherton (5 species).

Australian Entomologist 23 (2) September 1996 45

Host trees need to be infected with fungi to be suitable for P. muelleri larvae.

It would appear that the tree species is less important than that suitable white

rot must be present in the wood. In general, white rot is indicated by a pale

discolouration of the wood, a result of the darker coloured lignin being

removed. Colour ranges from white to pale brown and may include dark

zone lines and "pockets". White rot tends to have a "stringy" texture. White

rot chemistry is not uniform and we found that not all white rot wood is

suitable substrate for P. muelleri. Fungal determinations are also rendered

progressively more difficult as the substrate deteriorates. Table 2 gives

fungal species recorded in this study as associated with P. muelleri.

Pupation

The pupal cell is constructed in a similar manner to the other ecdysal cells

but takes up to a week to complete. In addition to the lining of faecal matter

the cell walls are coated with an amber fluid. The anal segment of the

abdomen is used as a "brush" in this process. The source and purpose of this

coating is unknown.

The pupa has previously been illustrated. Hancock (1970) gives a line

drawing and Brunet (1983) gives colour photographs of a pupa and newly

emerged adult.

Hancock (1970) reported moult to pupa occurring approximately two weeks

after completion of the pupal cell. The pupa can rotate in the pupal cell on

its longitudinal axis using of a pair of lateral spines on the apical abdominal

segment (Fig. 11). These spines grip the cell wall and the pupa rotates as it

arches and twists sideways. The pupa changes position in the cell many

times during development. As the legs begin to harden prior to adult

emergence, the pupa rolls on to its back. When the legs become rigid the

pupal skin around them is transparent. After then rolling back on to its legs

the pupal skin is shed. This commences with the pupal skin splitting down

the middle of the dorsal surface of the pronotum. The adult may remain

within the pupal cell for up to 8 months before emergence.

Adult Morphology and Behaviour

Males vary in length (measured from tips of mandibles to elytral apex) from

24-72 mm and females from 23-46 mm. As an expression of percentage of

total length, male mandibles vary between 19 -32%, irrespective of beetle

size. Two males have been recorded with asymmetric mandibles. No

variation in mandibular development is apparent in females. A small

percentage of non-teneral adults display a shift in colour to either increased

red or green. Very rarely is either colour absent but several dark blue

specimens have been recorded. Dodd (1971) also recorded rare examples

that were very dark in colour.

46 Australian Entomologist 23 (2) September 1996

In breeding large numbers of specimens of this species in this study, females

were found to outnumber males by approximately 4%. Dodd (1971),

however, stated that in the field males greatly outnumber females.

Adults break out of their pupal cells using the mandibles and legs. Males

with well developed mandibles use the base of these for chewing, raking

excavated material out with the tarsi. Upon emergence from the pupal cells

adults disperse in search of food, mates and oviposition sites. Adults are

known to live for up to 18 months in captivity. Light trapping records

indicate peak activity at and just after dusk from September to April.

Adult feeding has been observed on Eucalyptus sp. blossom, on the fruit of

Calamus moti Bailey (Arecaceae) (M. Walford-Huggins pers. comm.) and on

sap flows associated with insect damage (E. Adams pers. comm.). Dodd

(1971) noted that adults were taken on Glochidion ferdinandi (J. Muell.)

F.M. Bail. (Euphorbiaceae) where hepialid or other wood borers had left a

sap-exuding injury. Another apparently less favoured food plant is

Caesalpinia sp. (Caesalpiniceae), a robust thorny leguminous creeper; in this

case the beetle occurring on terminal shoots (Dodd 1971). All records except

the first involved feeding during the day.

Oviposition occurs throughout the year in captivity and in the wild. Males

have been recorded in the company of ovipositing females and mating has

been observed on numerous-occasions in captivity. Mating, together with

conflict between males, also has been observed at feeding aggregations in

captivity.

Adults being handled rarely fold in their legs and remain rigid; they usually

claw with the tarsi and may attempt to bite. Males may adopt a threatening

stance by rising up on the mid and hind pairs of legs, with forelegs

outstretched and mandibles working like scissors.

Males use their mandibles as levers when in conflict with one another. Two

protagonists approach each other with the mandibles lowered. Each beetle

tries to pass beneath its opponent’s body or legs, at which point the

mandibles are raised in an attempt to dislodge the rival. On vertical surfaces

combatants may be thrown into the air. On horizontal surfaces one may be

rolled over and in the grappling that ensues tarsi are sometimes bitten off.

Males also have been observed using their mandibles to facilitate mating. If

the female is in a position where she cannot be successfully approached, the

male may use his mandibles to lever her into a better position. Where escape

or concealment is attempted the male may use his mandibles to violently

throw her about or carry her elsewhere.

Relations with other beetles and parasites.

Other lucanid species found in the same logs in close proximity to P.

muelleri include Lamprima latreillei W.S. Macleay, Rhyssonotus nebulosus

Australian Entomologist 23 (2) September 1996 47

(Kirby), Prosopocoilus torresensis (Deyrolle), Figulus sp., Aegus jansoni

Boileau, Syndesus cornutus (Fabricius), and Cacostomus squamosus

Newman. The first three species appear to be white rot associated but tend to

utilise different niches. L. latreillei prefers drier sites and is more common

in dead standing timber than in logs on the ground. R. nebulosus tends to be

most common in sapwood and P. torresensis is less common with increasing

elevation. Although they can be found in the same timber as P. muelleri, A.

jansoni and S. cornutus are brown rot associated species. Brown rot is the

result of fungal attack in which the cellulose has been destroyed leaving the

dark coloured lignin. This is evidenced by zones of darker colour in the

wood, dark brown to rust-red in colour. Brown rot does not have a "stringy"

texture but tends to be powdery. C. squamosus is found in detritus in cracks

in wood and under logs and is not clearly associated with either type of rot.

Larvae of the cetoniine Schizorhina atropunctata (Kirby) are commonly

found with white rot-feeding lucanid larvae, feeding on their faeces and

fragmented wood. Larvae of another cetoniine, Lenosoma sp. also have been

found feeding on P. muelleri larval frass.

Larvae of P. muelleri are parasitised commonly by the fly Amphibolia

ignorata Paramonov (Tachinidae) (Fig. 9). They are also parasitised by the

wasp Liacos insularis (Smith) (Scoliidae) (Fig. 9): larvae attach to final-

instar P. muelleri larvae and pupae (Fig. 10), the egg being laid on the

underside, just behind the legs. L. insularis was only observed on the edge

of rainforest. The beetle Mordella elongatula (Macleay) (Mordellidae) (Fig.

9), one specimen of which emerged from a pupal cell of L. insularis, is

possibly a hyperparasite but this habit is not usual for mordellid beetles.

Acknowledgments

The authors thank Geoff Monteith and Bryan Cantrell for identifying

parasites of P. muelleri, Victoria Gordon, Ian Hood, John Lawrence, Alan

Mills and M. Ramsden for identifying fungal samples and Bernie Hyland for

determination of leaf samples. Geoff Monteith is especially thanked for

detailed notes on taxonomic history of Phalacrognathus and on larval hosts.

Thanks are also extended to David Hancock for discussion on larval host

trees. Timber samples were identified by Garry Hopewell and Graham

Hughes. Particular gratitude is extended to Graham who determined the bulk

of the samples. We also thank the Conservator, Queensland Forest Service

for permission to collect in State Forests.

References

ALDERSON, J. 1975a. Descriptions of the larvae of four species of Lucanidae (stag beetle)

Victorian Naturalist 92: 71-79.

ALDERSON, J. 1975b. Description of the larva of Ceratognathus niger (Westw.) Coleoptera:

Lucanidae (stag beetle). Victorian Naturalist. 92: 217-221.

48 Australian Entomologist 23 (2) September 1996

ANON. 1885a. Notes and exhibits. Proceedings of the Linnean Society of New South Wales

10: 187.

ANON. 1885b. Notes and exhibits. Proceedings of the Linnean Society of New South Wales

10: 553.

ANON. 1892. Notes and exhibits. Proceedings of the Linnean Society of New South Wales

17: 20.

BELL, F.C., WINTER, J.W., PAHL, L.I. & ATHERTON, R.G. 1987. Distribution, area and

tenure of rainforest in northeastern Australia. Proceedings of the Royal Society of Queensland

98:27-39.

BRITTON, E.B. 1970. Coleoptera. Pp 495-621 in CSIRO The Insects of Australia. Melbourne

University Press, Melbourne.

BROOKS, J.G. 1970. The old "red-timbered" log. News Bulletin of the Australian

Entomological Society 6: 38.

BRUNET, B.L. 1981. Some observations of the north Queensland stag beetle. Lucanidae.

North Queensland Naturalist. 45 (178): 12-13.

BRUNET, B.L. 1983. One Step Closer Please. 95 pp. View Productions Pty Ltd, Sydney.

CSIRO 1970. The Insects of Australia. 1029 pp Melbourne University Press, Melbourne.

DODD, A.P. 1971. The stag beetle, Phalacrognathus muelleri. News Bulletin of the

Entomological Society of Queensland. 74: 12.

HANCOCK, D.L. 1970. Rearing Phalacrognathus muelleri, Australia's finest stag beetle.

News Bulletin of the Entomological Society of Queensland. 72: 15-16.

HAWKESWOOD, T.J. 1987. Beetles of Australia. 248 pp. North Ryde, N.S.W. Angus and

Robertson.

LAWRENCE, J.F. 1981. Notes on Larval Lucanidae (Coleoptera). Journal of the Australian

Entomological Society 20: 213-219.

MACLEAY, W.J. 1885a. Revision of the genus Lamprima of Latreille, with descriptions of

new species. Proceedings of the Linnean Society of New South Wales 10: 129-140.

MACLEAY, W.J. 1885b. A new genus of the subfamily Lamprimides of Lacordaire.

Proceedings of the Linnean Society of New South Wales 10: 473-474.

MOORE, B.P. and CASSIS, G. 1992. Lucanidae. Pp. 4-19 in Houston, W.W.K. (ed)

Zoological Catalogue of Australia. Coleoptera: Scarabaeoidea. Canberra: AGPS Vol. 9, pp.

xii + 544.

MUSGRAVE, A. 1932. Bibliography of Australian Entomology 1775-1930 With Biographical

Notes on Authors and Collectors. 380 pp. Royal Zoological Society of New South Wales,

Sydney.

QUICK, W.N.B. 1984. Notes on the longevity of Phalacrognathus muelleri Macleay

(Coleoptera: Lucanidae, Lampriminae). Victorian Entomologist. 14: 23-25.

SHIPP, J.W. 1893. On a new species of the genus Phalacrognathus, MacLeay. Transactions

of the Entomological Society of London 1893: 223-225.

TILLYARD, RJ. 1926. The Insects of Australia and New Zealand. Angus & Robertson,

Sydney.

TOOHEY, E. 1994. Kie Daudai. Notes and Sketches from Cape York. 324 pp. Privately

Published.

WOOD, G.A. and HASENPUSCH, J. 1990. A simple method of elevating artificial light

sources to attract insects. Queensland Naturalist. 30:36-37.

Australian Entomologist 23 (2) September 1996 49

THE LIFE HISTORY OF THE WESTERN AUSTRALIAN

SKIPPER MESODINA CYANOPHRACTA LOWER

(LEPIDOPTERA: HESPERIIDAE)

Andrew A.E. Williams! and Andrew F. Atkins?

! Department of Conservation and Land Management, W.A. Wildlife Research Centre,

P.O. Box 51, Wanneroo, W.A. 6065

? Design Department, The University of Newcastle, Callaghan, N.S.W. 2308

Abstract

The life history of Mesodina cyanophracta Lower is described and illustrated. The early

stages of M. cyanophracta and M. halyzia (Hewitson) are compared.

Introduction

Mesodina cyanophracta Lower (Figs 7, 8) was recognised only recently as

distinct from M. halyzia (Hewitson) (Edwards 1987). M. cyanophracta is

restricted to south-western Australia where it occurs from 36 km west of

Binnu near Geraldton, south to Albany and inland to the Stirling Range;

adults have been taken from late October to March (Common and Waterhouse

1981, Dunn and Dunn 1991). We have recent records from Condingup Peak,

65km east of Esperance (33?45'41"S 122?32'53"E), and from Mount Ragged

in Cape Arid National Park (33?28'02"S 123?27'31"E). Around Perth the

peak flying time is in November and the foodplant is Patersonia occidentalis

R.Br. (Iridaceae) (Williams et al. 1993). The larval and pupal stages of M.

cyanophracta have not been described in detail, although Common and

Waterhouse (1981) and Edwards (1987) stated that the early stages are similar

to M. halyzia.

Life history

Egg (Figs 1, 9, 10). Diam. 1.75 mm, hemispherical, off white changing to

pale green with maroon dorsal blotch, acentric to micropyle. Surface covered

with a delicate lace-like ribbed structure.

First instar larva (Fig. 2). Length 3.5-4.0 mm. Head shiny black, surface

faintly pitted and covered with variable pale setae; prothoracic plate shiny

black; collar between head and prothoracic plate bright orange-red. Body

tapered, yellowish, last segment pinkish; covered with variable white setae,

some slightly clubbed; posterior setae long and slender.

Mature larva (Fig. 3). Length 20-26 mm. Head (Fig. 4) large and rounded,

greyish black in colour, surface granulated and covered with variable pale

setae. Body greyish-brown with short clubbed setae, posterior with long

whitish setae. A narrow blackish dorsal line extends almost the length of the

body. Medium-sized to mature larvae are covered with a white waxy powder

which tends to obscure the skin coloration.

Pupa (Fig. 6). Length 18-22 mm, broad anterior tapering to posterior with

small cremaster. Frons (Fig. 5) more or less smooth with small elliptical

operculum pointed laterally. Colour somewhat variable; fresh pupae with

50 Australian Entomologist 23 (2) September 1996

thorax and wing cases dull green, abdomen cream to yellow cream and frons

grey-brown. As development progresses the pupae darken and some

individuals turn almost black.

Observations and Discussion

Eggs were found at Moore River National Park, 100 km north of Perth, in

mid November 1993 and transferred to P. occidentalis plants suitably located

for observation near Wanneroo. Larvae hatched after several.days and

consumed their egg casings before constructing shelters. Most individuals

made shelters by drawing together the outer edges of a single leaf of the

foodplant to form a partial tube, which was then sealed at the top. Others

sewed together the extreme tips of two leaves to form a shelter. Early stage

larvae invariably fed on the upper edges of the leaves in close proximity to

their shelters.

As larvae increased in size they abandoned their first shelters and constructed

larger ones by sewing together three or four leaves of the foodplant to form a

tent-like structure. These shelters were closed at the top and lined with silk

with the entrance underneath. Larvae rested head downwards within the

shelters and fed during the day, as do larvae of M. halyzia (Common and

Waterhouse 1981). They produced distinctive feeding scars by cutting wedge-

shaped sections out of the leaves; in some cases this resulted in leaves taking

on a saw-toothed appearance. Larvae fed actively in November, December and

again from May to October. They remained largely inactive during the hot

dry summer months. Larvae pupated head downwards within their shelters.

Before pupating they sealed the entrance with a horizontal pad of silk. Pupal

duration was approximately 35 days.

Differences were found between the pupae of M. cyanophracta and M. halyzia.

The most distinctive feature of M. cyanophracta is the comparatively small

elliptical pupal cap (operculum), which is pointed laterally, unlike that of M.

halyzia which is broadly elliptical, rounded laterally and with a more

roughened sclerotized area on the upper frons. Mature larvae of M.

cyanophracta also differ in colour from those of M. halyzia which are pale

greenish in colour (Common and Waterhouse 1981). Scanning electron

micrographs of the eggs of M. cyanophracta and M. halyzia (Figs 9-12) show

subtle differences in their delicate lace-like ribbing. In M. cyanophracta the

cross ribs are more prominent and the micropyle pattern more diffuse and

complex than in M. halyzia. The delicate ribbed structure of Mesodina eggs

is unique in the Trapezitinae (AFA pers. obs.).

Observations over several years at Wanneroo indicate that female M.

cyanophracta prefer to oviposit on young vigorously growing foodplants.

They will often favour sites which have recently regenerated after fire, or

select young P. occidentalis plants growing alongside tracks or firebreaks.

Similar behaviour has been recorded for Trapezites sciron sciron Waterhouse

& Lyell near Perth, where ovipositing females showed a distinct preference

Australian Entomologist 23 (2) September 1996 51

Figs 1-8. Juvenile and adults of Mesodina cyanophracta Lower from Wanneroo,

W.A. (1) egg; (2) Ist instar larva; (3) final instar larva; (4) final instar larval

head; (5) frons of pupa and operculum; (6) pupa; (7) adult male, upperside and

underside; (8) adult female, upperside and underside. Scale bars: (1, 2) = 1 mm, (3-

6) = 5 mm, (7, 8) = 10 mm.

52 Australian Entomologist 23 (2) September 1996

Figs 9-10. Scanning electron micrographs (SEM) of egg of Mesodina

cyanophracta Lower from Wanneroo, W.A. (9) showing delicate lace-like ribbed

structure, (10) detail of micropyle pattern. Scale bar = 100 m.

Australian Entomologist 23 (2) September 1996 53

Figs 11-12. Scanning electron micrographs (SEM) of egg of Mesodina halyzia

(Hewitson) from Catherine Hill Bay, N.S.W. (11) showing delicate lace-like

ribbed structure, (12) detail of micropyle pattern. Scale bar = 100 m.

54 Australian Entomologist 23 (2) September 1996

for young vigorously growing Lomandra caespitosa (Benth.) foodplants

(Williams et al. 1992). Larvae of M. cyanophracta are frequently attacked by

a slender endoparasitic wasp, Casinaria sp. nr. meridionalis (Turner)

(Ichneumonidae), that eventually kills them in the final instar (AFA

identification - see Gauld 1984).

Voucher specimens pertinent to this paper are lodged in the Insect Collection

of the Western Australian Department of Conservation and Land Management

and in Andrew Atkins' private collection.

Acknowledgment

We thank Mr Garry Weber of The University of Newcastle for the scanning

electron micrographs.

References

COMMON, LF.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Pp. xiv + 682.

Angus and Robertson, Sydney.

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

and taxonomy. Part 2: Family Hesperiidae. Privately published, Melbourne.

EDWARDS, E.D. 1987. A new species of Mesodina Meyrick from the Northern Territory

(Lepidoptera: Hesperiidae). Australian Entomological Magazine 14: 4-12.

GAULD, I.D. 1984. An introduction to the Ichneumonidae of Australia. British Museum

(Natural History), London.

WILLIAMS, M.R., WILLIAMS, A.A.E. and ATKINS, A.F. 1992. The life history of the

Sciron Skipper Trapezites sciron sciron Waterhouse and Lyell (Lepidoptera: Hesperiidae:

Trapezitinae). Australian Entomological Magazine 19: 29-32.

WILLIAMS, A.A.E., WILLIAMS, M.R., HAY, R.W. and TOMLINSON, A.G. 1993. Some

distributional records and natural history notes on butterflies from Western Australia.

Victorian Entomologist 23: 126-131.

Australian Entomologist 23 (2) September 1996 55

SEASONALITY OF CICADAS (HEMIPTERA) ON THE

NORTHERN TABLELANDS OF NEW SOUTH WALES

Marc Coombs

Department of Zoology, University of New England, Armidale, N.S.W. 235]

Present address: CSIRO Division of Entomology, PMB 3, Indooroopilly, Qld 4068

Abstract . ,

Seasonality records are provided for 16 cicada species at sites near Armidale NSW, observed

over 4 summers (1990-91 to 1993-94). Cicadas were active at these sites from late October to

late February. The greatest number of species was present during December and early

January. There was a significant, positive correlation of species number with day length in all

years. Neither temperature nor rainfall consistently correlated with species number.

Introduction

A conspicuous feature of cicada biology is their predictable occurrence each

year during summer. Though seasonality records are available for Australian

species over their geographic range (Moulds 1990), there are no studies

documenting seasonal patterns of occurrence on a regional basis. Regional

differences in seasonality would be expected to occur as a consequence of

regional climatic differences. This study was undertaken to document the

seasonal activity of cicada species at sites near Armidale on the Northern

Tablelands of New South Wales. Armidale experiences relatively mild

summers (January average daily maximum, 27.1?C), and winters are cool to

cold (July average daily maximum 12.4°C). The frost free period extends

from November to March (30 y av.) (Bureau of Meteorology, unpubl. data).

The Armidale climate is broadly representative of the New England

Tablelands, a region extending from the township of Bendemeer (70 km

south-west of Armidale) to Dorrigo (110 km east) and north to the

Queensland border (Wallangarra).

Methods

The following localities were used as sampling sites in this study: The

University of New England campus (30?29'S, 151?39'E), the 'Pinnacle'

(30°30'S, 151?30'E), Dangars Falls (30°41'S, 151?44'E) and 'The Devils

Pinch' (30?20'S, 151°40'E). All sites are within 20 km of Armidale and vary

from ca 1000 to 1200 m above sea level. Each was visited during the second

and fourth weeks of each month from August to April of the summers 1990-

91, 1991-92, 1992-93 and 1993-94. The cicada species present on each

sampling date were identified by sight or by song pattern. Cicada songs are

species specific (Young 1972, Ewart 1989) and can be readily learnt for a

given locality. To assess the influence of climate on seasonal activity,

fortnightly averages of climatic and regional variables (max. and min.

temperature, rainfall and photoperiod) were obtained for Armidale for the

study periods. Data from the four sampling localities were pooled and

analysed (Pearson product-moment correlation) against climatic and regional

variables for each year of the study.

56 Australian Entomologist 23 (2) September 1996

Table 1. Seasonality records for cicada species at Armidale, New South Wales,

during the period October to February of the years 1990/91 to 1993/94. Each

month is scored twice to represent sampling in the second and fourth week.

Species Year Month

OR ORB NS Nie De ae) ae) gee Fee

Cicadetta 90/91 uo 4E we x) 0

waterhousei 91/92 * S * >

(Distant) 92/93 wR n

93/94 * * * * * *

Cicadetta 90/91 V VU k * wi

labeculata 91/92 wo * * x

(Distant) 92/93 vov 3 D €

93/94 c RE m o £

Cicadetta 90/91 wo WR Rd

tristrigata 91/92 ME Lf

(Goding & 92/93 o RB GE 1

Froggatt) `- 93/94 Meter Voip fua

Cicadetta 90/91

landsboroughi 91/92 Apto. er UR

(Distant) 92/93 tio ERO M UE

93/94 uo eee d

Cicadetta puer 90/91 o Ch RR GRE CEP XS S

(Walker) 91/92 e M EP RE D € €

92/93 * * * * * * *

93/94 * * * * * * * *

Urabunana 90/91 wo 8. wq P Y

marshalli 91/92 eh ES REL €

Distant 92/93 wt o cu

93/94 B te ER

Urabunana 90/91 A te LE ES

wollomombii 91/92 * * * *

Coombs 92/93 * * * x

93/94 wo dE. £5

Birrima varians 90/91 o £j x * * AES

(Germar) 91/92 vo 44 45 — 9. Q og €

9 2/9 3 * * * * * * *

93/94 * * * * * * *

Pauropsalta 90/91 Soo EB KS. $$ o0

corticinus 91/92 S: * * x

Ewart 92/93 we wo Sd, 5

93/94 SO RD ER cU

Australian Entomologist 23 (2) September 1996 57

Pauropsalta 90/91 o 0 2 € —G 1 4 7

collina Ewart 91/92 E 2 * * * * Ok

92/93 * * * * * * * *

93/94 * * * * * * * *

Notopsalta sp. 90/91 3 0 $9

91/92 * *

92/93 * *

93/94 * *

Cystosoma 90/91 S uu id €

saundersii 91/92 * * * * *

Westwood 92/93 * * * * *

93/94 * * * * *

Cyclochila 90/91

australasiae 91/92 * * * *

(Donovan) 92/93 * * * * *

93/94

Macrotristria 90/91

angularis 91/92

(Germar) 92/93 * * * *

93/94

Psaltoda plaga 90/91 * o * RO 0X* — 0k — 0X

(Walker) 91/92 * * * xD

92/93 * * * xk

93/94 * * * G 8 *

Psaltoda 90/91 * * * * ë *

moerens 91/92 * * * *

(Germar) 92/93 * * * *

93/94

Results

In general, cicadas were active at the study sites from late October to late

February. Within this period, species appeared at differing times and were

active for varying durations over the summer. Seasonality records are

provided for 16 species of cicada for each year of the study (Table 1). Most

adults were active from late November to late January. Pauropsalta collina

Ewart (all years) and Cicadetta puer (Walker) (1992-93 only) were the most

persistent species, occurring from late October to early February. Only

Psaltoda plaga (Walker) occurred until the end of February and none was

present by March. A species tentatively assigned to the genus Notopsalta

was active for the shortest period, from late December to late January. All

species were remarkably consistent from year to year in their times of

appearance and persistence through the season. Not all species, however,

were present in all study years. Macrotristria angularis (Germar) was present

in 1992-93 only, whilst Cyclochila australasiae (Donovan) and Cicadetta

58 Australian Entomologist 23 (2) September 1996

-=> 1990/91

-m- 1991/92

—— 1992/93 —_

45 |. —*-1993/94 ,

Number of cicada species

| Ja 1

0 : Hn—u

| Lg

Sept. Oct. Nov. Dec. Jan. Feb. Mar.

Sampling date (2nd and 4th week of

each month)

Fig. 1. Seasonal activity curve of cicada species at sites in the vicinity of

Armidale, NSW during the period September to March of the years 1990-91, 1991-

92, 1992-93 and 1993-94.

landsboroughi (Distant) were absent during 1990-91.

Figure 1 shows a plot of species number versus sampling date for each year

of the study. Species number peaked during December to early January, and

declined rapidly thereafter, with most species absent by late January. Pearson

correlations and significance levels for species number against climatic and

Australian Entomologist 23 (2) September 1996 59

Table 2 Pearson correlations for species number against maximum and

minimum temperature (°C), rainfall (mm) and photoperiod (hours of daylight).

Year Max. Min. Rainfall Photoperiod

1990-91 r 0.783 0.684 0.337 0.865

t(12) 4.36 3.25 1.24 5.97

P « 0.01 « 0.01 > 0.2 < 0.01

1991-92 r 0.673 0.395 0.331 0.815

1(12) 3.15 1.49 1.21 4.87

P «001 >0.1 > 0.2 < 0.01

1992-93 r 0.427 0.581 0.738 0.822

t(12) 1.63 2.47 3.79 5.00

P > 0.1 < 0.05 < 0.01 < 0.01

1993-94 r 0.511 0.526 0.556 0.852

t(12) 2.06 2.14 2.32 5.64

P >0.05 > 0.05 < 0.05 < 0.01

regional variables are shown in Table 2. Species number was significantly

and positively correlated with day length in all years and with average

maximum daily temperature during 1990-91 and 1991-92 but not 1992-93 or

1993-94. Correlations of species number with average minimum daily

temperature were significant during 1990-91 and 1992-93 only. Species

number was significantly correlated with rainfall during 1992-93 and 1993-94

but not 1990-91 or 1991-92.

Discussion

The seasonal activity of cicadas on the New England Tablelands appears to

reflect dominant climatic conditions. The adverse (cool) season is spent

below ground as immatures feeding on roots. The favourable (warm) season

(as indicated by the frost free period) is relatively short, extending from

November to March. The first adult cicadas appear during late October. The

species per fortnight curve (Fig. 1) shows a rapid increase during November,

peaking during December / January with a rapid decline in late January /

February. Although significant correlations of species number with

maximum or minimum daily temperature and rainfall occurred in some years,

only photoperiod (hours of daylight) was significantly and positively

correlated with species number in all years. Adult cicadas are strongly

heliothermic. Adult reproductive behaviour is restricted to daylight hours in

all species except the crepuscular Cystosoma saundersii Westwood. Clearly

species are timing their seasonal appearance to coincide with the greatest

duration of daylight hours. Such behaviour would enable individuals to

maximise the time spent undertaking reproductive activities.

60 Australian Entomologist 23 (2) September 1996

As adult cicadas are thought to be relatively short lived (usually about 2-4

weeks: Moulds 1990), variation observed in seasonal persistence between

species may be a consequence of differences in the synchrony of adult

emergence. Species present for several months (e.g. P. collina, Birrima

varians) may have a staggered emergence, whereas shorter periods of seasonal

activity (e.g. Notopsalta sp., U. wollomombii) may reflect greater

synchrony in emergence.

The species activity curve presented here for cicadas on the New England

Tablelands closely parallels activity patterns of other insect taxa at localities

with short summers (Shapiro 1975, Wolda 1988). A short period of seasonal

activity and a well defined seasonal peak appear to be typical of many insect

faunas occupying either the higher latitudes or those occurring at altitude.

Shapiro (1975) provides data on numbers of butterfly species in the Sierra

Mountains (7000' altitude) of California largely restricted to a four month

flight period. On average, cicadas on the New England Tablelands become

active approximately one month later and disappear approximately one to two

months earlier than would be expected when extracting seasonality data for

the same species from Moulds (1990). Ewart (1989) provides seasonality

records for Pauropsalta spp. from Queensland as extending from September to

May.

Acknowledgment

S. Winterton is thanked for critically commenting on the manuscript.

References

EWART, A. 1989. Revisionary notes on the genus Pauropsalta Goding and Froggatt

(Homoptera: Cicadidae) with special reference to Queensland. Memoirs of the Queensland

Museum 27: 289-375.

MOULDS, M.S. 1990. Australian cicadas. New South Wales University Press, Kensington.

SHAPIRO, A.M. 1975. The temporal component of butterfly species diversity. Pp 181-195 in:

Cody, M.L. and Diamond, J.M. (eds.), Ecology and Evolution of Communities. Harvard

University Press, Cambridge.

WOLDA, H. 1988. Insect seasonality: why? Annual Review of Ecology and Systematics 19:

1-18.

YOUNG, D. 1972. Analysis of songs of some Australian cicadas (Homoptera: Cicadidae).

Journal of the Australian Entomological Society 11: 237-243.

Australian Entomologist 23 (2) September 1996 61

SEASONAL ABUNDANCE, DISTRIBUTION, HOSTS

AND TAXONOMIC PLACEMENT OF

DIPTEROPHAGUS DACI DREW & ALLWOOD

(STREPSIPTERA: DIPTEROPHAGIDAE)

A.J. ALLWOOD! and R.A.I. DREW?

1 Regional Fruit Fly Project, c/- South Pacific Commission, Private Mail Bag, Nabua, Suva, Fiji

2 Department of Primary Industries, Plant Protection Unit, Meiers Road, Indooroopilly, Qld

4068 (address for correspondence)

Abstract

Nineteen species of dacine fruit flies are recorded as hosts of Dipterophagus daci Drew and

Allwood, a strepsipteran parasite which has been reared only from Tephritidae. Aspects of

the ecology of D. daci in northern Australia, in relation to the seasonal abundance of its two

most abundant hosts, Bactrocera aquilonis (May) and B. tenuifascia (May), are reported.

Monthly captures of the two hosts over a 12 month period indicated that their populations

increased with the onset of higher temperatures, moisture levels and availability of host fruits.

Numbers of D. daci peaked about one month after B. aquilonis and two months after B.

tenuifascia and evidence indicated that the seasonal activity of D. daci was dependent upon the

availability of its host and rainfall. Higher levels of parasitism occurred in B. aquilonis than in

B. tenuifascia and rapid increase in host populations was probably one significant factor in the

prevention of the parasite from causing a high level of parasitism during the period of high

fruit fly population. The fruit flies and their parasites were more abundant in wet than dry

habitats. D. daci is recorded for the first time from the Solomon Islands. The family

Dipterophagidae is reinstated.

Introduction

Most species of Strepsiptera occur in the Palaeotropical region of the world

which incorporates the tropics and subtropics (Kinzelbach 1978). Little is

known about the biology and ecology of these insects and no information has

been reported on the seasonal abundance of species or levels of parasitism

achieved in nature. The most comprehensive biological study was on a

strepsipterous parasite of Antestia spp. (Pentatomidae) by Kirkpatrick (1937).

Brief accounts of general biology and life histories have been reported by

Perkins (1905), Ogloblin (1939), Bohart (1941) and Riek (1970), while

Raatikainen and Heikinheimo (1974) studied the flying times of strepsipteran

males at different latitudes in Finland.

Fruit flies (Tephritidae: Dacinae) are endemic to northern and eastern

Australia. Considerable research has been undertaken on the ecology of

Dacinae in endemic tropical and subtropical rainforest habitats and in

cultivated orchards (Fitt 1981, Drew and Hooper 1983, Fletcher 1987). The

influence of hymenopterous parasites and various predators was investigated

by Drew (1987) in rainforest habitats. Major reductions in fly populations

were due to fruit-eating vertebrates and hymenopterous parasites had only a

minor effect.

Dipterophagus daci Drew and Allwood is unique in being the only

strepsipteran species so far described which is a parasite of Diptera. It is

most abundant in the Northern Territory where it parasitises a number of fruit

fly species. Because of the economic importance of fruit flies to Australia,

including some of the hosts of D. daci, data have been collected on host

62 Australian Entomologist 23 (2) September 1996

Garden Point 3 Lf

Site M007

Fig. 1. Location of trapping sites and meteorological stations used in this

study.

records, seasonal abundance and geographic distribution of the parasite in its

endemic habitats. Percent parasitism levels have been calculated also.

Taxonomic note

The family Dipterophagidae was established for D. daci based on a

combination of male, female and first stage larval characters (Drew and

Allwood 1985). Kathirithamby (1989) treated it as a subfamily of

Halictophagidae on the basis of a number of characters that D. daci and

known halictophagids have in common. However, other families of

Strepsiptera also possess some of these characters and families such as

Bohartillidae were separated on the basis of one of these characters (number of

antennal segments).

Kathirithamby (1989, 1992) noted that most species placed in the

Halictophagidae are parasites of Hemiptera and that all (except D. daci) had

males with seven antennal segments and lateral flabella on more than one

segment (except in the Tridactylophaginae which has one lateral flabellum

and parasitises Orthoptera) and females with abdominal segments 1-5 with

one genital aperture each and the cephalothorax flattened. A measurement of

the 7th antennal segment of D. daci was given (Kathirithamby 1989, p. 78)

but this species has only six such segments. Also it was stated that the

female of D. daci possessed genital pores on abdominal segments 4-6

(Kathirithamby 1989, pp 76, 78) or 3-6 (Kathirithamby 1992, p. 166).

However, this species has genital pores on abdominal sterna 3-5 (Drew and

Allwood 1985).

The following combination of characters is unique to D. daci: male with six

antennal segments and a lateral flabellum on segment 3 only; female with a

bell-shaped (rounded) cephalothorax and genital openings on abdominal sterna

3-5. These characters, plus the host, render it very distinct from all true

halictophagids and the family Dipterophagidae is reinstated.

Australian Entomologist 23 (2) September 1996 63

Garden Point, Melville 1.

(Sites M007, M033)

500 Middle Point — .— ...... TE

(Site AHO14)

400 Pine Creek moss

(Sites DROO4, MRF)

Monthly Rainfall (mm)

Mean Daily Temperature (°C)

Mean Relative Humidity (%) at 9 a.m.

AUG SEP OCT NOV DEC JAN FEB MAR APR MAY JUN JUL

Fig. 2. Meteorological data (monthly rainfall, mean maximum and minimum

daily temperatures and mean relative humidities at 9 am) at weather stations that

represent trapping sites.

64 Australian Entomologist 23 (2) September 1996

FIG.3. SITE M007 - MELVILLE ISLAND

No. of B. tenuifascia males e——e

in methyl eugenol trap/month

No. of B, aquilonis males o-—o

in cue lure trap/month

TOTAL NO. FLIES

FIG.4. No. parasitised B. tenuifascia — e———e

males in methyl eugenol trap/month

No. parasitised B. aquilonis 0——o

males in cue lure trap/month

160

140

120

100

NO. PARASITISED FLIES

AUG SEP OCT NOV DEC JAN FEB MAR APR MAY JUN JUL

Figs. 3 and 4. (3). Number of male Bactrocera aquilonis and B. tenuifascia per

trap month at site M007 on Melville Island; (4). Number of male Bactrocera

aquilonis and B. tenuifascia per trap month parasitised by Dipterophagus daci at

site M007 on Melville Island.

Australian Entomologist 23 (2) September 1996 65

FIG.5.

SITE M007 - MELVILLE ISLAND

% parasitism of B. tenuifascia e——e

males by Strepsipteran/month

96 parasitism of B. aquilonis 0—o

males by Strepsipteran/month

PERCENTAGE PARASITISM

AUG SEP OCT NOV DEC JAN FEB MAR APR MAY JUN JUL

Fig. 5. Percent of male Bactrocera aquilonis and B. tenuifascia per trap month

parasitised by Dipterophagus daci at site M007 on Melville Island.

Materials and Methods

The work was carried out in a region across the north of the Northern

Territory. Two sites were selected on Melville Island and three on the

mainland (Fig. 1). Taracumbi Falls (Site M033) and the site 7 km N of Paru

Village are both wet habitats on Melville Island, about 40 km apart. The site

at Moline Rock Falls (Site MRF) ca 70 km NE of Pine Creek is a wet

habitat. The site near the Wildman River (West Branch) ca 100 km E of

Darwin (Site AH014) and that ca 70 km NE of Pine Creek (Site DR004) are

classed as dry habitats. The wet sites are characterised by the presence of

surface water for the whole year and situated in or near monsoonal rainforest.

The dry sites are devoid of surface water for part of the year and are situated in

open eucalypt woodland.

The climate of the study area is classified as semi-arid tropical (Williams et

al. 1985). It is characterised by having a distinct “wet” season during

December to April and a "dry" season for the remainder of the year. Mean

daily maximum and minimum temperatures, mean relative humidities at 9

am and monthly rainfall data were obtained from weather stations at Garden

Point (Melville Island), Middle Point and Pine Creek, selected because of

their proximity to the trapping locations (Fig. 2).

Fruit fly populations were monitored at ca 100 localities representative of the

endemic vegetation. This was part of a broad surveillance strategy for exotic

fruit flies carried out under the auspices of the North Australian Quarantine

Survey. At each locality two Steiner type fruit fly traps were set, one

66 Australian Entomologist 23 (2) September 1996

FIG.6. SITE MO33 - MELVILLE ISLAND

No. of B. tenuifascia males e———e

in methyl eugenol trap/month

No. of B. aquilonis males Qe)

in cue lure trap/month

TOTAL NO. FLIES

[^]

FIG.7. No. parasitised B. tenuifascia 6— —o

males in methyl eugenol trap/month

No. parasitised B. aquilonis o—o

males in cue lure trap/month

NO. PARASITISED FLIES

JUN JUL

AUG OCT NOV DEC JAN FEB MAR APR MAY

Figs. 6 and 7. (6). Number of male Bactrocera aquilonis and B. tenuifascia per

trap month at site M033 on Melville Island; (7). Number of male Bactrocera

aquilonis and B. tenuifascia per trap month parasitised by Dipterophagus daci at

site M033 on Melville Island.

SEP

containing methyl eugenol (4 ml + 2 ml 50% w/v malathion e. c.), the other

containing cue lure (4 ml + 2 ml 50% w/v malathion e. c.). This study was

carried out for one year from August 1977. All traps were cleared of flies at

ca one-month intervals at which time the lure plus insecticide baits were

changed. The trapped flies (all males) were identified, counted and examined

under a stereo microscope for presence of strepsipteran parasites. At five

localities, selected as wet or dry habitats, the number and percentage of

parasitised flies were calculated for two species, Bactrocera aquilonis (May)

and B. tenuifascia (May).

Results

Seasonal abundance

Data are presented for B. aquilonis from three sites (M007, M033, AH014)

and for B. tenuifascia from five sites (M007, M033, MRF, DR004, AH014).

At each site flies were present for the entire one year trapping period. Data

Australian Entomologist 23 (2) September 1996 67

FIG.8. SITE MRF - MOLINE ROCK FALLS (PINE CREEK) - WET AREA

SITE DROO4 - DALY RIVER ROAD - DRY AREA

No. of B. tenuifascia males at MRF/month o

No. of B. tenuifascia males at DROO4/month 4———a

a 1200

TOTAL NO. FLIE

FIG.9. % parasitism of B, tenuifascia males by

Strepsipteran at MRF 4————4

3 and DROO4 2———

PERCENTAGE PARASITISM

AUG SEP OCT NOV DEC JAN FEB MAR APR MAY JUN JUL

Figs. 8 and 9. (8). Number of male Bactrocera tenuifascia per trap month at

sites MRF and DR004 in the Northern Territory; (9). Percent of male Bactrocera

tenuifascia per trap month parasitised by Dipterophagus daci at sites MRF and

DRO004 in the Northern Territory.

for B. aquilonis from MRF and DRO004 have not been presented but support

conclusions drawn from the other data.

At site M007 the numbers of B. aquilonis increased from August to reach a

peak (ca 6000 trapped flies per month) in mid-October and early November,

then declined to mid-January after which they remained at a low level (Fig.

3). At site M033 B. aquilonis had two population peaks, in mid-October (ca

6000) and mid-May (ca 4000), the latter being for a shorter period (Fig. 6).

The fruit fly population at M033 showed similar rates of increase and decline

in mid-October to that at M007. At site AH014 B. aquilonis also had two

peaks, one in mid-November and the other in mid-May (each ca 1500).

At site M007 the numbers of B. tenuifascia increased from mid-September to

reach a peak (ca 4000) in early December, then declined to mid-March after

which they remained at a low level (Fig. 3). At site M033 the numbers of B.

tenuifascia were low, with a maximum of 1100 in mid-August: before

declining to ca 100 in mid-November, after which they remained at

approximately the same level for the remainder of the study period (Fig. 6).

At site MRF the numbers of B. tenuifascia increased from August to reach a

68 Australian Entomologist 23 (2) September 1996

peak (ca 1800) in mid-September, then entered a slow decline to June (Fig.

8). At sites DROO4 (Fig. 8) and AH014 (Fig. 10) B. tenuifascia reached

peaks in mid-August (ca 600 and 4000 respectively), after which they entered

a slow decline.

Flies parasitised by D. daci generally were restricted to shorter periods of the

year. At site M007 the number of parasitised B. aquilonis increased in mid-

September to reach a peak (ca 165) in mid-November, then declined to mid-

February (Fig. 4). Parasites were present for ca 10 months. At the peak of

activity of B. aquilonis (October-November), the level of parasitism was

2.6% and this reached a peak of 7% in mid-January when the fly population

had declined (Fig. 5). Concurrently, the number of parasitised B. tenuifascia

increased slowly from mid-October to reach a peak in mid-February (ca 15),

then declined to mid-March (Fig. 4). Parasites were present for 5 months.

At the peak of activity of B. tenuifascia (early December), the level of

parasitism was 0.25% and this peaked at 6.5% in mid-April when the fly

population had declined markedly (Fig. 5).

At site M033 the number of parasitised B. aquilonis increased from mid-

August to peak at mid-December (ca 80 flies and 7% parasitism) (Fig. 7).

During the second peak of B. aquilonis in mid-May there was almost no

parasite activity (ca 0.1% parasitism) (Fig. 7). At this site the number of

parasitised B. tenuifascia was at a very low level but followed the population

trend of the host species (Fig. 7).

At site MRF the level of parasitism in B. tenuifascia was 0.7% at the peak of

the fly population, increasing to ca 3.5% in mid-December when the fly

numbers had declined to a low level (Fig. 9). At site DROO4 the level of

parasitism in B. tenuifascia was ca 0.25% at the peak of the fly population,

increasing to ca 1.5% in mid-December when the fly population was well

into its decline (Fig. 9).

At site AH014 the number of parasitised B. aquilonis reached a peak in mid-

November (Fig. 11). The level of parasitism was ca 0.896 during the peak of

the fly population in mid-November and 1.3% in December when the host

population was low. There was no parasite activity for eight months from

mid-January (Fig. 11). At this site the period of parasite activity in B.

tenuifascia was only four months, from mid-August to mid-December (Fig.

11). The level of parasitism in this fly species was ca 0.15% at the peak of

its population in mid-August and 0.4% in mid-November when the fly

population had declined.

Geographic distribution

D. daci is known in Australia from Melville Island, coastal and subcoastal

areas of both the Northern Territory and Cape York Peninsula, some Torres

Strait islands and Townsville (northern Qld), Mt Glorious, Palmwoods and

Australian Entomologist 23 (2) September 1996 69

FIG.10. SITE AH014 - ARNHEM HIGHWAY

4000 No. of B. tenuifascia males @——®

in methyl eugenol trap/month

No. of B. aquilonis males o——o

3000 in cue lure trap/month

2000

TOTAL NO. FLIES

1000

FIG.11. No. parasitised B. tenuifascia 6———9

males in methyl eugenol trap/month

No. parasitised B. aquilonis o_o

males in cue lure trap/month

NO. PARASITISED FLIES

AUG SEP OCT NOV DEC JAN FEB MAR APR MAY JUN JUL

Figs. 10-11. (10). Number of male Bactrocera aquilonis and B. tenuifascia per

trap month at site AH014 in the Northern Territory; (11) Number of male

Bactrocera aquilonis and B. tenuifascia per trap month parasitised by

Dipterophagus daci at site AH014 in the Northern Territory.

Redland Bay (SE Qld) (Drew and Allwood 1985, Drew unpublished data). It

occurs also on Guadalcanal, Solomon Islands (new record).

Host records

D. daci has been recorded from 19 dacine hosts: Bactrocera aquilonis (May),

B. cacuminata (Hering), B. decurtans (May), B. mayi (Hardy), B.

neohumeralis (Hardy), B. peninsularis (Drew & Hancock), B. tenuifascia

(May), B. tryoni (Froggatt), Dacus bellulus Drew & Hancock (Drew and

Allwood 1985), plus B. abscondita (Drew & Hancock), B. aeroginosa (Drew

& Hancock), B. breviaculeus (Hardy), B. frauenfeldi (Schiner), B. jarvisi

(Tryon), B. musae (Tryon), B. perkinsi (Drew & Hancock), Dacus aequalis

Coquillett (new records from Australia), B. froggatti (Bezzi) and B. umbrosa

(Fabricius) (new records from Solomon Islands). B. aquilonis, B. musae, B.

neohumeralis and B. tryoni are major pest species in Australia.

70 Australian Entomologist 23 (2) September 1996

Discussion

The seasonal activity of dacine fruit flies is dependent upon temperature,

rainfall and the state of development of the host fruit (Bateman 1968, Drew

and Hooper 1983). Drew and Hooper (1983) also demonstrated that male lure

trap catches provided an accurate assessment of the seasonal changes in dacine

populations.

Both B. aquilonis and B. tenuifascia were trapped throughout the year at all

sites but their populations increased in the August-December period prior to

the onset of the wet season. This was probably due to increasing temperature

and relative humidity and increased wild host fruit production, as found for B.

cacuminata (Hering) by Drew and Hooper (1983). The major wild host fruits

of B. aquilonis are Glycosmis trifoliata (Blume) Sprengel, Micromelum

minutum (Forster f.) Wight and Arn. and various species of Syzygium

(Smith et al 1988). Their peak fruiting period is October-November,

immediately prior to the onset of the wet season. Second peaks in

populations of B. aquilonis occured from March-May at sites M033 and

AH014, coinciding with the fruiting period of Terminalia ferdinandiana Exell

(March-June). Populations of B. tenuifascia also reflected the availability of

its major hosts; Planchonella pohlmaniana (F. Muell.) fruits April-November

while P. arnhemica (F. Muell.) P. Royen fruits June-November (Fitt 1981).

Rainfall is probably more important later because of its influence on the

survival of pupae and emerging adults (Bateman 1972). The populations of

B. aquilonis were higher than those of B. tenuifascia at all study sites.

Populations of B. aquilonis were 3-4 times larger in the wet habitats (M007

and M033) than in the dry habitat (AH014). Similarly, populations of B.

tenuifascia were usually higher in the wet habitats but this difference was not

as consistent as in B. aquilonis.

The increase in parasite activity followed that of the host flies. However,

there was a one month lag period for the parasite in B. aquilonis and a two

month lag period in B. tenuifascia. The activity of the parasites coincided

with the onset of the wet season and even when there was a second B.

aquilonis population peak at M033, in the dry season, the parasites were

virtually absent. There was a shorter period of parasite activity and a lower 96

peak parasitism level in both host species in the dry habitats. The level of

parasitism in B. tenuifascia was consistently lower than that in B. aquilonis,

indicating that the latter species is a better host for the parasite.

The seasonal activity of D. daci was dependent upon the availability of its

hosts and rainfall. However, in spite of the availability of large fruit fly

populations, the parasite was not efficient in inducing high rates of

parasitism. There were very low percentages of parasitism when the fly

populations were at their peaks and higher levels only when the fly

populations declined markedly. This may be explained by the fact that

Bactrocera species are r selected species, undergoing rapid increases in

Australian Entomologist 23 (2) September 1996 71

population when host fruits are available and environmental conditions

permit (Bateman 1972, Drew and Hooper 1983). The rapid population

increases of the host species appear too large for the rate of increase of the

parasite. Although no evidence exists, low levels of parasitism by D. daci

may be explained by low survival rates of triungulins, small numbers of

triungulins actually coming in contact with fruit fly adults, or male dacines

being less favoured as hosts than females.

There was no apparent effect on the external appearance of the host and on the

size and colour of the testes, similar to Corioxenos antestiae Blair

parasitising Antestia spp. (Pentatomidae) in Africa (Kirkpatrick 1937).

However, in Antestia the stylopised females never produced mature eggs and

the stylopised males were incapable of fertilising eggs even when they

copulated. Even if D. daci has a similar effect on its fruit fly host species, it

seems that it will never have a marked influence on population reduction.

Kirkpatrick (1937) suggested that the combination of an egg parasite and a

strepsipteran may have a better chance of inducing larger population

reductions in Antestia.

Gregarious parasitism, recorded by Drew and Allwood (1985), occurs with

larger numbers of parasites per host than that reported by Kirkpatrick (1937).

We have observed the fungus infection in the empty male pupal cases

recorded by Kirkpatrick (1937) and Bohart (1941). One female parasite

collected at Palmwoods, SE Qld, had over 3000 triungulin larvae indicating

that they have a very large reproductive rate. This is probably essential as the

triungulins must be exposed to severe environmental stresses between

emerging from the female and finding a host.

Dipterophagus daci has now been recorded in 19 dacine host species and more

frequently in Cape York Peninsula than SE Queensland. There is no evidence

to suggest that it is going through a southward expansion of its distribution.

It predominates in the northern tropics and it is probable that recordings in

SE Qld are related to increased collections of flies for other ecological studies

and bait spray trials.

Acknowledgments

Mrs M.C. Romig, QDPI, prepared the figures. The specimens were collected

during the North Australian Quarantine Survey and in Solomon Islands by

the RFFP and ACIAR projects. The study was partly funded by the

Australian Biological Resources Study. This support is gratefully

acknowledged.

References

BATEMAN, M.A. 1968. Determinants of abundance in a population of the Queensland fruit

fly. Symposium of the Royal Entomological Society of London 4: 119-131.

BATEMAN, M.A. 1972. The ecology of fruit flies. Annual Review of Entomology 17: 493-

518.

72 Australian Entomologist 23 (2) September 1996

BOHART, R.M. 1941. A revision of the Strepsiptera with special reference to the species of

North America. University of California Publications in Entomology 7: 91-159.

DREW, R.A.I. 1987. Reduction in fruit fly (Tephritidae: Dacinae) populations in their

endemic rainforest habitat by frugivorous vertebrates. Australian Journal of Zoology 35: 283-

288.

DREW, R.A.I. and ALLWOOD, A.J. 1985. A new family of Strepsiptera parasitising fruit

flies (Tephritidae) in Australia. Systematic Entomology 10: 129-134.

DREW, R.A.I. and HOOPER, G.H.S. 1983. Population studies of fruit flies (Diptera:

Tephritidae) in south-east Queensland. Oecologia 56: 153-159.

FITT, G.P. 1981. Ecology of northern Australian Dacinae (Diptera: Tephritidae). II.

Seasonal fluctuations in trap catches of Dacus opiliae and D. tenuifascia, and their relationship

to host phenology and climatic factors. Australian Journal of Zoology 29: 885-894.

FLETCHER, B.S. 1987. The biology of dacine fruit flies. Annual Review of Entomology 32:

115-144.

KATHIRITHAMBY, J. 1989. Review of the Order Strepsiptera. Systematic Entomology 14:

41-92.

KATHIRITHAMBY, J. 1992. Descriptions and biological notes of Halictophagidae

(Strepsiptera) from Australia, with a checklist of the world genera and species. Invertebrate

Taxonomy 6: 159-196.

KINZELBACH, R.K. 1978. Strepsiptera. Die Tierwelt Deutschlands Teil 65, 166 pp.

KIRKPATRICK, T.W. 1937. Studies on the ecology of coffee plantations in East Africa. II.

The Autecology of Antestia spp. (Pentatomidae) with a particular account of a strepsipterous

parasite. Transactions of the Royal Entomological Society of London 86: 247-343.

OGLOBLIN, A.A. 1939. The Strepsiptera parasites of ants. Proceedings of the 7th

International Congress of Entomology 2: 1277-1284.

PERKINS, R.C.L. 1905. Leaf-hoppers and their natural enemies (Pt. III. Stylopidae). Bulletin

of the Hawaiian Sugar Planters' Association Experiment Station 1: 86-111.

RAATIKAINEN, M. and HEIKINHEIMO, O. 1974. The flying times of Strepsiptera males

at different latitudes in Finland. Annales entomologici fennici 40: 22-25.

RIEK, E.F. 1970. Strepsiptera, pp. 622-635 in The insects of Australia. Melbourne University

Press.

SMITH, E.S.C., CHIN, D., ALLWOOD, A.J. and COLLINS, S.G. 1988. A revised host list of

fruit flies (Diptera: Tephritidae) from the Northern Territory of Australia. Journal of

Agricultural and Animal Science 45: 19-28.

WILLIAMS, J., DAY, K.J., ISBELL, R.F. and REDDY, S.J. 1985. Constraints to agricultural

development: soils and climate, pp. 31-92 in Agro-research for the semi-arid tropics: North-

West Australia. University of Queensland Press.

Australian Entomologist 23 (2) September 1996 73

NOTES ON THE LIFE HISTORY OF

NACADUBA KURAVA FELSINA WATERHOUSE & LYELL

(LEPIDOPTERA: LYCAENIDAE)

C.E. MEYER

10 Anne Clark Avenue, Nicholls, ACT, 2913

Abstract

Notes are given on the life history of Nacaduba kurava felsina Waterhouse & Lyell and the

food plant Embelia curvinervia (Myrsinaceae) recorded.

Introduction

The white line blue Nacaduba kurava felsina is known to occur in the

Northern Territory from Darwin south to Mataranka (Common and

Waterhouse 1981, Dunn and Dunn 1991). The life history of the eastern

subspecies N. kurava parma Waterhouse & Lyell has been described

(Common and Waterhouse 1981) but, apart from one record of a larva feeding

on the young foliage of a rainforest tree growing on the banks of the

Katherine River (Common and Waterhouse 1981), little is known of the life

history of N. k. felsina.

During the past two years adults were prevalent in early June at Oolloo

Crossing on the Daly River. Eggs and early instar larvae were discovered in

late May 1994 and reared on cuttings of the host plant.

Life History

Host Plant: Embelia curvinervia (Family Myrsinaceae) (Reynolds 1991).

The descriptions of the immature stages of N.k. parma given in Common and

Waterhouse (1981) are similar to those of N. k. felsina, with variation

occurring in colouring and markings as noted below.

Egg (Fig. 1): Small, white, mandarin shaped; dense clockwise and

anticlockwise series of ridges and pits radiating from micropylar depression;

ridges intersect at coarse erect spines; 0.55 mm diameter x 0.25 mm high.

Fig. 1. Nacaduba kurava felsina, eggs.

First instar larva: Olive green with purple dorsal lines from head to last

abdominal segment. Dorsal lines give early instar larva the appearance of

74 Australian Entomologist 23 (2) September 1996

being purple but on closer examination base colouring can be seen. Head

pale brown.

Final instar larva: Olive green or purple, strongly humped above and

prominently segmented with purple middorsal and subdorsal lines; head pale

brown. Size 9-12 mm (n=40).

Pupa: Covered with very short erect hairs and black spots; head and thorax

brown with pinkish brown abdomen; black spotted middorsal line present

from head to mesothorax; first abdominal segment with two conspicuous

dorsolateral black spots; attached by anal hooks and central girdle. Average

size 9 mm x 3.5 mm.

Discussion

The host plant is known from several sites in the Northern Territory and near

the Claudie River, Cape York Peninsula (Reynolds 1991). In the Northern

Territory it and associated butterfly colonies have been found at Oolloo

Crossing on the Daly River, the Adelaide River bridge on the Daly River road

and at the Marrakai Road jungle, 50 km south of Darwin. The host plant is a

rambling woody vine which appears to grow near permanent water.

Eggs generally are laid singly on new growth or flowers of the host plant.

Larvae at first were collected from new growth of the host plant in late May

and from the flowers a fortnight later. Larvae appeared to prefer flowers but

these are short-lived. During this period larval growth is rapid, reaching final

instar in 11 to 14 days. Pupal duration was 6 to 11 days.

Larvae occur throughout the year but are more numerous from May to July,

indicating a preference for the dry season (winter). Their appearance at other

times depends upon the presence of new growth or flowers on the host plant.

Adult females flutter around the host plant and rest on twigs or branches of

the vine. Adult males are territorial, settling head downwards on twigs or

small branches higher up in the canopy adjacent to the host plant.

Acknowledgments

I thank Ian Cowie of the Northern Territory Herbarium, Palmerston for

identifying the food plant, Dave Wilson for his company and assistance in the

field in May 1994 and Richard Weir for the egg photography.

References

COMMON, I.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Pp xiv + 682.

Angus and Robertson, Sydney.

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

and taxonomy. Part 3. Family Lycaenidae. Pp iii + 336-512. Privately published by the

authors, Melbourne.

REYNOLDS, S.T. 1991 The Genus Embelia N. Burman (Myrsinaceae) in Australia.

Austrobaileya 3(3): 361-367.

Australian Entomologist 23 (2) September 1996 75

LEPIDOPTERA BREEDING RECORDS FROM

ALPHITONIA SPECIES (RHAMNACEAE)

AT PALUMA, NORTH QUEENSLAND

R.V. JACKSON

Department of Zoology, James Cook University, Townsville, Qld 4811

(Present address: C.R.C. Tropical Rainforest Ecology and Management, Department of

Zoology, James Cook University, Townsville, Qld 4811)

Abstrac

Fourteen ea of Lepidoptera (12 previously unrecorded) feeding upon three species of

Alphitonia (Rhamnaceae) from the Paluma area, North Queensland, are documented.

Introduction

Three species of Alphitonia grow in the Paluma area (19°00'S 146°12'E,

altitude 892 m, 80 km north-west of Townsville, North Queensland). A.

petriei C.T.White & Braid is a pioneer tree, common along disturbed and

undisturbed rainforest margins. A. whitei Braid is more shade tolerant and

grows in small gaps and the understorey, while A. excelsa (A. Curn. ex

Fenzl) Reissek ex Benth. occurs in dry sclerophyll communities. To date 11

Lepidoptera species have been recorded as herbivores of Alphitonia species.

All but one are from A. excelsa. These are: Bucculatrix sp. (Bucculatricidae),

which is a leaf-miner in early stages but later feeds exposed on the leaf

surface; Carmenta chrysophanes (Meyrick) (Sesiidae), larvae of which bore

the inner bark; Casbia rectaria Walker (Geometridae), a leaf feeder; the leaf-

miner Leucoptera sp. (Lyonetiidae); Opodiphthera austrophela (Walker)

(Saturniidae), another leaf feeder (Common 1990); plus 5 species of

Lycaenidae which feed on the leaves or flower buds (Common and

Waterhouse 1981). The recorded distributions of. C. chrysophanes and C.

rectaria cover the Paluma area. The remaining species, Aenetus mirabilis

(Rothschild) (Hepialidae), has a stem-boring larva on an unspecified species

of Alphitonia (Common 1990) but its described location at the edge of

rainforest and its geographical range suggest its host is probably A. petriei.

At Paluma I observed and reared specimens of 7 lepidopteran families

comprising 14 species. These are listed below by family (Table 1). All are

folivores and all except one are new records. These are also the first

confirmed records from A. petriei and A. whitei. Voucher specimens of both

plants and insects are deposited at the Department of Zoology, James Cook

University of North Queensland.

Acknowledgments

Thanks go to Ted Edwards (CSIRO Division of Entomology, Canberra) for

identifying the geometrids, E. ?postvittana, S. janetta, O. mendosa and E.

epidela, and to an anonymous reviewer for constructive criticism which

improved the manuscript.

76 Australian Entomologist 23 (2) September 1996

Table 1. Lepidoptera observed feeding on Alphitonia species and reared to

maturity.

SPECIES FAMILY HOST MONTH(S)

PLANT OBSERVED

Casbia rectaria Walker Geometridae A. petriei* — Feb-Aug;

A. excelsat Oct-Nov

A. whitei

C. calliorma Turner Geometridae A. whitei Feb-May

A. petriei

C. didymosticta Turner Geometridae A. petriei Jun; Aug

C. scardamiata (Warren) Geometridae A. petriei Apr May;

uly

Ectropis rufobrunnea Warren Geometridae A. petriei Aug; Dec

Epiphyas ?postvittana (Walker) Tortricidae A. petriei Aug-Oct

A. excelsa

Syntherata janetta (White) Saturniidae A. excelsa Feb; Dec

Opodiphthera eucalypti (Scott) Saturniidae A. excelsa Mar

Olene mendosa Hübner Lymantriidae A. petriei Apr

Euproctis ?epidela Turner Lymantriidae A. excelsa Feb; Aug

Orgyia papuana Riotte Lymantriidae A. petriei Mar

Mecytha fasciata (Walker) Zygaenidae A. excelsa Dec

Anaxidia lozogramma Turner Limacodidae A. petriei Dec

Danis hymetus (C.& R. Felder) Lycaenidae A. petriei Feb-June

* where >l species of Alphitonia hosted a herbivore, they are listed in order of

relative frequency of use

1 previously recorded host plant

References

COMMON, LF.B. 1990. Moths of Australia. Melbourne University Press, Melbourne.

COMMON, L.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia (revised ed.)

Angus and Robertson, Sydney.

Austrralian Entomologist 23 (2) September 1996 77

SMOKE FLIES (DIPTERA: PLATYPEZIDAE)

AND THE SYDNEY BUSHFIRES

Daniel J. Bickel

Australian Museum, College Street, Sydney, NSW 2000; e-mail: danb@amsg.Austmus.gov.au

Abstract

Swarms of the smoke fly Microsania australis Collart were found at a smoking log in Royal

National Park, New South Wales, three weeks after the area was burnt by severe bushfires.

The attraction of Microsania to smoke for mating assembly is discussed.

Introduction

In early January 1994, bushfires burnt large regions of eastern New South

Wales, including some Sydney suburbs. The bushfires were severe in Royal

National Park, a large area of natural bushland south of metropolitan Sydney.

More than 80% of the park was burnt, including most of the northern half.

On January 29, three weeks after the fires, I visited the Park. The landscape

was open and desolate, with the dense mallee heathland reduced to a thick grey

ashbed and a few charred sticks. In the dry sclerophyll woodland many trees

were dead and fallen and standing trees were badly seared.

The bushfires took a heavy toll of fauna, including insects. A few ants from

surviving underground colonies were foraging, otherwise there was no

observable ground fauna. Native cockroaches, beetles and centipedes had been

baked in place under sandstone slabs. I saw few flying insects during a ten

kilometre walk through burnt heath and woodland. Near the Curra Moors

track roadhead I noticed smoke issuing from a log. This was unusual since

the fires had been "out" for almost three weeks and it had rained during that

time. A large fallen Angophora costata trunk and its stump were both

smoldering and the log was almost entirely burnt out. Swarms of tiny flies

were present in smoke from both the log and stump, with some 150-200

individuals in each swarm. Of 68 individuals taken in a sweep, there were 66

males and 2 females of Microsania australis Collart (Platypezidae), confirmed

by the genitalic figure in Collart (1938).

Microsania Zetterstedt is a cosmopolitan genus with four described Australian

species (Chandler 1994). They are commonly known as "smoke flies"

because they form mating swarms in woodsmoke. Indeed, collectors suggest

lighting smoky fires with green wood to attract them. Kessel (1960a, b)

notes that Microsania is attracted to cold smoke used by bee-keepers and even

to shirts previously worn in woodsmoke.

Discussion

Although Microsania australis 1s attracted to woodsmoke, its concentration in

large numbers in such a devastated landscape raises questions:

1. Where did they come from? Nothing is known of the immature stages of

Microsania (Kessel 1987) but other platypezid genera have fungus-feeding

larvae. Nevertheless, the intensity of the bushfires would have destroyed

most larvae within the main burn. The nearest source of unburnt or slightly

78 Austrralian Entomologist 23 (2) September 1996

burnt bushland was Bola Creek, 3-4 km to the west.

2. When did they arrive? The main bushfire would have been too hot and

turbulent to attract the flies. I suspect M. australis came into the area after

the main blaze was extinguished but while logs were still smoldering. As

fires died out, the smoke flies probably gathered at remaining smoke, and the

smoldering trunk was possibly the last source over a wide area.

The sex ratio of the M. australis sample (66 CC’, 2 Q9) is typical of an aerial

mating swarm (Downes 1969). In many insects, aerial swarming is initiated

by environmental cues such as light intensity and relative humidity.

However it seems that Microsania spp. use smoke in two ways: as an

aggregating "scent" to concentrate dispersed individuals of both sexes and as a

swarm marker, the actual smoke plume delimiting the swarm boundary

(individuals rarely leave the plume and follow as it shifts).

Kessel (1989) suggested that attraction to smoke was a positive response to

concentrate individuals in areas recently burnt by forest fires, and that

Microsania bred in fungi specifically associated with fire-scarred wood.

However, this hypothesis has yet to be documented, as larvae and their hosts

are unknown. If this were the case, one would expect Microsania to be co-

adapted with fire-ecology vegetation, such as the eucalypt forests of Australia

or chaparral of California. However, smoke flies also occur in northwestern

Europe and Central African rainforest where natural fires are exceedingly rare.

In these moist regions they often appear in large numbers around smoky fires

(e.g. Chandler 1978). Therefore, although Microsania is distinctly attracted

to smoke for mating, it must have other means of aggregation and swarming

in the absence of fire. Possibly they are attracted to certain plant aromatics

which also facilitate aggregation.

References

CHANDLER, P.J. 1978. Some dipterous opportunists at Windsor Forest, Berks.: the attractions

for flies of bonfires, wood ash and freshly cut logs. Entomologists Gazette 29: 253-257.

CHANDLER, P.J. 1994. The Oriental and Australasian species of Platypezidae (Diptera).

Invertebrate Taxonomy 8: 351-434.

COLLART, A. 1938. Description d'un Microsania nouveau d'Australie (Diptera:

Platypezidae). Bulletin Musée royal d'Histoire naturelle de Belgique 14 (16): 1-4.

DOWNES, J.A. 1969. The swarming and mating flight of Diptera. Annual Review of

Entomology 14: 271-298.

KESSEL, E.L. 1960a. The response of Microsania and Hormopeza to smoke (Diptera:

Platypezidae and Empididae). Pan-Pacific Entomologist 36: 67-68.

KESSEL, E.L. 1960b. Microsanias attracted to cold smoke. Wasmann Journal of Biology 18:

312-313.

KESSEL, E.L. 1987. Platypezidae. Pp 681-688, in J.F. McAlpine, et al., Manual of Nearctic

Diptera Vol 2. Research Branch Agriculture Canada Monograph 28, Ottawa, 1332 pp.

KESSEL, E.L. 1989. Autobiographical Anecdotes. Myia 2, San Francisco, 231 pp

Austrralian Entomologist 23 (2) September 1996 79

A NEW RECORD OF NESOLYCAENA CAESIA

D'APICE & MILLER (LEPIDOPTERA: LYCAENIDAE)

FROM NORTH-EASTERN WESTERN AUSTRALIA

C.E. MEYER

10 Anne Clark Avenue, Nicholls, A.C.T., 2913

Abstract

Nesolycaena caesia d'Apice and Miller is recorded from near Kununurra, Western Australia.

A single female Nesolycaena caesia d'Apice and Miller was taken at a roadside

soak, 1 km N of Amalia Gorge (15°57'S 128°01'E), via Kununurra, Western

Australia, in April 1995. d'Apice and Miller (1992) recorded this butterfly

from three localities near Kalumburu in the eastern Kimberley Region of

Western Australia. This record extends the known eastward distribution of

this butterfly by approximately 250 km.

Acknowledgments

I thank S.J. Johnson for his initial thoughts on the specimen and M.F. Braby

(ANIC) for genitalia identification.

Reference

D'APICE, J.W.C. and MILLER, C.G. 1992. The genus Nesolycaena Waterhouse and Turner

(Lepidoptera: Lycaenidae) with a description of a new species. Australian Entomological

Magazine 19: 75-80.

80 Austrralian Entomologist 23 (2) September 1996

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AUSTRALIAN ENTOMOLOGY

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1995 The taxonomy, phylogeny and biogeography of the cicada genus Gymnotympana Stal, 1861 (Homoptera:

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BRABY, M.F. :

1995 Seasonal changes in relative abundance and the spatial distribution of Australian lowland tropical satyrine

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1995 World revision of the genus Encyrtoscelio Dodd (Hymenoptera: Scelionidae). Invert. Taxon. 9: 1021-1045.

CLARK, S. and GREENSLADE, P.

1996 Review of the Tasmanian Hanseniella Bagnall (Symphyla: Scutigerellidae). Invert. Taxon. 10: 189-212.

DANGERFIELD, P.C. and AUSTIN, A.D.

1995 Revision of the Australasian species of the Cardiochilinae (Hymenoptera: Braconidae). Invert. Taxon. 9: 387-

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1995 The Richmond birdwing butterfly: what's in a name? ANIC News 7: 12-15.

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1995 Phylogeny of the Ensifera (Orthoptera): a hypothesis supporting multiple origins of acoustical signalling,

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1996 When an insect is more like a plant. Nature Aust. 25 (4): 30-38.

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1993 A new species of the genus Calodema Laporte & Gory 1838 from Queensland, Australia (Coleoptera:

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ENTOMOLOGICAL NOTICES

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THE AUSTRALIAN

Entomologist

Volume 23, Part 2, 30 September 1996

CONTENTS

ALLWOOD, AJ. and DREW, R.A.I.

Seasonal abundance, distribution, hosts and taxonomic placement of

Dipterophagous daci Drew and Allwood (Strepsiptera: Dipterophagidae). 61

BICKEL, DJ.

Smoke flies (Diptera: Platypezidae) and the Sydney Bushfires. 77

COOMBS, M.

Seasonality of cicadas (Hemiptera) on the northern tablelands of

New South Wales. 55

De BAAR, M. and JOHNSON, SJ.

Notes on the food plant of Deudorix epirus agimar Fruhstorfer

(Lepidoptera: Lycaenidae). 36

JACKSON, R.V.

Lepidoptera breeding records from Apbitonia species (Rhamnaceae)

at Paluma, north Queensland, 75

MEYER, C.E.

Notes on the life history of Nacaduba kurava felsina Waterhouse & Lyell

(Lepidoptera: Lycaenidae). 73

MEYER, C.E.

A new record for Nesolycaena caesia d'Apice & Miller (Lepidoptera: Lycaenidae) f

rom north-eastern Western Australia. 79

SEMMENS, T.D.

Flower visitation by the bumble bee Bombus terrestris (L.) (Hymenoptera: Apidae)

in Tasmania. 33

WILLIAMS, A.A.E. and ATKINS, A.F.

The life history of the Western Australian skipper Mesodina cyanophracta Lower

(Lepidoptera: Hesperiidae) 49

WOOD, G.A., HASENPUSCH, J. and STOREY, R.I.

The life hostory of Phalacrognathus muelleri (Macleay) (Coleoptera: Lucanidae) 37

RECENT LITERATURE

An accumulative bibliography of Australian entomolo 80

ENTOMOLOGICAL NOTICES Inside back cover.

ISSN 1320 6133