THE AUSTRALIAN

ntomologist

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THE ENTOMOLOGICAL SOCIETY OF QUEENSLAND

Volume 25, Part 1, 5 June 1998

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THE AUSTRALIAN ENTOMOLOGIST

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Cover: Anthrax maculata (Diptera: Bombyliidae) described by Macquart in 1846

has been collected commonly throughout eastern Australia, across northern

Australia and south-western W.A. Specimens have been collected flying around

burnt trees and mud wasp nests. Females are a common sight in suburban

Brisbane, patrolling brick walls searching for mud wasp nests. Illustration by

Chris Lambkin, Department of Entomology, University of Queensland.

Australian Entomologist, 1998, 25 (1): 1-6 1

OBSERVATIONS ON THE BIOLOGY OF ARHOPALA WILDEI

MISKIN (LEPIDOPTERA: LYCAENIDAE) AND ITS HOST ANT

POLYRHACHIS QUEENSLANDICA EMERY (HYMENOPTERA:

FORMICIDAE)

Rod Eastwood’ and Allan J. King’

150 Broadwater Terrace, Redland Bay, Qld 4165

?PO Box 1302, GPO Townsville, Qld 4810

Abstract

Observations on the biology of Arhopala wildei Miskin and Polyrhachis queenslandica Emery

are recorded, as well as details of their interactions. Ant assisted cuticle removal during ecdysis

of A. wildei larvae is recorded for the first time, as well as pupal stridulation and pupal

oscillation (vibration) in A. wildei.

Introduction

The myrmecophagous early stages of the lycaenid butterfly, Arhopala wildei

Miskin, have been described recently by King and Ring (1996). The

butterfly deposits its eggs near the nest entrance of the arboreal ant

Polyrhachis queenslandica Emery and the first instar larvae are carried into

the nest by the ants. The larvae then remain in the nest where they feed on

the ants’ brood. Additional behavioural observations reported in this paper

were made independently by the present authors at the same location near

Waughs Pocket, north of Innisfail and at other sites in northern Queensland.

Observations

Adult butterfly behaviour

Males of A. wildei are active in early morning sunshine (c >0600h EST),

often flying close to the ground in clearings whilst “dogfighting” with other

males. During the early part of the day males often search the foliage of vine

forest trees, especially near P. queenslandica nests, possibly in search of

freshly emerged females. They later retreat into the foliage and are seen only

occasionally. Pairs in copula were observed on exposed foliage in mid-

morning (c 0900h EST), adjacent to areas of male activity. When disturbed,

mating pairs drift to a lower perch, seemingly unable to sustain flight. The

mating ritual was not observed.

Females are seen most commonly investigating the foliage between mid-

morning (0900h) and mid-afternoon (1500h) and have been observed

depositing eggs between these times. Whilst searching for egg laying sites,

females adopt a slow flight pattern with much fluttering in and around the

foliage. On occasion, they will fly quickly out of the foliage with a short

burst of fast circling flight — apparently after being disturbed. Some older

females remain in the vicinity of P. queenslandica nests for extended periods

(3-4 hrs), occasionally flying off and returning to rest nearby.

Adult butterflies were not seen to visit any flowers in the study area.

However, they were seen in the company of other Arhopala Boisduval

2 Australian Entomologist, 1998, 25 (1)

species, A. micale amphis Waterhouse and A. madytus Fruhstorfer and other

insects (wasps and ants), feeding on exudates at the leaf petiole junction of

the large-leafed vine Merremia peltata (L.) Merr. (Convolvulaceae).

The ant nest

Nests of P. queenslandica ranged in size from very small (c 1 cm’),

containing just a Queen, sometimes with a few early stages, up to large nests

(>100 cm’) with more than 100 adult ants and a similar number of immature

stages. Most nests were around 30 cm’ and contained 40-50 mature ants. A

typical P. queenslandica nest is constructed of two, or sometimes three

overlapping leaves sealed along the sides with what appears to be masticated

bark. The inside of the nest has no compartments and all surfaces are

covered by a thin layer of light brown silk. A dead pupa of A. wildei in one

P. queenslandica nest was likewise webbed over. Two or three tube-like

access holes of 3 mm diameter are built into the webbing at opposite ends of

the nest. During the day, guard ants can be seen in these access holes.

P. queenslandica also nest opportunistically. One group nested in a cavity

under a plastic table using nesting material removed from a damaged nest. In

the wild they have been found nesting in an abandoned wasp nest (S.

Robson, pers. comm.).

Nest sites for P. queenslandica varied in height from 30 cm above ground to

high in the intermediate canopy in protected positions. Most of the nests

were situated 3-5 m above the ground in the overhanging lower canopy of a

forest boundary. This part of the canopy is protected from strong winds and

is also where most of the A. wildei females were observed to fly. Ant nests

may be up to 30 m above the ground as some A. wildei females were seen

investigating foliage at this level. P. queenslandica nests were found on both

windward and leeward sides of forest boundaries, in foliage with and without

green tree ants Oecophylla smaragdina (Fabricius) and other Polyrhachis F.

Smith species.

P. queenslandica nests are very clean inside. Remains of ant eggs, larvae

and pupae after consumption by A. wildei larvae are ejected from the nest by

the ants. Dead ants and A. wildei larval frass also are ejected. However,

some nests contained a number of fluffy, white, papery objects of various

sizes which proved to be the masticated exuviae of A.wildei larvae. No

empty A. wildei pupal cases were found in an active nest and it would appear

that the ants destroy or eject empty butterfly pupal shells.

Ant behaviour

When disturbed, P. queenslandica adopt a defensive threatening posture.

They curl their gaster forward under the mesosoma, striking it repeatedly on

the substrate for one to two seconds, producing an audible rattling or

drumming sound. In daylight hours, the ants are reluctant to leave the nest,

even when disturbed. If the nest is breached, workers will form a line, face

Australian Entomologist, 1998, 25 (1) 3

the exposed breach in the defensive posture and spray fluid from the gaster at

any intruder. Observations made on a number of disturbed nests showed that

nests suffering minor damage were quickly repaired the same night, while the

majority of those that had sustained major damage were found abandoned the

next day.

When two ant nests taken from about 50 m apart were placed in the same

container, these nests merged into one with the expulsion of one queen and

three worker attendants. The exiled queen was later killed by two other

workers and the three attendants returned to the new single nest. On other

occasions, ant brood from one nest was placed in a container housing another

ant nest taken from a different location. The introduced ant brood was

picked up by the other ants and placed inside the nest. P. queenslandica

from different nests were not aggressive to one another as is the case with

many other species of ants (Hdlldobler and Wilson 1990). A. wildei larvae

transferred from one nest to another were similarly accepted without

hesitation.

Butterfly oviposition

A. wildei eggs are laid singly or in small groups of two to four on the ant nest

material, usually near the tubular access holes. The host ants were not seen

to attack females laying eggs. A small number of P. queenslandica nests,

with and without A. wildei eggs attached, were found to be abandoned and

one abandoned ant nest was found to contain an empty pupal shell of A.

wildei. Nests without eggs attached were occasionally found to contain one

or more semi-mature A. wildei larvae. More often there were fewer butterfly

larvae in the ants’ nest than hatched eggs on the outside. The majority of

nests with butterfly eggs were around 20-30 cm’ and had 3-5 eggs attached.

Some of the large-leafed evergreen trees in which larger nests were found,

hold their foliage for at least two years and it is possible that these nests have

supported continuous generations of A. wildei over that period of time. Only

one intact A. wildei egg was found with a hole cut in the side, indicating that

egg parasitoids may be present.

A. wildei larvae in the ant nest

In similar fashion to another myrmecophage, Acrodipsas illidgei Waterhouse

& Lyell (Samson 1989), freshly emerged first instar A. wildei larvae are

carried into the nest by the ants and therein remain very difficult to detect.

Eggshells are neither eaten by the freshly emerged larvae nor removed by the

ants and oophagy was not observed. A. wildei larvae of less than 2 mm in

length were seen clinging to the ants’ silken pupal cases or attached to ant

eggs in the brood batch. Ant eggs and larvae appear to be coated with a thin,

viscid layer. This substance enables the ant eggs and larvae to adhere to each

other and to the nest wall, and facilitates the movement of batches of brood

by individual ants. Small A. wildei larvae also adhere to the ant brood and

are likewise moved by the ants. In the event of a major disturbance the ants

4 Australian Entomologist, 1998, 25 (1)

pick up the ant brood, with lycaenid larva attached, and retreat to the darker

corners of the nest. Ants will often pick up A. wildei larvae instead of their

own brood.

The host ants do not show any aggressive behaviour towards any instar

larvae of A. wildei. A. wildei larvae have been observed consuming P.

queenslandica eggs and larvae and there is circumstantial evidence that they

also eat ant pupae. Trophallactic feeding was not observed between the

butterfly larvae and adult ants. However, it may occur since the. ants. seem to

treat A. wildei larvae, in all other respects, like their own. Butterfly larvae

are regularly attended by the ants with particular attention being paid to the

Newcomer’s organ (NO). Exudates from this organ collect in the concave

anal depression of the A. wildei larva and often, when approached by an ant,

the larva will raise its posterior end to a vertical position, facilitating direct

access to the NO by the ant. Similar behaviour has been observed in

Acrodipsas illidgei by Samson (1989). The queen ant also attends the NO of

A. wildei.

A. wildei larval ecdysis

Two attendant P. queenslandica were observed to carefully remove the

moulting cuticle from a second instar A. wildei larva. The action was

performed at first with both ants pulling and then with one ant holding the

larva in its mandibles, while the other ant slowly peeled back the loose

exuvium. This was followed by a lengthy session of meticulous grooming

with the larva being turned over several times, apparently under duress. The

exuvium was then picked up by one of the attendant ants and thoroughly

masticated before being dropped. It was not ejected from the nest. This

entire process was performed by the same two ants without intervention from

others nearby.

Pupal stridulation and oscillation (vibration)

A. wildei pupae stridulate but, unlike other lycaenids that stridulate with

intermittent bursts of “ticks” or “burrs”, they emit a prolonged “burrr” that

may last for two to three seconds. This was particularly noticeable after a

pupa had been removed from and subsequently reintroduced to an ant nest.

The pupa made no noise until it was “attended” by the ants. This

phenomenon of pupal stridulation being immediately induced by contact with

an attendant ant also occurs in other species of lycaenids, e.g. Jalmenus

evagoras (Donovan) (RE, unpublished observation).

A. wildei pupa were also observed to oscillate. This oscillation consisted of a

rapid dorso-ventral movement of the anterior end of the pupa. It was not

determined if the pupal oscillation coincided with the sound production but

they appeared independent. Interestingly; the frequency of the pupal

oscillations also appeared to be the same as the frequency with which the

host ants tapped on the substrate when alarmed.

Australian Entomologist, 1998, 25 (1) 5

Discussion

Adults of A. wildei dramatically vary in size with wingspans of males and

females ranging from 25-41 mm. The smallest adults may result from semi-

mature larvae that had exhausted their available food supply or were left

behind when a nest was abandoned by the ants, forcing the lycaenid larvae to

pupate early. There is no evidence to indicate that larger A. wildei larvae

migrate to new ant nests as is the case with another arboreal

myrmecophagous lycaenid, Liphyra brassolis Westwood (Dodd 1902, J.

Young pers. comm.). Smaller A. wildei larvae may be carried by P.

queenslandica to a new site should the ant nest be destroyed or abandoned.

However, larger larvae are very vulnerable and desiccate quickly outside the

ant nest and may not be able to survive an extended journey.

Another measure that may be used by the butterfly to ensure an adequate

food supply is cannibalism. In smaller ant nests or under adverse conditions

A. wildei larvae seem to be able to regulate their numbers, and hence the

available food supply, by this method. This hypothesis is supported by

circumstantial evidence, including the fact that some smaller P.

queenslandica ant nests were found to contain only one lycaenid larva yet

had 2 or 3 recently eclosed eggs attached.

A very similar behaviour to the ant-assisted cuticle removal was recorded by

Brewster (1913), where two Polyrhachis ammon (Fabricius) ants assisted the

eclosion of a winged ant from its pupa. After being removed from the pupa

the two ants continued to assist the imago “...one holding while the other

pulled the wings clear of their sticky covering.” Traniello (1982) and

Hölldobler & Wilson (1990) also describe the care that ants bestow on their

eggs, larvae and pupae including that of assisting the larval molt by licking

the ecdysial skin free. Eclosion of an A. wildei adult in the presence of ants

was not observed.

Pupal noise, produced by hammering on the substrate, has been recorded in

six species of lycaenids (Downey 1966) and he suggested that the function

was to frighten away small predators. However, DeVries (1990) suggested

that lycaenid pupal noises mimic vibrations used by the ants for

communication. It is possible that the pupal oscillation of A. wildei is a

specialised form of communicative stridulation or alarm response similar to

that of the P. queenslandica ants. l

Pierce et al. (1987) deduced that significant extra numbers of new workers of

Iridomyrmex anceps (Roger) can be produced by ants that are tending

lycaenids and Nash (1989) was able to provide direct evidence that ant

colonies exhibited significantly higher growth rates when they were able to

tend lycaenid larvae. It would be reasonable to assume that larvae of A.

wildei can similarly stimulate P. queenslandica. In addition, the NO of late

instar larvae of A. wildei is proportionally larger than those observed on other

myrmecophilous lycaenids, such as J. evagoras (Kitching 1983). If it is the

6 Australian Entomologist, 1998, 25 (1)

NO that produces the compounds that stimulate fecundity in the attendant

ants, then this organ would be a very valuable asset to the A. wildei larva and

may account for its increased size and prominence. Of course it is entirely

possible that the exudates from the A. wildei organs are simply to ‘bribe’ the

ants to prevent predation (Malicky 1970), while the larva consume the ants’

brood. Whatever the purpose of the exudates, they are evidently very

attractive to the ants. The exudates are also persistent, apparently remaining

on the moulted A. wildei exuviae in sufficient amounts so that the ants are

reluctant to eject these “foreign” objects from the nest.

Acknowledgments

We gratefully acknowledge Lance and Lyn Vievers for permission to study

and film A. wildei on their property at Waughs Pocket. Thanks also to Ann

Fraser for helpful comments on an earlier draft of this paper and to Rupert

Barrington and Alan Hayward (both BBC London).

References

BREWSTER, M.N. 1913. Observations on ants. Australian Naturalist 2: 178-179.

DEVRIES, P.J. 1990. Enhancement of symbioses between butterfly caterpillars and ants by

vibrational communication. Science 248: 1104-1106.

DODD, F.P. 1902. Contributions to the life-history of Liphyra brassolis, Westw. The

Entomologist 35: 184-188.

DOWNEY, J.C. 1966. Sound production in pupae of Lycaenidae. Journal of the

Lepidopterists’ Society 20(3): 129-155.

HOLLDOBLER, B and WILSON, E.O. 1990. The ants. Belknap Press of Harvard University

Press: Cambridge; 732pp.

KING, A.J. and RING, L.R. 1996. The life history of Arhopala wildei wildei Miskin

(Lycaenidae). Australian Entomologist 23(4): 117-120.

KITCHING, R. 1983. Myrmecophilous organs of the larva and pupa of the lycaenid butterfly

Jalmenus evagoras (Donovan). Journal of Natural History 17: 471-481.

MALICKY, H. 1970. New aspects on the association between lycaenid larvae (Lycaenidae)

and ants (Formicidae, Hymenoptera). Journal of the Lepidopterists’ Society 24: 190-202.

NASH, D. 1989. Cost-benefit analysis of a mutualism between lycaenid butterflies and ants.

Dissertation. University of Oxford, Oxford, UK.

PIERCE, N.E., KITCHING, R.L., BUCKLEY, R.C., TAYLOR, M.F.J. and BENBOW, K.F.

1987. The costs and benefits of cooperation between the Australian lycaenid butterfly,

Jalmenus evagoras, and its attendant ants. Behavioral Ecology and Sociobiology 21: 237-248.

SAMSON, P. 1989. Morphology and biology of Acrodipsas illidgei (Waterhouse and Lyell), a

myrmecophagous lycaenid (Lepidoptera: Lycaenidae: Theclinae). Journal of the Australian

Entomological Society 28: 161-168.

TRANIELLO, J.F.A. 1982. Population structure and social organization in the primitive ant

Amblyopone pallipes (Hymenoptera: Formicidae). Psyche 89(1-2): 65-80.

Australian Entomologist, 1998, 25 (1): 7-12 7

A NEW SPECIES OF TRAPEZITES HUBNER (LEPIDOPTERA:

HESPERIIDAE) FROM WESTERN AUSTRALIA

Andrew A.E. Williams‘, Matthew R. Williams’ and R.W. Hay’

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

PO Box 51, Wanneroo, WA 6065

*Department of Conservation and Land Management, 50 Hayman Road, Como, WA 6152

’8 Klem Avenue, Manning, WA 6152

Abstract

Trapezites atkinsi sp. nov. is described from south-western Western Australia and, on present

knowledge, is-limited to a small area of coastal heathland at Windy Harbour. The pupal stage

is described and illustrated. The larval foodplant is Acanthocarpus preissii Lehm.

(Dasypogonaceae).

Introduction

In reviewing the Western Australian species of Trapezites Hiibner, Mayo and

Atkins (1992) drew attention to a form from Windy Harbour, which they

tentatively assigned to the T. sciron complex. Specimens of this skipper

were first collected by David Yeates on 11 November 1989; subsequently

further specimens were taken in 1989, 1990 and 1995. Examination of this

additional material confirmed earlier suspicions that the original specimens

represented a previously unrecognised species of Trapezites. T. atkinsi sp.

nov. is closely allied to three other Western Australian species of Trapezites:

T. sciron Waterhouse & Lyell, T. argenteoornatus (Hewitson) and T.

waterhousei Mayo & Atkins.

Abbreviations

The following abbreviations refer to institutional and private collections:

AA = Andrew Atkins Collection, Newcastle; RWH = R. W. Hay Collection,

Perth; HHB = H. H. Bollam Collection, Chittering; WADA = Western

Australian Department of Agriculture Collection, Perth; CALM =

Department of Conservation and Land Management Collection, Perth;

WAM = Western Australian Museum, Perth.

Trapezites atkinsi sp. nov.

(Figs 1-9)

Types. WESTERN AUSTRALIA: Holotype d', Windy Harbour, 29.x.1995,

low coastal heath above cliffs with Acanthocarpus, 34°50’13”S

116°00’49”E., A.A.E. Williams, Reg. No. WAM 1998/0013 (in WAM).

Paratypes (11 0”, 8 2): 1 9, Windy Harbour, 14.xi.1995, ex pupa in shelter on

Acanthocarpus preissii, 34°50’13”S 116°00’49”E., A.A.E. Williams, Reg.

No. WAM 1998/0014 (in WAM); 1 2, Windy Harbour, 8.xi.1995, ex pupa in

shelter on Acanthocarpus preissii, 34°50°13”S 116°00’49”E., A.A.E.

Williams; 1 0’, Windy Harbour, 8.xi.1990, M.R. Williams; 1 0%, Windy

Harbour, 29.x.1995, M.R. Williams; 2 9, Windy Hbr., 34°50’13”S

116°00’49”E., ex pupa, 2 & 7.x1.1995, M.R. Williams (all in CALM); 1 ©’,

8 Australian Entomologist, 1998, 25 (1)

Windy Harbour, 26.xi.89, R.H.; 1 d, Windy Harbour, 8.xi.1990, M.R.

Williams (RWH); 3 0’, Windy Harbour, Pt D’Entrecasteaux, 19.xi.1989, D.

Yeates, each with an additional label, Agriculture (Dept) Western Australia,

with an identifying database number 41347, 41348 or 41349; 1 9, same data

but number 41350, with genitalia in attached vial; 2 0’, Windy Harbour,

24.xi.1989, H. Bollam, both with additional labels as above but numbered

41345 and 41346, both with genitalia in attached vials (all in WADA); 1 CO’,

Windy Harbour form, xi.1989, H. Bollam; 1 0’, Windy Harbour, 8.xi.1990,

M.R. Williams; 1 9, Windy Harbour, 29.x.1995, reared ex pupa, A. Atkins, 1

9, Windy Harbour, 1.xi.1995, reared ex pupa, A. Atkins; 1 9, Windy

Harbour, 6.xi.1995, reared ex pupa, A. Atkins (all in AA).

Male (Fig. 4). Head dark brown with hair-tufts of light brown, beneath

greyish-cream; antennal shaft dark brown above, segmented with yellowish

scales, greyish-cream beneath with darker segments, club dark brown to

black, nudum 14-15 segmented black; labial palpus black above, cream

beneath, third segment short and black. Thorax above black with brown hair

scales, posterior with long yellowish-grey hair-scales; legs pale brown, hind

leg often with two pairs of spurs. Abdomen brown, with distal edges of

segments pale yellowish-brown, posterior hair-tuft pale brown, beneath

yellowish-grey. Forewing length 14 mm, with costa almost straight, apex

pointed, termen almost straight; above centrally mid-brown to dark brown

with pale greyish yellow hairs on termen, base and central (median) area with

yellowish grey hairs, three to four subapical pale yellow spots in area

between R3, R4, R5 and M1 diagonally placed across wing, a yellowish cell

spot and two yellowish-orange spots, one between M3 and CuAl and a

slightly larger spot between CuAl and CuA2, two confluent yellowish-

orange spots forming an arc or crescent in median area between CuA2 and

1A+2A, sometimes reduced or nearly absent; cilia pale grey-brown; beneath,

costa, apex, inner margin to tornus and small area above median spots

yellowish-grey, distad of which are two dark spots, median area dark brown

to black, all spots as above but slightly paler; cilia cream with slightly darker

chequering. Hindwing slightly rounded, above dark brown, yellowish hairs

arising from base to median area and extending along inner margin to

subtornal area, a large central yellow-orange patch of scales; cilia pale

greyish-yellow; underside pale yellowish-grey, basal area greyish-brown to

grey, central patch as above but pale yellow, six subterminal and three

Figs 1-9. Pupal stage and adults of Trapezites atkinsi sp. nov. (1) frons of pupa; (2)

lateral and dorsal view of pupa; (3) pupal setae; (4) adult male (left upperside, right

underside); (5) adult female (left upperside, right underside); (6) female genitalia

(dissected from specimen #41350 in WADA collection); (7) male genitalia including

outside left valva (dissected from specimen labelled Windy Harbour form, xi.1989,

H. Bollam, in AA collection); (8) uncus (ventral); (9) posterior section of outside

right valva. Scale bars: Figs 1-2 = 5 mm; Fig 3 = 0.5 mm; Figs 4-5 = 10 mm; Fig 6 =

1 mm; Figs 7-9 = 2 mm.

Australian Entomologist, 1998, 25 (1)

10 Australian Entomologist, 1998, 25 (1)

median black spots centred with silvery-white scales (the largest subterminal

spot being near tornus); cilia cream with slightly darker chequering.

Genitalia (Figs 7-9). Uncus/tegumen broad with uncus (Fig. 8) tapered,

rounded and blunt, slightly toothed at ‘T’-shaped tip, gnathos broadly

rounded and moderately toothed distally, dorso-lateral flanges short, simple

and distally rounded, directed slightly posterior; saccus short, beak-shaped,

connected to a simple, broadly curved vinculum; left valva broad, toothed

and sclerotized posterio-dorsally, harpe, broad, slightly dentate and short

sacculus directed dorsally, ampulla rounded and blunt, right valva slightly

more dentate and toothed; aedeagus short and broad, posterior tip (postzone)

expanded, semi-cupped shape; juxta simple and saddle-shaped.

Female (Fig. 5). Similar to male, but forewing and hindwing with apex and

termen more rounded, spots on forewing larger, with the two confluent spots

forming a crescent between CuA2 and 1A+2A prominent. Spots on both

wing surfaces larger and more prominent. Forewing length 15 mm.

Genitalia (Fig. 6). Papilla analis broad and triangular, apophysis long,

straight and slender; sterigma plates with lamella postvaginalis elliptical,

long, thin and steeply ‘V’-shaped with deeply rounded central groove,

lamella antevaginalis simple, broadly ‘U’-shaped and slightly sclerotized;

caudal chamber moderately broad and short; ductus bursae short and

spreading; corpus bursae broadly ovoid with creased and slightly sclerotized

base and a short, spherical accessory pouch attached by a short, narrow neck.

Distrbution

Trapezites atkinsi is known only from a small area of coastal heathland at

Windy Harbour, Point D’Entrecasteaux, in south-western Western Australia.

Etymology

The species is named in honour of Mr Andrew Atkins, in recognition of his

contributions to Australian entomology over many years and particularly his

contribution to the study of Hesperiidae.

Variation

There is slight variation in the size of the median spot along the inner margin

of the forewing; generally this is distinctively crescent-shaped. In some

specimens the underside of both wings is more evenly pale yellowish-brown.

There is also slight variation in the size of the spots on the underside of the

hindwing.

Life History

Foodplant. Acanthocarpus preissii Lehm. (Dasypogonaceae).

Pupa (Fig. 2). Length 18 mm; greyish-brown covered with a prominent

mottled dark brown maculation and branched setae (Fig. 3); pupal cap

(operculum) (Fig. 1) rough, sclerotized and subelliptical to quadrate, lightly

Australian Entomologist, 1998, 25 (1) 1]

covered with branched setae and mottled maculation, upper area extended to

quadriform ‘turret’, lower area strongly dentate; cremaster short, tapered and

decurved with rounded tip.

Discussion

T. atkinsi resembles all other Western Australian species of Trapezites in

wing-shape, general pattern and colour of both upperside and underside. It is

most similar to T. argenteoornatus in having prominent yellowish-orange

maculation, especially that of the upperside of the hindwing. However, T.

atkinsi is somewhat larger, the wings lack prominent chequered cilia, and the

silver spots on the underside of the hindwing are smaller and on a yellowish-

grey ground.

The broadly rounded valvae tips, ampulla, broadly rounded gnathos and

dorso-lateral processes of the uncus in the genitalia distinguish males of this

species from other Western Australian Trapezites species (Mayo and Atkins

1992, Andrew Atkins, pers. comm.). Female genitalia of T. atkinsi are

differentiated by the very long, flat ovoid extensions to the lamella post

vaginalis plate and a broad ‘U’-shaped lamella antevaginalis plate. In profile

the genitalia of both sexes appear closer to T. sciron eremicola Burns than to

T. argenteoornatus.

The pupa of T. atkinsi is strongly mottled and resembles that of T.

argenteoornatus, but the operculum is higher and dentate at the base and has

a distinctive raised dorsal area. Larval and pupal shelters are similar to those

of T. argenteoornatus. Both species are recorded on the foodplant A. preissii

but T. argenteoornatus has not been recorded south of Bunbury.

Larvae and pupae have been found in shelters on the foodplant or on nearby

vegetation. The foodplant grows along coastal dunes and on the top of

limestone headlands. Adults fly in areas around the foodplants during late

October and November. They “patrol” in sunshine and often settle on

sheltered sandy patches.

The discovery of T. atkinsi brings the number of Trapezites species found in

south-west Western Australia to four, with three of these being restricted to

this area. The phylogeny of this interesting group would no doubt reward

further study, as they all are very closely related. Three (T. waterhousei, T.

atkinsi and T. argenteoornatus) are allopatric, whereas the fourth (T. sciron)

may be sympatric or parapatric. Closer study of this group may shed further

light on the complex variation observed in T. sciron and T. argenteoornatus

by Mayo and Atkins (1992).

Conservation

T. atkinsi is found within D’Entrecasteaux National Park, but its apparently

restricted distribution places this skipper in a vulnerable category. Our

searches south-east of Windy Harbour and in seemingly identical habitats

12 Australian Entomologist, 1998, 25 (1)

between Yallingup and Cape Naturaliste have failed to detect further

populations of the skipper.

Common name

In order to ensure consistency with proposed changes to the common names

of Australian butterflies, we suggest the common name “Heath Ochre” for T.

atkinsi.

Acknowledgments

We are grateful to David Yeates for information and discussion, to Hugh

Bollam for records and material, and to John van Schagen for lending

material held in the Western Australian Department of Agriculture collection.

We thank Andrew Atkins for genitalia dissections and for providing the

illustrations. Valuable comments on the manuscript were received from

Michael Braby.

Reference

MAYO, R. and ATKINS, A. 1992. Anisyntoides Waterhouse (Lepidoptera: Hesperiidae): a

synonym of Trapezites Hiibner, with description of a new species from Western Australia.

Australian Entomological Magazine 19: 81-88.

Australian Entomologist, 1998, 25 (1): 13-22 13

NEW LARVAL FOOD PLANTS FOR AUSTRALIAN HAWK MOTHS

(LEPIDOPTERA: SPHINGIDAE)

M.S. MOULDS

Entomology Department, Australian Museum, 6-8 College St, Sydney, NSW 2000

Abstract

New food plants are listed for 31 species of Australian hawk moths. Fifty-five of these food

plant records are of native plant species and 17 are exotics. Three previously published food

plant records for Cephonodes kingii (W.S. Macleay) and one for Psilogramma menephron

(Cramer) are disputed, together with one for Hopliocnema brachycera (Lower) that originates

from a labelled museum specimen. A brief overview of the diversity of Australian hawk moth

food plants is given.

Introduction

In previous papers (Moulds 1981, 1984) I summarised larval food plants for

Australian hawk moths affecting garden ornamentals and commercial crops.

However, many Australian hawk moth species have larvae that feed on

neither ornamentals nor crops and this paper lists known food plants for a

number of these in addition to unrecorded food plants for species treated

previously. Food plants listed by Common (1990) are not repeated.

The following abbreviations are used when listing names of observers with

several records: AJG, Alan Graham; GS, G. Sankowsky; JO, J. Olive; MSM,

M. S. Moulds.

Nomenclature for sphingid species follows that of Moulds (1996). Plant

names follow Henderson (1997), otherwise Brock (1988) or Bailey et al.

(1976). Exotic plant species are marked by an asterisk (*).

Food plant identifications originating from H. Beste, AJG, GS, D. Lane,

MSM, JO, C. Pratt, P. Valentine and A. Walford-Huggins were provided by

Tony Irvine, CSIRO, Forest Research Station, Atherton or Garry Sankowsky,

Yuruga Nursery, Walkamin. Moth identifications were obtained by rearing

larvae to adults; doubtful adult identifications were confirmed by the author.

New records

Acosmeryx miskini (Murray)

VITACEAE

Cayratia clematidea (F. Muell.) Domin "wild grape" [GS, Tolga, Qld; A.

Hiller, Mt Glorious, Qld]

Agrius godarti W.S. Macleay

CONVOLVULACEAE

*Ipomoea batatas (L.) Lam. "sweet potato" [MSM - Larvae were not found

naturally on this plant but several reared from eggs successfully developed to

adults. ]

14 Australian Entomologist, 1998, 25 (1)

Cephonodes hylas (Linnaeus)

Add to the records listed by Moulds (1984) the following:

RUBIACEAE

Pavetta granitica Bremek. [D. Lane and MSM, Dimbulah; GS, Tolga, Qld]

Psychotria sp. "wild coffee" [Anne Garrett, Rockhampton district, Qld]

Cephonodes kingii (W.S. Macleay)

Add to the records listed by Moulds (1984) and Common (1990) the

following:

RUBIACEAE

Gardenia ochreata F. Muell. "scented gardenia bush" [GS, Chillagoe and

Georgetown, Qld]

Gardenia ovularis F. M. Bailey [GS, Tolga, Qld]

Tarenna sp. [GS, Tolga, Qld]

NOTE: Jones and Elliot (1995) list Cissus, Grevillea and Oreocallis as food

plants. Oreocallis was previously known as Embothrium and now as

Alloxylon. All stem from old inaccurate records and should be disregarded

(see Moulds 1984: 60).

Cephonodes picus (Cramer)

Add to the records listed by Moulds (1984) and Common (1990) the

following:

RUBIACEAE

Aidia racemosa (Cav.) Tirveng. [Anne Garrett, Rockhampton district, Qld]

Coenotes eremophilae (T. P. Lucas)

Add to records listed by Moulds (1984) and Common (1990) the following:

ACANTHACEAE

*Barleria cristata L. "Philippine violet" [E. A. Henty, Kununurra, WA]

RUBIACEAE

*Mussaenda sp. [E. A. Henty, Kununurra, WA]

VERBENACEAE

Stachytarpheta urticifolia (Salisb.) Sims "ratstail" "dark blue snakeweed" [E.

A. Henty, Kununurra, WA]

Vitex glabrata R. Br. [Cliff Meyer, Kununurra, WA]

Coequosa triangularis (Donovan)

Add to the records listed by Moulds (1981) the following:

PROTEACEAE

Grevillea asplenifolia x caleyi 'ivanhoe' [N. Marks, Pennant Hills, NSW]

Australian Entomologist, 1998, 25 (1) 15

Hakea spp. [McMaugh (1985, 1986)]

Persoonia levis (Cav.) Domin [Bruce White, Doyalson, NSW - Rose (1975)

fed P. levis to newly hatched larvae which died. The discovery of larvae

feeding naturally on this plant by White confirms it is a larval food plant.]

Daphnis hypothous (Cramer)

APOCYNACEAE

Alstonia actinophylla (A. Cunn.) K. Schum. "milkwood" [G. Brown and

MSM, Darwin, NT]

Alstonia constricta F. Muell. "bitterbark" [GS, Tolga, Qld]

Alstonia muelleriana Domin "hard milkwood" [MSM, Julatten, Qld]

Alstonia scholaris (L.) R. Br. "milky bean", "milky pine" [MSM, Julatten,

Qld]

RUBIACEAE

*Anthocephalus chinensis (Lam.) A. Rich ex Walp. [Gary Fitt, Darwin, NT -

a tree grown experimentally for commercial use near Darwin; larvae

completely defoliated some plants.]

Nauclea orientalis (L.) L. "Leichhardt tree" [Gary Fitt, Kununurra, WA; P.

Valentine, Townsville, Qld]

Daphnis placida (Walker)

Add to the records listed by Common (1990) the following:

ALANGIACEAE

Alangium villosum subsp. polyosmoides (F. Muell.) Bloemb. [J. Stockard,

Wingham Brush, NSW]

Alangium villosum subsp. tomentosum (F. Muell.) Bloemb. [GS, Burnett R.

and Wallaville, Qld]

APOCYNACEAE

Alstonia actinophylla (A. Cunn. ) K. Schum. [C. Pratt, MSM, Cooktown, Qld;

G. Brown, Darwin, NT]

Alstonia muelleriana Domin "hard milkwood" [MSM, Julatten, Qld]

Alstonia scholaris (L.) R. Br. "milky bean", "milky pine" [MSM, Julatten,

Qld]

*Tabernaemontana divaricata (L.) R. Br. "mock gardenia" [GS, Tolga, Qld]

Ochrosia elliptica Labill. [GS, Tolga, Qld] _

Daphnis protrudens (Felder)

RUBIACEAE

Timonius timon (Spreng.) Merr. var timon [GS, Windsor Tableland, Qld]

16 Australian Entomologist, 1998, 25 (1)

Eupanacra splendens (Rothschild)

Add to the records listed by Moulds (1984) the following:

ARACEAE

*Monstera deliciosa Liebm. "monstera" [Dennis Kitchen, JO, Cairns, Qld]

Rhaphidophora australasica F. M. Bailey [H. Beste, Julatten, Qld]

Hippotion boerhaviae (F.)

RUBIACEAE

Hedyotis sp. [JO, Trinity Beach, Qld]

*Pentas lanceolata (Forssk.) Deflers "pentas" [JO, Trinity Beach, Qld -

larvae were not found naturally on Pentas but readily accepted it when

transferred from Hedyotis]

Hippotion brennus (Stoll)

DILLENIACEAE

Hibbertia scandens (Willd.) Gilg "golden guinea vine

Kuranda, Qld]

RUBIACEAE

*Pentas lanceolata (Forssk.) Deflers "pentas" [JO, Trinity Beach, Qld]

Pogonolobus reticulatus F. Muell. [JO, Trinity Beach, Qld]

mow

snake vine" [JO,

Hippotion rosetta (Swinhoe)

RUBIACEAE

*Pentas lanceolata (Forssk.) Deflers "pentas" [D. Lane, Atherton, Qld]

*Richardia brasiliensis Gomes "Mexican clove" "white eye" [AJG, Yorkeys

‘Knob, Qld]

*Richardia scabra L. [AJG, Yorkeys Knob, Qld]

Spermacoce exserta Benth. [Cliff Meyer, Darwin, NT]

VITACEAE

Cayratia clematidea (F. Muell.) Domin "wild grape" [AJG, Yorkeys Knob,

Qld]

‘Hippotion scrofa (Boisduval)

Add to the records of Moulds (1981, 1984) the following:

ASTERACEAE

*Xanthium spinosum L. "Bathurst burr" [G. Brown, Cootamundra, NSW -

identification uncertain; record requires confirmation]

RUBIACEAE

Hedyotis sp. [JO, Trinity Beach, Qld]

Spermacoce exserta Benth. {Cliff Meyer, Darwin, NT]

Australian Entomologist, 1998, 25 (1) 17

Hopliocnema brachycera (Lower)

MYOPORACEAE

Eremophila willsii F. Muell. (MSM, Wallara Stn, NT)

Eremophila exotrachys Kraenzlin (MSM, Wallara Stn, NT)

CASUARINACEAE

A data label attached to a specimen in the South Australian Museum states

that it was reared from Casuarina. As larvae feed on Eremophila it is most

unlikely that Casuarina is a food plant and the record is here disregarded.

Hyles livornicoides (Lucas)

Add to the records listed in Moulds (1981) the following:

FABACEAE

*?Medicago [sativa] L. "lucerne" [Lower (1897) - a doubtful record that

requires confirmation]

Leucomonia bethia (Kirby)

VERBENACEAE

Clerodendrum floribundum R. Br. "lJollybush" [GS, Walsh River, Mareeba

district, Qld]

Macroglossum alcedo (Boisduval)

RUBIACEAE

Hodgkinsonia frutescens C. T. White [GS, MSM. Tolga, Wongabel S. F. and

Yungaburra, Qld]

Macroglossum dohertyi (Rothschild)

RUBIACEAE

Myrmecodia platytyrea subsp. antoinii (Becc.) C. R. Huxley & Jebb "ant

plant" [D. Lane, Iron Range, Qld]

Macroglossum micaceum (Walker)

RUBIACEAE

Canthium sp. [GS, JO, Forty Mile Scrub, Qld]

Macroglossum prometheus (Boisduval)

RUBIACEAE

Morinda citrifolia L. "cheesefruit" "great morinda" [AJG, Yorkeys Knob,

Qld]

Macroglossum tenebrosa (T.P. Lucas)

RUBIACEAE

Morinda salomonensis Engl. [D. Kitching, Kuranda, Qld]

Macroglossum vacillans (Walker)

LOGANIACEAE

18 Australian Entomologist, 1998, 25 (1)

Strychnos lucida R. Br. "strychnine tree" [GS, Myall Ck (=York Downs),

Cape York Pen., Qld]

Meganoton rufescens (Butler)

Add to the records listed in Moulds (1984) the following:

ANNONACEAE

*Annona muricata L. "soursop" [R. Straatman and MSM, Kuranda, Qld]

Psilogramma menephron menephron (Cramer)

Add to the records listed in Moulds (1981, 1984) the following:

BIGNONIACEAE

Deplanchea tetraphylla (R. Br.) F. Muell. "golden bouquet tree" [JO, Trinity

Beach, Qld]

*Radermachera sinica (Hance) Hemsl. [J. McMaugh and C. Cassar, Sydney,

NSW]

CASUARINACEAE

Walker (1856) formalized the name Macrosila casuarinae from a Boisduval

manuscript name; this is now considered a junior synonym of Psilogramma

menephron. Boisduval [1875] gives Casuarina as a larval food plant. A

complete absence of other Australian records for this common moth from

such an abundant plant suggests Boisduval's record is erroneous. Further, I

have been unable to persuade young larvae to feed on Casuarina. However,

Robinson.,(1975) records Psilogramma jordana B-Bkr in Fiji as feeding on

Casuarina nodiflora. Despite this contradiction I believe the evidence

dismissing Casuarina as a food plant for P. menephron outweighs the

likelihood of this ancient record being correct and I here disregard the record

in the absence of further evidence.

OLEACEAE

Jasminum didymum subsp. lineare (R. Br.) P. S. Green [MSM, Townsville

district, Qld]

*Osmanthus fragans (Thunb.) Lour. "fragrant olive" [J. McMaugh, Sydney,

NSW]

VERBENACEAE

*Citharexylum hydalglense [A. B. Rose, Forster, NSW]

Clerodendrum cunninghamii Benth. [GS, Mt Gravatt, Qld]

Clerodendrum tomentosum (Vent.) R. Br. "lolly bush" [C. N. Smithers,

Sydney, NSW]

*Duranta repens L. "golden-dewdrop" "pigeon berry" [A. B. Rose, Forster;

J. McMaugh, Sydney, NSW]

Vitex glabrata R. Br. [Cliff Meyer, Kununurra, WA]

Australian Entomologist, 1998, 25 (1) 19

Vitex trifolia L. [Gordon Jones, Roebourne, WA; J. McMaugh, Sydney,

NSW]

Synoecha marmorata Rothschild & Jordan

MYOPORACEAE

Eremophila mitchellii Benth. "sandalbox", "budda", "false sandalwood",

"bastard sandalwood", "emu bush" [Lucas 1891]

Theretra latreillii (W.S. Macleay)

Add to the records listed in Moulds (1981, 1984) the following:

RUBIACEAE

*Pentas lanceolata (Forssk.) Deflers "pentas" [JO, Cairns, Qld - larvae were

not found on pentas but those transferred from other food plants readily

accepted it.]

VITACEAE

Cissus opaca F. Muell. "pepper vine" [J. Moss, Brisbane, Qld]

Theretra oldenlandiae (F.)

Add to the records listed in Moulds (1981, 1984) the following:

ARACEAE

*Zantedeschia aethiopica (L.) K. Spreng. "arum lily" [MSM, Brisbane, Qld]

RUBIACEAE

Hedyotis sp. [JO, Trinity Beach, Qld]

Theretra queenslandi (T.P. Lucas)

URTICACEAE

Dendrocnide excelsa (Wedd.) Chew "giant stinging tree" "fibrewood" [Clyne

(1980); John Stockard, Wingham, NSW; AJG, Toowoomba, Qld; A. Hiller,

Mt Glorious, Qld]

Dendrocnide moroides (Wedd.) Chew "gimpi gimpi" "gympie" [A. Walford-

Huggins, Tully Falls, Qld]

Dendrocnide photinophylla (Kunth) Chew "shining-leaved stinging tree"

"mulberry-leaved stinging tree" "fibrewood" [GS, Mt Tamborine, Qld; Anne

Garrett, Rockhampton district, Qld.]

Pipturus argenteus (G. Forst.) Wedd. "native mulberry" [GS, Tolga, Qld;

AG, Kuranda, Qld.]

Theretra silhetensis (Walker)

ONAGRACEAE i

*Lugwigia octovalvis (Jacq.) P. H. Raven "water primrose" [J. Moss,

Brisbane, Qld; GS, Eurimbulah, Qld]

RUBIACEAE

Hedyotis sp. [JO, Trinity Beach, Qld]

20 Australian Entomologist, 1998, 25 (1)

*Pentas lanceolata (Forssk.) Deflers "pentas" [JO, Trinity Beach, Qld -

larvae not found naturally on pentas but readily accepted it when transferred

from Hedyotis]

VITACEAE

Cayratia clematidea (F. Muell.) Domin [AJG, Yorkeys Knob, Qld]

Discussion

Forty-three of the 64 Australian hawk moth species now have larval food-

plant records of Australian origin. A further five species have food-plant

records from localities beyond Australia, viz.: Hippotion rosetta (Swinhoe)

on Boreria and Oldenlandia (Holloway 1987); Macroglossum corythus

(Walker) on Strychnos, Guettarda, Morinda and Paederia (Holloway 1987);

M. heliophila (Boisduval) on Morinda and Psychotria (Holloway 1987); M.

insipida (Butler) on Hedyotis, Borreria, Spermacoce and Corchorus

(Holloway 1987); and Theretra nessus (Drury) on Dioscorea, Ipomoea,

Amaranthus, Impatiens, Citrullus, Arachis, Boerhavia, Knoxia, Morinda,

Oldenlandia, Spermacoce, Glossostigma and Camellia (Holloway 1987,

Mackey 1975, Seitz 1928-29). Food-plant records for the remaining 15

Australian hawk moth species are lacking.

Known Australian food plants now total 196 species in a remarkable 122

genera and 43 families. The Rubiaceae (34 spp) has by far the highest

representation followed by Bignoniaceae (15 spp), Vitaceae (15 spp) and

Oleaceae (14 spp). The Rubiaceae support 22 Australian hawk moth species,

more than for any other plant family. In comparison, the Vitaceae support 11

species but the Bignoniaceae only four and the Oleaceae just one.

One third of the known Australian food plants are introduced, 66 species in

all spread across 23 families. Previously I have discussed the polyphagous

nature of some Australian hawk moths (Moulds 1981), notably

Gnathothlibus erotus (Cramer), Hippotion celerio (L.), Psilogramma

menephron, Theretra latreillii and T. oldenlandiae. These five species

account for 49 of the introduced plants and P. menephron alone for 25.

These moths are all wide-ranging species with distributions extending at least

throughout the Indo-Australian region.

The 17 hawk moths endemic to Australia (a quarter of all Australian species)

together feed on 66 species in 21 families but only nine of these plants are

exotics. Coenotes eremophilae is an exception amongst the Australian

endemic hawk moths, in that it has a large food-plant range (27 species in 13

families) accounting for 26 of the 66 species, and 7 of the 21 families above.

However, the plants that it feeds on are in families that are for the most part

closely related, and all but three are Australian natives.

To draw conclusions at this time about the diversity of Australian hawk moth

food plants would be premature in view of the incomplete knowledge

available. However, two trends are worth noting. Firstly, those hawk moths

Australian Entomologist, 1998, 25 (1) 21

with the widest distributions also tend to have the broadest range of food

plants. Hippotion celerio, for example, is a cosmopolitan moth reaching the

British Isles and North America, and from Australia alone it is recorded

feeding on 25 species in 10 families. Among the Australian endemics

Coenotes eremophilae has by far the widest distribution occurring throughout

much of mainland Australia, and it has by far the highest food plant diversity

as noted above. An exception to this trend seems, at first, to be Agrius

convolvuli (L.) which, like H. celerio, is cosmopolitan but has comparatively

few Australian food-plant records. However, throughout its world-wide

distribution its larvae feed on a wide range of plants covering 7 families

(Holloway 1987). In contrast, narrow-range species such as Hopliocnema

brachycera, Synoecha marmorata (T. P. Lucas) and Coequosa triangularis

are each known to feed on plants in just one family.

Secondly, there is a tenuous correlation between the abundance of a species

and its food-plant diversity. Common species tend to have a range of food

plants spanning several families. For example, Psilogramma menephron (38

species in 6 families), Theretra oldenlandiae (25 species in 8 families) and

Gnathothlibus erotus (17 species in 6 families). Less abundant hawk moths

have food plants which tend to be closely related. For example Coequosa

triangularis is known to feed on nine different plants but all within the

Proteaceae. The seven Australian Macroglossum species for which food

plants are known are all uncommon; together they have 10 known food

plants, nine of which are in the Rubiaceae and one in Loganiaceae.

One would expect some kind of correlation between the systematic positions

of hawk moths and their food plants but there is no obvious broad-based

association in this regard. This subject is a complex one and falls beyond the

scope of this paper and is not pursued here.

Acknowledgments

I am sincerely grateful to those who have provided food plant records: Hans

Beste, Graham Brown, C. Cassar, Densey Clyne, Garry Fitt, Jim Frazier,

Anne Garrett, Alan Graham, the late E. A. Henty, Tony Hiller, Gordon Jones,

Dennis Kitchen, David Lane, N. Marks, Judy McMaugh, Cliff Meyer, John

Olive, Clive Pratt, Tony Rose, Garry Sankowsky, John Stockard, Courtenay

Smithers, the late Ray Straatman, Peter Valentine, Allan Walford-Huggins

and Bruce White. The assistance of Tony Irvine and Garry Sankowsky in

providing plant identifications is gratefully acknowledged. I am also

indebted to my wife Barbara for typing the manuscript.

References

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McMAUGH, J. 1986. What garden pest or disease is that? Companion edition. Lansdowne

Press, Sydney/Auckland /London. 302 pp.

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commercial crops in Australia. General and Applied Entomology 13: 69-80.

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ROBINSON, G.S. 1975. Macrolepidoptera of Fiji and Rotumu: a taxonomic and

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ROSE, A.B. 1975. Moths of the families Sphingidae, Notodontidae and Agaristidae observed

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Australian Entomologist, 1998, 25 (1): 23-27 23

PHACONEURA (HOMOPTERA: MEENOPLIDAE)

ATTENDED BY ANTS OF THE GENUS PARATRECHINA

(HYMENOPTERA: FORMICIDAE) IN CAVES

W.F. HUMPHREYS

Western Australian Museum, Francis Street, Perth, WA 6000

Abstract

Cave inhabiting species of Phaconeura Kirkaldy from the Kimberley and Cape Range, Western

Australia, are attended by ants of the genus Paratrechina Motschoulsky, which themselves

probably are especially adapted for cave life.

Introduction

The Australian tropics have been found recently to contain a diverse

troglobitic fauna, in both humid (Howarth 1988) and arid (Humphreys

1993b) regions. One of the more diverse groups of animals found in the

caves is planthoppers (Homoptera) of the fulgoroid families Cixiidae

(especially the genus Solonaima Kirkaldy: Hoch 1988, Hoch and Howarth

1989a, 1989b, 1989c) and Meenoplidae (especially species of the genus

Phaconeura Kirkaldy: Hoch 1990, 1993). By 1989 North Queensland had

recorded the highest concentration of cave-adapted Fulgoroidea in the world

(Hoch and Asche 1989).

Until 1990 only one cave adapted meenoplid was known in Australia,

Phaconeura pluto Fennah from Nambung National Park in Western Australia

(Fennah 1973). Subsequently four new species of Phaconeura were

described from North Queensland caves (Hoch 1990) and another (Hoch

1993) from caves in the Cape Range of Western Australia (Humphreys

1993a, 1993b). Since then several more undescribed cavernicolous species

of Phaconeura (H. Hoch, pers. comm.) have been found in the area of Cape

Range and in the Kimberley. Cave-adapted meenoplids have been found

exclusively in limestone caves in Australia, whereas elsewhere they are only

known from lava tubes (Hoch 1990) in the Canary Islands (Remane and

Hoch 1988) and Western Samoa (Hoch and Asche 1988). Ant-Homoptera

associations are well known (Schaefer 1987, Bourgoin in press) but have so

far been reported only from epigean species.

The distribution of obligatory cave dwelling (troglobitic) apterous species of

Phaconeura, between isolated but neighbouring towers of the Chillagoe

Karst, led Howarth (pers. comm. in Hoch 1990) to presume the species to be

attended by ants in order to explain their dispersal; species of Paratrechina

Motschoulsky occur in these caves (Howarth 1988). Here support is

provided for Howarth's supposition by reporting the attendance of

meenoplids by ants in caves in the Kimberley and Cape Range, Western

Australia.

24 Australian Entomologist, 1998, 25 (1)

Observations

During surveys of cave fauna in northwestern Australia (Humphreys 1993c,

1995) meenoplids were found to occur widely but the observations reported

here are from two main caves in the Kimberley and Cape Range.

Cave KNI-9 is in the Ningbing Range, Kimberley (15°17'S; 128°37'E),

formed in the exposed Devonian reef system. The cave is mainly horizontal

and is ca 136 m long; it is one of many in the southern Ningbing Range.

Cave C-29 is in Cape Range, Carnarvon Basin (22°06'S; 114°O1'E) in an

anticline of Miocene limestones. The cave comprises a 24 m deep chamber

from which two passages extend for 105 m (R. D. Brooks, pers. comm.) and

where the ants and meenoplids were found. The chamber is often dry but

seepages sometimes moisten the cave soil at the extremities of the passages.

In the caves meenoplids are typically found wandering over the substrate or

feeding on fine growing roots of undetermined plants, probably often Ficus,

usually on damp soil covered surfaces. However, at two places in cave KNI-

9, nymphs of a yet undescribed species of Phaconeura (H. Hoch, pers.

comm. 1994) were found in groups on roots, either on the surface or in soil

cavities, with ants in attendance which appeared to groom the nymphs. The

ants became agitated when disturbed and began to collect the nymphs, both

from the surface roots and the underground cells, and transport them away

from the area. Within several minutes after disturbance no nymphs were left

in the area. In Cape Range (C-29) nymphs of Phaconeura sp. were found in

groups on roots, either on the surface or in soil cavities, with ants in

attendance (A. Amarkey, pers. comm.); nymphs cannot be determined to

species but Phaconeura proserpina Hoch is known from nearby caves.

In both areas the ants were species of Paratrechina, a genus that requires full

revision before species-level identification will be possible (S. Shattuck, pers.

comm. 1994). These ants have very small eyes and it "seems likely that this

is one of the few ants known that is especially adapted for cave life" (S.

Shattuck, pers. comm. 1995). Small-eyed species of Paratrechina have also

been collected from Cutta Cutta Cave, N.T., in the same parts of the cave as

Phaconeura sp. indet. (W. Binks, R. D. Brooks and B. Vine, pers. comm.),

although no clear association was reported.

Paratrechina is widely distributed, although much more common in eastern

and northern areas of Australia (S. Shattuck, pers comm. 1994). While

Paratrechina have been considered to be opportunists (Reichel and Andersen

1996), members of the genus are known to attend Hemiptera. The sugarcane

mealybug Saccharicoccus sacchari (Cockerell) (Pseudococcidae), is

consistently attended both above and below ground by ants, including

Paratrechina sp. prob. vaga (Forel), which were observed carrying the

mealybugs underground (Carver et al. 1987). In addition Paratrechina

obscura Mayr was involved in behaviour - removing mummies from nodes -

that has been interpreted as mutualistic (De Barro 1990).

Australian Entomologist, 1998, 25 (1) 25

Microclimate

The meenoplids were found in those parts of the cave that contained root

systems and where the air was nearly saturated with water (Table 1). In KNI-

9 these areas were in high parts of the cave where the warmer air collected.

In C-29 the meenoplids were found at the extremities of the tunnels where

humidity is greatest. The fauna of tropical caves, even in arid zones, is

dependent on high moisture levels (Howarth 1980, Humphreys and Collis

1990, Humphreys 1991, Weinstein 1994).

Table 1. The temperature (°C) and relative humidity (%) associated with cave KNI-9

in May and June 1994. The mean is given with its standard deviation in parentheses.

Temperature and relative humidity were spot measured using a whirling hygrometer

(Brannan, England).

Location n °C %

Outside cave 3 29.2 (1.72) 44 (16)

Inside cave: no meenoplids 26 25.9 (1.76) 86 (15.9)

meenoplids 3 27.3 (0.46) 98 (0.6)

Discussion

The fauna found in the Kimberley caves is not predominantly troglobitic,

unlike that of tropical caves elsewhere in Australia (Howarth 1988,

Humphreys 1993b, 1993c). Only one of 14 species (7%) associated with the

meenoplids was troglomorphic, that is it had morphological adaptations

associate with cave life, namely Blattodea [Nocticola brooksi Roth

(Blattodea: Nocticolidae)]. By contrast, arid Cape Range contains a very rich

cave fauna with rainforest affinities (Humphreys 1993b, 1993c) and, of the

species known from C-29, half (n=8) were highly troglomorphic, namely

Nocticola flabella Roth (Blattodea: Nocticolidae), Ngamarlanguia luisae

Rentz & Su (Orthoptera: Gryllidae: Nemobiinae), Stygiochiropus communis

Humphreys & Shear (Diplopoda: Paradoxosomatidae) and Draculoides vinei

(Harvey) (Arachnida: Schizomida).

Cave restricted animals ultimately feed on photosynthetically derived energy,

except in special cases of chemoautotrophy. The energy is transferred to the

cave largely by water, as fine or gross plant material, by animals, such as bats

and rhaphidophorid cave crickets foraging outside the cave, and by plants by

means of root growth and sap transport. Cave restricted animals may eat the

roots alive or dead, feed on root exudates or directly on the sap, as do

meenoplids. The presence in these caves of a two level assemblage of

terrestrial species utilizing plant roots may be the first step to recognising

more complex root feeding assemblages in caves, as found recently for

26 Australian Entomologist, 1998, 25 (1)

aquatic invertebrates for which rich assemblages are supported by root mats

in caves (Jasinska et al. 1996).

Acknowledgments

It is my pleasure to acknowledge the identifications and comments of

Hannelore Hoch and Steve Shattuck and the invaluable support of Julianne

Waldock, Darren Brooks and Brian Vine. Facilities were made available by

Bob Shackles, Officer in Charge, Frank Wise Institute for Tropical

Agricultural Research in Kununurra. Professor Dr Hannelore Hoch and Dr

M. B. Malipatil identified Emesinae. Field work was funded under the

National Estate Grants Scheme.

References

BOURGOIN, T., in press. Habitat and ant attendance in Hemiptera Auchenorrhyncha, a

phylogenetic test. Memoires du Museum National d'Histoire Naturelle (Zoologie).

CARVER, M., INKERMAN, P.A. and ASHBOLT, N.J. 1987. Anagyrus saccharicola

Timberlake (Hymenoptera: Encyrtidae) and other biota associated with Saccharicoccus

sacchari (Cockerell) (Homoptera: Pseudococcidae) in Australia. Journal of the Australian

Entomological Society 26: 367-368.

DE BARRO, P.J. 1990. Natural enemies and other species associated with Saccharicoccus

sacchari (Cockerell) (Hemiptera: Pseudococcidae) in the Bundaberg area, southeast

Queensland. Journal of the Australian Entomological Society 29: 87-88.

FENNAH, R.G. 1973. ` Three new cavernicolous species of Fulgoroidea (Homoptera) from

Mexico and Western Australia. Proceedings of the Biological Society, Washington 86: 439-

446.

HOCH, H. 1988. Five new epigean species of the Australian planthopper genus Solonaima

Kirkaldy (Homoptera: Fulgoroidea: Cixiidae). The Beagle, Records of the Northern Territory

Museum of Arts and Science 5: 125-133.

HOCH, H. 1990. Cavernicolous Meenoplidae of the genus Phaconeura (Homoptera:

Fulgoroidea) from Australia. Occasional Papers of the Bishop Museum 30: 188-203.

HOCH, H. 1993. A new troglobitic planthopper species (Homoptera: Fulgoroidea:

Meenoplidae) from Western Australia. Records of the Western Australian Museum 16: 393-

398.

HOCH, H. and ASCHE, M. 1988. A new troglobitic meenoplid from a lava tube in Western

Samoa (Homoptera: Fulgoroidea: Meenoplidae). Journal of Natural History 22: 1489-1494.

HOCH, H. and ASCHE, M. 1989. Cave-dwelling planthoppers of Australia (Insecta:

Homoptera: Fulgoroidea). Jn Pearson, L. (ed.). Tropicon Conference, Lake Tinaroo, Far North

Queensland. 27-31 Dec. 1988. 67-75. Australian Speleological Federation, Cairns.

HOCH, H and HOWARTH, F.G. 1989a. Six new cavernicolous cixiid planthoppers in the

genus Solonaima from Australia (Homoptera: Fulgoroidea). Systematic Entomology 14: 377-

402.

HOCH, H. and HOWARTH, F.G. 1989b. Reductive evolutionary trends in two new

cavernicolous species of a new Australian cixiid genus (Homoptera: Fulgoroidea). Systematic

Entomology 14: 179-196.

Australian Entomologist, 1998, 25 (1) 27

HOCH, H. and HOWARTH, F.G. 1989c. The evolution of cave-adapted cixiid planthoppers in

volcanic and limestone caves in North Queensland, Australia (Homoptera: Fulgoroidea).

Mémoires de Biospéologie 16:17-24.

HOWARTH, F.G. 1980. The zoogeography of specialised cave animals: a bioclimatic model.

Evolution 34: 394-406.

HOWARTH, F.G. 1988. Environmental ecology of north Queensland caves: or why there are

so many troglobites in Australia. Jn Pearson, L. (ed.) 17th biennial conference, Australian

Speleological Federation Tropicon Conference, Lake Tinaroo, Far North Queensland 27-31

Dec. 1988. 76-84. Australian Speleological Federation, Cairns.

HUMPHREYS, W.F. 1991. Experimental re-establishment of pulse-driven populations in a

terrestrial troglobite community. Journal of Animal Ecology 60: 609-623.

HUMPHREYS, W.F. 1993a. The significance of the subterranean fauna in biogeographical

reconstruction: examples from Cape Range peninsula, Western Australia. Records of the

Western Australian Museum, Supplement 45: 165-192.

HUMPHREYS, W.F. 1993b. Cave fauna in semi-arid tropical Western Australia: a diverse

relict wet-forest litter fauna. Mémoires de Biospéologie 20: 105-110.

HUMPHREYS, W.F. (ed.). 1993c. The biogeography of Cape Range, Western Australia.

Records of the Western Australian Museum, Supplement 45: 1-248.

HUMPHREYS, W.F. 1995. Limestone of the east Kimberley, Western Australia - karst and

cave fauna. Report to the Australian Heritage Commission and the Western Australian

Heritage Committee. 190 pp.+ xix. Unpublished.

HUMPHREYS, W.F. and COLLIS, G. 1990. Water loss and respiration of cave arthropods

from Cape Range, Western Australia. Comparative Biochemistry and Physiology 95A: 101-

107.

JASINSKA, E.J., KNOTT, B. and McCOMB, A.J. 1996. Root mats in ground water: a fauna-

rich habitat. Journal of the North American Benthological Society 15: 508-519.

REICHEL, H. and ANDERSEN, A.N. 1996. The rainforest ant fauna of Australia's Northern

Territory. Australian Journal of Zoology 44: 81-95.

REMANE, R. and HOCH, H. 1988. Cave-dwelling Fulgoroidea (Homoptera

Auchenorrhyncha) from the Canary Islands. Journal of Natural History 22: 403-412.

SCHAEFER, C.W. 1987. The early habitat of the Auchenorrhyncha. In C. Vidano and A.

Arzone (eds). Proceedings of the 6th Auchenorrhyncha Meeting, Turin, Italy, Sept. 1987, 135-

146.

WEINSTEIN, P. 1994. Behavioural ecology of tropical cave cockroaches: preliminary field

studies with evolutionary implications. Journal of the Australian Entomological Society 33:

367-370.

28

Australian Entomologist, 1998, 25 (1)

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Dates: 26 - 29 September 1998 {

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will introduce the Symposia on :

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Australian Entomologist, 1998, 25 (1): 29-31 29

NOTES ON THE DISTRIBUTION OF THE DINOSAUR ANT

NOTHOMYRMECIA MACROPS CLARK (HYMENOPTERA:

FORMICIDAE) IN SOUTH AUSTRALIA

C.H.S. WATTS, A.J. McARTHUR and R. FOSTER

South Australian Museum, North Terrace, Adelaide, SA 5000

Abstract

Nothomyrmecia macrops Clark, the only surviving member of the ant subfamily

Nothomyrmeciinae and known previously from one restricted locality in South Australia and

one site in Western Australia, is recorded from 17 additional localities over a linear distance of

more than 400 km in northern Eyre Peninsula, South Australia. It appears to be a relatively

common nocturnal species, active only on relatively cold nights in regions of mature mallee.

Introduction

Nothomyrmecia macrops Clark is the only surviving member of the ant

subfamily Nothomyrmeciinae. It retains a large number of primitive

morphological and behavioural features (Holldobler and Taylor 1983) and

hence has attracted the common name ‘dinosaur ant’ or ‘living fossil ant’.

First recorded in 1934 from the vicinity of Esperance in Western Australia, it

was not collected again until it was discovered at Poochera on Eyre

Peninsula, South Australia in 1976 (Taylor 1978). Since then searches by

Taylor and others have failed to locate additional colonies (R. W. Taylor,

pers. comm.). The future of this phylogenetically isolated and hence

scientifically important ant was in doubt.

Methods

During the spring and early summer of 1995, with some guidance from Dr R.

W. Taylor, we conducted a search for additional colonies of N. macrops in

the Upper Eyre Peninsula region of South Australia (Fig. 1).

Trials of various survey methods (beating, hand searching, the use of

‘tanglefoot™’, pitfall traps and baiting with honey) were undertaken on the

known populations at Poochera. All of the above methods produced N.

macrops except the use of pitfalls. Beating bushes and baiting tree trunks

with honey proved equally effective. However, in terms of unit effort,

baiting was clearly superior, being a rapid and easy technique capable of

quickly surveying relatively large areas.

It may be indicative of the ant’s behaviour that no specimens were caught in

pitfall traps, although the traps were exposed for two months and were

scattered among the mallee, often only a metre or two from trees on which

the ant was observed climbing on most nights.

As previously demonstrated by Hölldobler and Taylor (1983), N. macrops is

nocturnal and is generally active only at temperatures below about 15°C

(although two individuals were caught on different nights at 20°C).

30 Australian Entomologist, 1998, 25 (1)

Based on our experiences at Poochera we adopted the following survey

method. During the day we located likely patches of mallee scrub. In the

late afternoon up to five sites, usually several kilometres apart, were baited

by smearing honey bait on tree trunks around eye height whilst walking in a

straight line or loop for twenty minutes, resulting in a bait trail of 200-500 m

on 50-80 tree stems. Soon after dusk the bait line was visited and samples of

any ants present collected. In general, sites were surveyed on only one night.

130 135

Fig. 1. Eyre peninsula (South Australia) survey sites for Nothomyrmecia macrops.

Filled squares indicate sites where N. macrops was collected. Shaded area refers to

open scrub woodland as given in Griffin and McCaskill 1986. (Because of the scale

of the map not all sites are distinguished).

Results

Seventy-four separate sites were surveyed between Lake Gilles in the east

and Nundroo in the west. Nothomyrmecia macrops was found at 17 sites in

addition to the known sites at Poochera (Fig. 1). (More specific details of

localities of sites surveyed are available from A. J. McArthur). Negative

results do not, of course, necessarily imply that it was not present. This is

particularly true for a species such as N. macrops which is known to be

inactive on warm nights and to be erratic in activity even at low

temperatures.

Australian Entomologist, 1998, 25 (1) 31

We did not attempt to measure the density of N. macrops, nor the geographic

extent of the individual colonies we located. Our impression was that where

it occurred it was in reasonable numbers, with 10-12 individuals at a bait

station not unusual.

To the best of our ability in the time available, we attempted to describe in

general terms each of the sites visited. Nothomyrmecia macrops appeared to

be associated with sites characterised by the following: ‘old growth’ mallee,

long unburnt with a mixture of tree sizes and at least a few large old trees;

mallee dominated by Eucalyptus oleosa, E. brachycalyx and E. gracilis alone

or more often in combination; loose, friable calcareous soils with a high

‘fines’ content; fairly bare ground with a thin, flat layer of litter and little

understorey and a high diversity of ant species, but no dominant aggressive

species.

Discussion

We found N. macrops to be easily surveyed using a dilute honey bait on cool

(<18°C) nights. Using this technique we found N. macrops to be a relatively

common species in areas of ‘old growth’ patchy mallee with sparse

understorey and a thin litter layer on friable soil, between Lake Gilles and

Penong on Eyre Peninsula, a distance of about 400 km. Within this area it

occurred as a member of several ant assemblages. We expect that the

distribution of the species will eventually be shown to be even greater. We

see no reason to consider the species endangered as long as no further major

clearing of mallee vegetation occurs. Remnant roadside mallee represents a

significant proportion of its remaining habitat. The species occurs in the

Lake Gilles Conservation Park and the Chadinga Conservation Reserve.

Acknowledgments

This work was supported by a grant from the Endangered Species Unit of

Environment Australia. Dr R. W. Taylor provided valuable advice from his

deep knowledge of the behaviour of this ant. Mr Adrian (Bill) Pyke is

thanked for his considerable contribution to the field work. Plant

identifications were done by the South Australian Herbarium. Collected

material is lodged in the South Australian Museum.

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TAYLOR, R.W. 1978. Nothomyrmecia macrops: a living-fossil ant rediscovered. Science

201: 979-984.

32 Australian Entomologist, 1998, 25 (1)

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The Australian Entomologist

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THE AUSTRALIAN

Entomologist

Volume 25, Part 1, 5 June1998

KEX

CONTENTS

EASTWOOD, R. and KING, AJ.

Observations on the biology of Arhopala wildei Miskin (Lepidoptera: Lycaenidae)

-and its host ant Polyrhachis queenslandica Emery (Hymenoptera: Formicidae)

WILLIAMS A.ALE., WILLIAMS, M.R. and HAY, R.W.

A new species of Trapezites Hübner (Lepidoptera: Hesperiidae) from

Western Australia -

MOULDS, M.S.

New larval food plants for Australian hawk moths (Lepidoptera: Sphingidae)

HUMPHREYS, W.F.

Phaconeura (Homoptera: Meenoplidae) attended by ants of the genus

Paratrechina (Hymenoptera: Formicidae) in caves

WATTS, C.H.S. , McARTHUR, AJ. and FOSTER, R.

Notes on the distribution of the dinosaur ant Nothomyrmecia macrops Clark

(Hymenoptera: Formicidae) in South Australia

CONFERENCE NOTICES

Australian Entomological Society Conference and Australasian Applied

Entomological Research Conference

RECENT LITERATURE

An accumulative bibliography of Australian entomology

_ ENTOMOLOGICAL NOTICES Inside back cover.

ISSN 1320 6133

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