THE AUSTRALIAN ENTOMOLOGIST

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Sustaining Associates

Arrest-A-Pest Pty Ltd, Rhone-Poulenc Rural Australia Pty Ltd.

Cover: Ornate false spider mites (Tuckerellidae) are early derivative members of

the economically important Tetranychoidea - spider mites, clover mites, flat

mites, etc.. This species, Tuckerella sp. nr. flabellifer Miller, feeds on the stems

of the introduced weed lantana and on a variety of native trees and shrubs.

Illustration by Juanita Choo, Department of Zoology and Entomology,

University of Queensland.

Australian Entomologist, 1999, 26 (2): 33-35 } f oa oa TA

A NEW SPECIES OF DELIAS HUBNER j

(LEPIDOPTERA: PIERIDAE) _ wee

FROM THE STAR MOUNTAINS, PAPUA NEW GUINEA ___

R.B. LACHLAN a

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

Abstract

Delias akrikensis sp. nov. is described from the south-eastern slopes of Mt Akrik, Star

Mountains, Western Province, Papua New Guinea. It is easily distinguished from D. alepa

Jordan and D. weiskei Ribbe, with which it flies, by wing markings on both the upper and

undersides of both sexes. It appears to be closely related to D. alepa.

Introduction

Three subspecies of Delias alepa Jordan are recognised (Yagishita et al.

1993), all restricted to West Irian: D. a. alepa from Mt Goliath and the Snow

Mountains, D. a. kunupiensis Joicey & Talbot from the Weyland Mountains

and D. a. orthobasis Roepke from the Central Mountains. Two subspecies of

Delias weiskei Ribbe are recognised (Yagishita et al. 1993), both confined to

Papua New Guinea: D. w. weiskei from the Aroa River area and

D. w. sayauriae Okano from the Kerowagi region.

Between 1991 and 1996 six males and a female of a species of Delias

Hübner, with characters similar to both D. alepa and D. weiskei, were

collected by RBL and M. S. Moulds from a single unnamed creek on the

south-eastern slopes of Mt Akrik. Although these specimens are similar in

general appearance and size to D. weiskei and D. alepa, they clearly differ in

several characters, notably the absence of the orange-yellow found on the

underside forewings of D. weiskei and the thin red submarginal line of the

hind wing reaching the costa, unlike D. alepa. The seven specimens taken

were found flying together with D. alepa and D. weiskei. The distinct nature

of these seven individuals and lack of specimens intermediate between them

and D. alepa or D. weiskei clearly support their status as a distinct species.

Delias akrikensis sp. nov.

(Figs 1-4)

Types. PAPUA NEW GUINEA: Holotype &, Mt Akrik, 1700 m, Star Mountains,

Western Province, 22.iii.1994, R.B. Lachlan. Paratypes: 4 00", 1 2, same locality,

l.iv.1991, 3.iv.1994, 2.iv.1994, R.B. Lachlan, 1 O”, same locality, 21.xi.1996, M.S.

Moulds. Holotype in Australian National Insect Collection, CSIRO, Canberra;

paratypes in the collection of RBL.

Description. Male. Basal two-thirds of forewing upperside white to just

beyond cell, its edge only slightly curved between the costa and tornus.

Upperside of hind wing white with very thin black marginal border. Pattern

of underside partly visible from above. Underside of forewing with white

basal part slightly less extensive than the white area above. Diffuse black

scaling on anterior third of cell, most intense basally. Four subapical yellow

spots, the anterior two nearly confluent and together forming a broad

34 Australian Entomologist, 1999, 26 (2)

rounded patch. Hind wing with black ground colour. A small round to

subovate cream discal patch is present between lower edge of discal cell and

vein Rs; its inner edge is diffuse and straight and passes through middle of

discal cell; its outer edge touches a thin red submarginal line which extends

from the costa above the discal patch to vein 1A+2A. The red submarginal

line has a small break at vein Rs. A short red subbasal stripe is present in

space Rs and is distally white. A small red diffuse spot is present between

middle of lower edge of cell and vein 1A+2A. Marginal area between the

red line and outer edge of wing white from costa to middle of space M1.

Length of forewing: 20-22 mm.

Female. Upperside of forewing with the white basal area slightly reduced

and not extended beyond end of discal cell; two small white subapical spots.

Hind wing with wider black border than male, particularly between veins M1

and CuA2. Veins Rs down to 1A+2A covered in black scales distally.

Underside as in the male. Length of forewing: 21 mm.

Etymology. D. akrikensis is named after Mount Akrik, the only known

locality for the species.

Discussion

Although D. akrikensis closely resembles D. weiskei and D. alepa,

D. akrikensis is unique in that both the upper and undersides of the forewing

have a smaller basal white area than D. weiskei and D. alepa. The upperside

of the male is almost identical to the male of D. w. sayuriae from Kerowagi,

central PNG. The underside of the hind wing of D. akrikensis has the thin

red submarginal line extending from the costa to vein 1A+2A, as in

D. weiskei, but this line on D. alepa reaches vein Rs and does not pass

through the cream discal patch that extends to the costal margin. The short

red subbasal stripe in space Rs is clearly distally white in D. akrikensis, to a

lesser degree in D. weiskei and entirely red in D. alepa. On the forewing

underside D. akrikensis lacks the orange-yellow proximal area found in

D. weiskei but the four yellow subapical spots of both these species are very

similar in size and shape. The two dominant subapical spots are similar in

size in all three species, but in D. alepa the minor spots are reduced to a thin

line extending to the margin.

In overall appearance D. akrikensis most resembles D. alepa. On the

upperside of the hind wing of the female of D. akrikensis, veins Rs to 1A+2A

are clearly covered in black scales distally. This was not seen in any female

specimen of D. weiskei or D. alepa collected at the same locality, nor does it

show in the specimens figured in D'Abrera (1990) and Yagishita et al (1993).

It is surprising to note that despite intensive collecting at many different sites

above and below 1700 m over several years in the upper Ok Tedi region,

D. akrikensis was taken only at the type locality.

Australian Entomologist, 1999, 26 (2) 35

Figs 1-4. Delias akrikensis sp. nov. (1) male upperside, (2) male underside,

(3) female upperside, (4) female underside.

Acknowledgments À

Without the support of Ok Tedi Mining Ltd between 1991 and 1996, both

myself and Mr Max Moulds (Australian Museum) could not have carried out

surveys of the region. I also wish to thank Max Moulds for his comments on

the draft manuscript and Mrs Barbara Moulds for typing the manuscript.

References

D'ABRERA, B. 1990. Butterflies of the Australian Region. 3rd ed. Hill House, Melbourne

and London; 416 pp.

TALBOT, G. 1930. A monograph of the pierine genus Delias. Part V. Pp 220-259, pls XL-

XLIII, LVIII, LIX. John Bale, Sons and Danielsson Ltd, London.

YAGISHITA, A., NAKANO, S. and MORITA, S. 1993. An illustrated list of the genus Delias

Hübner of the world. [Text] pp i-xiv; 1-384 (and 4 pp errata); [illustrations] pp i-ix; 1-409; i-vi.

Ed. Yasusuke Nishiyama. Khepera Publishers, Tokyo.

36 Australian Entomologist, 1999, 26 (2)

BOOK REVIEW

Australian Ants: Their Biology and Identification. By S. Shattuck. I-XI + 400 pages +

16 pages of colour plates, hardback. Price $A89.95. CSIRO Entomology Monographs

on Invertebrate Taxonomy Volume 3, December 1998. ISBN 0 643 06032 4.

At last there is a comprehensive book dealing with one of Australia's most diverse,

numerous and ecologically important groups of insects, the ants. Keys to the genera

of Australia's ants are available elsewhere, as are catalogues and checklists of the

fauna and there are a number of regional guides that include considerable information

on the biology of Australian ants. However Steven Shattuck is the first to bring all

this information and much more together within the one book that is comprehensive

both geographically and taxonomically.

The book is basically divided into three sections. The first 22 pages briefly but

adequately cover a range of general information on ant biology, morphology and

classification, discuss ants as pests and environmental monitoring tools and provide

useful notes on collecting methods and specimen preparation. There is also an

overview of the distribution and abundance of ants in Australia which is accompanied

by two informative but washed-out looking maps. There is also a map illustrating

some of the more important place names for those unfamiliar with Australia's

geography, although state borders would have been a useful inclusion.

The meat of the book is contained in the next two sections. Pages 21-57 consist of

excellent keys for the identification of all 103 known genera of Australian ants. A key

to the 11 Australian subfamilies is followed by separate keys to genera of each

subfamily, where necessary. Every couplet is provided with clean, concise line

drawings (except curiously the last couplet of the key to subfamilies), with labelled

arrows referring to specific characters mentioned in the couplets. I spent some

considerable time running specimens through the keys and found them quick and

very easy to use. If difficulties are encountered they normally stem from problems

with specimen preparation although in very rare instances the author has been forced

to use characters that are difficult to see or interpret. What novice ant workers will

welcome most is that a minimum of technical language is used and when it is, the

terms are either clearly illustrated in figures in the morphology section or they are

explained simply in a glossary.

The bulk of the book, pages 58-220, consists of individual treatments of all 103

genera, arranged in alphabetical order within their respective subfamilies. This

section is a tremendous summary of the current knowledge of the entire Australian

ant fauna. Each generic treatment includes notes on identification and a summary of

its biology which also refers to important taxonomic works on the genus. Notes on

distribution and habitat preferences and a checklist of described species, including

synonyms are provided. All accounts (except two) are illustrated by scanning electron

micrographs of the head and mesosoma of a representative species, or more if there is

considerable morphological variation, and a distribution map usually showing

individual collection sites is provided.

Steven Shattuck has done a first rate job and it is difficult to find fault with this

marvellous book. It is well written, richly illustrated, including 16 pages of mostly

good quality colour plates and well designed. This book is an absolute must for

anyone with an interest in Australian ants and should serve as a template for anyone

contemplating dealing with the ant fauna of other regions. The price tag of $89.95

may put it out of reach of students who might well be inspired to take up the study of

ants, but compared with some monographs, it is good value.

C.J. Burwell. Higher Entomology Section, Queensland Museum, PO Box 3300,

South Brisbane 4101.

Australian Entomologist, 1999, 26 (2): 37-44 37

A NEW HAWK MOTH FROM NORTHERN AUSTRALIA

WITH NOTES ON ITS LIFE HISTORY

(LEPIDOPTERA: SPHINGIDAE)

M.S. MOULDS' and D.A. LANE?

‘Australian Museum, 6 College St. Sydney, NSW 2000 (Email: maxm@amsg.austmus. gov.qu)

23 Janda St, Atherton, Qld 4883

Abstract

Psilogramma argos sp. nov. is described, figured and distinguished from the closely allied and

sympatric P. menephron (Cramer). The early stages of P. argos are described and the larva and

pupa figured. The larval foodplant is Gyrocarpus americanus Jacq., family Hernandiaceae.

Introduction

Three species have been recognised in the Indo-Australian genus

Psilogramma Rothschild & Jordan: P. menephron (Cramer), P. increta

(Walker) and P. wannanensis Meng (Bridges 1993). Both P. menephron and

P. increta have been recorded from Australia although the latter record

requires confirmation (Moulds 1996). Moulds and Lachlan (1998) recorded

a fourth species from Papua New Guinea, which remains undescribed.

Another species, described here, is known to occur in Australia, the first

specimens being taken by M.S. and B.J. Moulds in 1973 from north-eastern

Queensland. Subsequently, it was collected by E.D. Edwards from the

Northern Territory and by M.S. and B.J. Moulds from Western Australia;

more recently specimens again have been taken in north-eastern Queensland.

Placement of this new species in Psilogramma complies with generic

diagnoses given by Rothschild and Jordan (1903), D’Abrera [1987] and

Holloway (1987).

In 1997, one of us (DAL) found larvae of an unknown sphingid species near

Chillagoe, north-eastern Queensland and these produced adults of this

undescribed Australian Psilogramma species. Below we name this species

and describe its life history.

The following abbreviations are used: AM, Australian Museum, Sydney;

ANIC, Australian National Insect Collection, CSIRO, Canberra; DAL,

D.A. Lane; MSM, M.S. Moulds. Nomenclature for wing venation and

genital structures follows that of Common (1990).

Psilogramma argos sp. nov.

(Figs 1-7, 9-10, 13-15)

Types. Holotype d, 12 km W of Chillagoe, N. Queensland, 24.iii.1997, bred ex

larva, D.A. Lane (ANIC). Paratypes: QUEENSLAND: 1 O”, Mcllwraith Range,

13°42'S 143°17'E, 350 m, 17-18.i.1988, B. Hacobian (ANIC); 1 O”, Mt White, Coen,

13°58'S 143°11'E, 12.i.1994, E.D. Edwards and P. Zborowski (ANIC); 1 O”, Forty

Mile Scrub, approx 40 miles SSW of Mt Garnet, 9.i.1973, M.S. & B.J. Moulds (AM);

2 OG, Forty Mile Scrub, approx 40 miles SSW of Mt Garnet, 9.i.1973, M.S. & B.J.

Moulds (MSM); 1 O”, Chillagoe, 4.ii.1989, D.A. Lane (DAL); 1 O”, 1 ?, 12 km W of

38 Australian Entomologist, 1999, 26 (2)

Figs 1-3. Psilogramma argos sp. nov., Chillagoe, N. Qld: (1) male, ex pupa,

upperside; (2) same, underside; (3) 4th instar larva.

Australian Entomologist, 1999, 26 (2)

Figs 4-6. Psilogramma argos sp.nov., Chillagoe: (4) Sth instar larva; (5) pupa, lateral

view; (6) pupa, ventral view.

40 Australian Entomologist, 1999, 26 (2)

Chillagoe, 26, 30.iii.1997, bred ex larva, D.A. Lane (DAL); 1 O”, 2 99, 14 km W of

Chillagoe, 2, 6, 7.iv.1997, bred ex larva, D.A. Lane (DAL); 2 99, 15 km W of

Chillagoe 18,20.iii.1997, bred ex larva, D.A. Lane (DAL). NORTHERN

TERRITORY: 6 00", 7 km ESE of Smith Point, Cobourg Pen., 11°09'S 132°11'E,

23.i.1977, E.D. Edwards (ANIC); 1 O”, Black Point, Cobourg Pen., 11°09'S 132°09'E,

8.i.1977, E.D. Edwards (ANIC); 1 9, Darwin, 3-9.xii.1963, J. Sedlacek (MSM).

WESTERN AUSTRALIA: 1 O”, Broome, 7.ix.1978, M.S. & B.J. Moulds (MSM).

Male (Figs 1-2). Ground colour of fresh specimens light grey, in places

almost white, highlights and markings black or near black; worn specimens

off white, almost pale cream, with extensive areas of pale grey, highlights

and markings black to dark brown.

Head grey, palpi with a broad black band from eye to apex, light grey above

this band, white below. Antennae grey to black, darkest below.

Thorax above grey; a black longitudinal band near outer margin of tegulae,

and continued (although sometimes interrupted) along posterior edge of

metathorax forming a broad ‘U’, this band proximally edged with ochre on

the specimen from Broome. Thorax below milky white.

Wingspan 84-94 mm. Fore wing 38-44 mm; apex sharply rounded but not

pointed; termen strongly angled outwards at CuA> so that tornal angle

approaches 90°; above, light grey with black markings as detailed in Fig. 1.

Hind wing above brown to near black around base and on outer two thirds,

remainder (which includes tornus) grey, markings as in Fig. 1. Fore wing

and hind wing below dark brown, grey and off white, marked as in Fig. 2.

Abdomen above grey, lacking a black band along midline although some

specimens have faint remnants of hair-line, laterally black or brown, the

segmental margins marked white. Below milky white.

Male genitalia (Figs 7, 9, 10). Uncal lobes in lateral view claw-like, curved

evenly on ventral margin, gently so on dorsal margin with abrupt down-curve

towards a pointed apex; tapering throughout length but with apical region

tapering to a point from a broad base; apex black or nearly so. Gnathos short

and broad, margin gently curved. Valvae near parallel-sided, distally

rounded; sacculus ill-defined, harpe absent. Aedeagus with subapical

process very broad, flat, short, apically forked into one short and one long

pointed spine. Vinculum with anterior, rod-like extension much longer than

midline of basal triangle.

Female. Similar to male. Female genitalia not dissected; a wild-caught

female is required and we were reluctant to disfigure the single specimen

available.

Diagnosis. The abdomen lacks the prominent black dorsal midline present in

all other Psilogramma species and the male and female fore wing is strongly

curved outwards at CuA, producing a tornal angle near 90°. Adults have an

overall white appearance and the underside of the thorax and abdomen are

always pure milky white. Male genitalia also show marked differences from

Australian Entomologist, 1999, 26 (2) 41

those of P. menephron. The uncal lobes are, in lateral view, more thick-set

(compare Figs 7 and 8), uncal shape in dorsal view is clearly different

(compare Figs 9 and 11), the posterior rod-like extension to the vinculum is

much longer (compare Figs 10 and 12) and the aedeagal subapical process

terminates with one very short and one very long pointed spine (compare

Figs 7 and 8).

Etymology. Named from the Greek word ‘argos’ meaning white and

pertaining to the overall white appearance of this species, more evident than

in any other Psilogramma species.

Distribution. Northern Australia, from Broome in Western Australia, Darwin

and Cobourg Peninsula in the Northern Territory and McIlwraith Range, Mt

White, Chillagoe and Forty Mile Scrub National Park in north-eastern

Queensland.

Figs 7-8. Male genitalia in situ but with right valva removed, lateral view:

(7) Psilogramma argos sp.nov., paratype, Cobourg Peninsula, NT; (8) P. menephron,

Iron Range, N.Qld.

42 Australian Entomologist, 1999, 26 (2)

As the distribution of the larval foodplant includes many localities

throughout northern Australia, P. argos is probably spread widely through

the same region. The small number of specimens in collections most likely

reflects a lack of collecting in suitable areas during the wet season.

There are records of adults for September and from December to April.

Emergence of adults appears to be closely aligned to monsoonal rains, the

first individuals emerging with the onset of the wet season.

13 15

Figs 9-15. (9, 10) uncus in dorsal view, vinculum in ventral view, Psilogramma

argos sp.nov., paratype, Cobourg Peninsula; (11, 12) same, P. menephron, Iron

Range; (13-15) pupa, cremaster, dorsal, lateral and ventral views, P. argos sp.nov.,

Chillagoe.

Australian Entomologist, 1999, 26 (2) 43

Biology

Habitat. In the Chillagoe area, northern Queensland, the species is closely

associated with dry vine scrub patches straddling and lying between

limestone outcrops and ridges that rise to some 30 m above the surrounding

countryside. Such habitat extends from approximately 10 km east to 15 km

west of Chillagoe. The habitat at Forty Mile Scrub National Park is dry vine

scrub on red soil. The larval food plant occurs at both localities.

Foodplant. Gyrocarpus americanus Jacq., “Twirly Whirly Tree” or

“Stinkwood” (Hernandiaceae). Widespread across monsoonal northern

Australia.

Egg. Only hatched eggs observed, 1.5 mm in diameter, laid singly on the

underside of leaves.

Larva (Figs 3-4). First instar unknown, others described below.

Early instars: Similar to last instar. Spiny protuberances on thoracic

segments proportionately larger, pale yellow. Caudal horn nearly straight.

4th instar (Fig. 3): Similar to last instar. Caudal horn straighter, 14 mm long.

Sth instar (Fig. 4): Pale green with numerous pale yellow spots; dorsal and

subdorsal regions with variable coverage of dark brown to purplish blotches

scattered irregularly, some larvae almost devoid of such colouring, others

heavily marked. Seven oblique, pale yellow stripes laterally on abdominal

segments 1-8, each stripe arising anterior of a spiracle, at about level of

spiracle or just below it; these terminating on dorsal surface of abdomen;

posterior-most stripe reaching base of caudal horn. Thoracic segments each

dominated by transverse rows of conical protuberances, pale yellow at base,

becoming light brown at apex, those on segment 2 largest. Head pale green

with darker green vertical band either side of midline, these bands edged pale

yellow along their outer margins. Legs and prolegs pale green. Anal plate

and anal prolegs with many scattered protuberances, similar in colour to

background surface. Spiracles pale yellow. Caudal horn 9-10 mm long,

gently curved backwards, pale green to turquoise at base merging to pale

yellow at apex; adorned with scattered, prominent, brown protuberances,

near conical in shape. Length of larva at maturity approximately 75 mm.

Larval behaviour. The large bell-shaped leaves of Gyrocarpus are often

adorned with numerous brown blotches, together with irregular patterns of

chewed holes of various origins. Larvae rest on the undersides of these large

leaves, often along the midrib, and are remarkably well camouflaged.

In captivity, prior to pupation, mature larvae turned an overall purplish

colour and wandered continuously until placed in separate containers, with a

bedding of potting mix with soil and bark chips to a depth of about 150 mm.

Some larvae burrowed immediately, others did so at various times over a 24

hour period. All burrowed to the bottom of the container and formed a cell

chamber of soil/potting mix lined with silk, in which they pupated.

44 Australian Entomologist, 1999, 26 (2)

As the host plant mostly grows on the limestone outcrops at Chillagoe, larvae

probably utilise the numerous cracks and fissures that run through these

rocky areas. These cracks and fissures often fill with leaves and debris and

may provide a suitable pupation site.

Pupa (Figs S, 6, 13-15). Similar to that of Psilogramma menephron. Light

brown. Haustellum case well developed and about one third length of pupa.

Metathoracic plate broken by deep transverse grooves and pits. Abdominal

segments 9 and 10 lacking deep pits as on segments 1-8. Cremaster (Figs

13-15) in dorsal view near to an equilateral triangle; terminating in a pair of

distally-directed sharp spines, slightly diverging; dorsal surface knarled, no

portion dominating in height. In lateral view near straight and more or less

parallel-sided; tilting upwards from body axis. Ventrally broadly walled

along outer margins, midline sharply grooved.

Distinguished from P. menephron by having abdominal segment 9 lacking

pits; metathoracic plate bearing deep grooves and pits; the cremaster straight

in lateral view, that of menephron down-turned along ventral margin; and the

cremaster depressed along ventral midline while that of menephron is raised.

Larvae collected in February pupated during March and produced adults in

March and April.

Acknowledgments

For helpful discussion we thank Ted Edwards (ANIC) and Jan Kitching

(Natural History Museum, London). We are grateful to Sally Beech for

preparing the line drawings. Ted Edwards kindly provided access to

specimens in the ANIC. Photographs of the larvae and pupae were prepared

by Clifford Frith (Butchers Creek via Malanda, Qld) and those of the adult

moth by Stewart Humphreys (AM); to both we express our thanks. For the

foodplant identification we thank Gary Sankowsky (Tolga, Qld).

References

BRIDGES, C.A. 1993. Catalogue of the family-group, genus-group and species-group names

of the Sphingidae of the world. Privately published by the author; 296 pp.

COMMON, LF.B. 1990. Moths of Australia. Melbourne University Press, Carlton; vi, 535 pp,

32 pls.

D’ABRERA, B. [1987]. Sphingidae Mundi, hawk moths of the world. Based on a checklist by

Alan Hayes and the collection he curated in the British Museum (Natural History). E. W.

Classey, Faringdon; ix, 226 pp.

HOLLOWAY, J.D. 1987. The moths of Borneo: superfamily Bombycoidea: families

Lasiocampidae, Eupterotidae, Bombycidae, Brahmaeidae, Saturniidae, Sphingidae. Malayan

Nature Society and Southdene Sdn. Bhd., Kuala Lumpur; pp 1-199, figs 1-163, pls 1-20.

MOULDS, M.S. 1996. Sphingidae. Jn: Nielsen, E.S., Edwards, E.D. and Rangsi, T.V. (eds),

Checklist of the Lepidoptera of Australia. Monographs on Australian Lepidoptera 4. CSIRO,

Australia; pp 266-270, 365-366.

MOULDS, M.S. and LACHLAN, R.B. 1998. An annotated list of the hawk moths

(Lepidoptera: Sphingidae) of Western Province, Papua New Guinea. Australian Entomologist

25(2): 45-60.

ROTHSCHILD, L.W. and JORDAN, K. 1903. A revision of the lepidopterous family

Sphingidae. Novitates Zoologicae 9 (Suppl.): cxxxv, 972 pp, 67 pls.

Australian Entomologist, 1999, 26 (2): 45-52 45

EVIDENCE FOR UNPALATABILITY IN THE GENUS DELIAS

HUBNER (LEPIDOPTERA: PIERIDAE) AND ITS ROLE IN

MIMETIC ASSEMBLAGES

A.G. ORR

Environmental Sciences, Griffith University, Nathan, Qld 4111

Abstract

Evidence is presented which strongly suggests that butterflies of the pierine genus Delias

Hiibner are highly distasteful or even toxic to vertebrate predators, but may be consumed

without ill effects by invertebrate predators. The role of Delias species as models for Batesian

and Mullerian mimetic assemblages is discussed, with particular reference to the probable

Batesian mimics of the genus Mynes Boisduval. Itis suggested that the inexact resemblance of

Batesian mimics of Delias is due to their fluttering flight, which transmits a distinctive flash

pattern but few fine details of colour and pattern. A similar argument provides an explanation

for the lack of discernible Mullerian assemblages in the genus. The relatively few mimics of

Delias may also be a result of the probable Gondwanan origin of the genus and its present day

preference for temperate or montane climates, which has meant there has been insufficient

contact with potential mimics from the Oriental fauna for many associations to form.

Introduction

The mimetic resemblances of butterflies of the genus Delias Hübner and

other pierids, notably Cepora Billberg and the nymphalid genus Mynes

Boisduval are well known (Dixey 1918). However the true nature of the

relationship has never been clarified. Certain evidence, direct and

circumstantial, suggests that Delias species, almost all of which feed in the

larval stage on mistletoe (Loranthaceae), are distasteful or even toxic to

vertebrates, are aposematically patterned as adults and may serve as the focus

for Batesian and/or Mullerian assemblages throughout their geographic

range. To date however there has been no hard evidence, by way of

chemical analysis or bioassay, that these butterflies are unpalatable and

evidence for the chemical basis of toxicity based on the hostplant chemistry

is also ambivalent (Samuelsson 1966, 1969). In this paper I present some

new evidence that Delias species are unpalatable or toxic to vertebrates, but

probably palatable to invertebrates. This is provided by direct observation of

predation and rejection in the field and by a simple experiment. I also review

patterns of apparently Delias-centred mimetic assemblages which occur

throughout the geographic range of the genus. In particular, I examine the

relationship between Delias and Mynes, many species of which appear to be

Batesian mimics of various Delias species

Circumstantial evidence of unpalatability

The larvae of Delias are exposed and gregarious and presumably vulnerable

to predation. While none of the Australian species is especially colourful,

they are not cryptic. Larvae of some Oriental species such as D. henningia

Eschscholtz are strikingly patterned with dark brown and yellow bands.

Pupae are characteristically bright yellow and not concealed. The adults are

slow flying and conspicuous. Moreover, the undersides of most species are

very brightly. coloured, making them extremely conspicuous when at rest.

46 Australian Entomologist, 1999, 26 (2)

The butterflies roost quite openly. Although occasionally I have seen beak

marks on a high proportion of specimens, I have very seldom seen the

characteristic symmetrical damage caused by birds or other vertebrate

predators attempting to take the insect while at rest. This negative evidence

is based on many hundreds of observations on D. nigrina (Fabricius) and

D. argenthona (Fabricius), which are present for most of the year in large

numbers in my garden at Caloundra, Queensland. Similar observations apply

to over 300 specimens of more than 50 species I have collected in Papua

New Guinea and Southeast Asia.

General evidence of rejection by vertebrates

Direct evidence of avian predation is often very difficult to obtain. For

example Robbins (1980), in an extensive investigation of the role of avian

predation in the evolution of “false heads” in lycaenids, did not witness a

single attack in a two year study. It is to be expected that attacks on

distasteful species would be even less frequently observed. I first began

observing Delias seriously in 1974 and have, in the 25 years since, watched

many thousands of individuals. The following observations are the total of

interactions with predators I have witnessed in that time.

In late September 1987 a young Lewin’s honeyeater (Meliphaga lewinii)

fairly recently fledged from a nest in my garden, was observed taking a

D. argenthona male in flight. It held it by the body for about 25 seconds,

then released it intact but badly injured. A few days later a similar

interaction between a spangled drongo (Dicrurus hottentottus) and a male

D. nigrina was observed. As indirect evidence of avoidance, about the same

period I sampled 30 male D. nigrina and 10 male and female D. argenthona.

Of these, nine (23%) had beak marks on their wings, including two

D. argenthona and one D. nigrina which had symmetrical damage indicating

they had been attacked at rest. At that time many newly fledged

insectivorous birds were about. I took another sample of the same size three

weeks later in mid October, by which time none of the specimens showed

any sign of attack suggesting the young birds were rapidly educated to avoid

them. It should be noted that the maximum lifespan of D. nigrina in the wild

at this time of year seldom exceeds two weeks (Orr 1988) hence this sample

would have mostly represented a later brood of butterflies. In March 1978 a

Lewin’s honeyeater, which was feeding on insects at the time, was seen to fly

within 20 cm of a resting D. argenthona, apparently investigate it briefly,

then fly off. In April 1985 little wattlebirds (Anthochaera chrysoptera) were

twice observed to fly swiftly and deliberately at fluttering D. nigrina males,

only to pull out of the attack at the last moment, swerving abruptly when

about 30 cm from the butterfly. In neither case did the butterfly take any

evasive action before or after the aborted attack. In January 1979 spinetailed

swifts (Hirundapus caudacutus) were seen swooping down from a great

height and snatching various butterfly species which were hilltopping on

Gunong Jassar, West Malaysia. Twice I saw an apparent attack on Delias

Australian Entomologist, 1999, 26 (2) 47

pasithoe parthenope (Wallace), where the bird swerved at the last moment,

much as described above.

In an outdoor insectary small populations of D. argenthona and D. nigrina

were maintained for over a year where at night they roosted communally on

the walls. Small garden skinks, which frequently captured and ate small to

medium sized lycaénids and satyrids, including Melanitis leda bankia

(Fabricius), were on three occasions observed to attack roosting Delias,

grasping them by the thorax or abdomen, but releasing them after a few

seconds.

Experimental evidence of the distastefulness or toxicity

In 1996, a magpie (Gymnorhina tibicen) regularly came into my house to

accept scraps of meat, Spodoptera Guenée larvae and various geometrid

larvae from plants believed to be low in noxious chemicals. These were

always eaten enthusiastically. When it was offered mature Delias nigrina

larvae it swallowed the first avidly, then took another somewhat less

willingly, but after about five minutes showed visible signs of distress, flew

to a nearby branch and vomited. It was only after some days and much

coaxing that it returned to feed and, when offered unadulterated mince, ate it

enthusiastically, but when offered pieces of mince mixed with freshly ground

adult D. nigrina or D. argenthona, it briefly tasted them then rejected them.

The Magpie was almost certainly a naïve subject, since its ordinary method

of feeding would not have brought it into contact with either larval or adult

Delias. Clearly this is not statistical proof of unpalatability of Delias, being

based on a sample size of one, but taken in conjunction with other evidence it

does suggest a strong prima facie case for distastefulness in adult and larval

stages.

Further evidence is provided by a simple investigation of the reponses of

wattlebirds (Anthochaera chrysoptera) to Delias larvae presented at a feeding

table to which they had become habituated. When palatable Spodoptera

larvae were offered, they were eaten, whereas the Delias larvae of similar

size and colour were sometimes investigated but never taken. When both

types of larvae were presented only the Spodoptera were taken.

Palatability to invertebrate predadators:

There is no evidence that Delias are unpalatable to invertebrate predators.

Between 1982 and 1988 I maintained captive populations of up to 12

D. nigrina and/or D. argenthona in a large outdoor insectiary at Caloundra.

Both species were regularly taken and eaten by Thomisus Walkenauer sp.

(Araneae: Thomisidae), living in flowerheads. On one occasion an

unidentified preying mantis captured a D. nigrina female, which it consumed

normally and without ill effect. A more critical test was provided by Nephila

Leach sp. (Araneae: Argiopidae), which reject or partially reject any butterfly

feeding on Apocynaceae, presumably on account of sequestered pyrrolizidine

alkaloids (Brown 1984), as well as Aristolochea feeding troidine

48 Australian Entomologist, 1999, 26 (2)

swallowtails. Nephila showed no tendency at all to reject D. argenthona or

nigrina (n=34). It is worth noting that Danaus affinis (Fabricius) bred on

Ischnostemma carnosum and Euploea core (Cramer), which had fed on

Ischnocarpus frustescens, both asclepiadaceous foodplants rich in

cardenoloids but lacking P.A.s, were also eaten by Nephila, but they rejected

E. core which had fed on Parsonsia straminea (Apocynaceae), which does

contain P.A.s (Orr et al. 1996).

Evidence of Batesian mimicry

When on the wing, butterflies of the genus Mynes, especially the females,

bear a striking resemblance to Delias, particularly to members of the

D. nigrina group. It must be stressed that in terms of details of pattern they

are quite imprecise mimics. But exact resemblance may not be necessary as

both Mynes and Delias share a constantly fluttering mode of flight, as

opposed to the sailing flight of many danaids and heliconiids which reveals

clear details of pattern and colour when the butterfly is on the wing. The

flash pattern transmitted by Mynes can be very deceptive. Mynes species

feed on various Urticaceae, which are usually associated with palatability.

M. geoffroyi guerini Wallace is found in northeastern Australia where its

distribution and habitat preferences overlap with those of Delias nigrina. In

flight, the female strongly resembles the female of D. nigrina (cf Common

and Waterhouse 1972). Males vary in their underside coloration, but the

darker forms likewise appear to mimic the male of D. nigrina. This species

is evidently the frequent target of unsuccessful attacks by birds. For example

in a series of 37 M. geoffroyi guerini I collected in southern Queensland, 14

(38%) had large pieces missing from the wings suggestive of avian attack.

Ten of these were males of the pale form (59% of all males, n=17), which are

not obvious mimics. In most cases only one wing was damaged, indicating

that the attack probably took place while the butterfly was flying. A high

incidence of wing damage is a well known characteristic of Batesian mimics,

presumably resulting from tentative attacks by birds. By contrast, in a series

of 83 D. nigrina collected at the same times and places as the Mynes, only

two showed faint beak marks and none had large pieces of wing missing.

Moreover more than half the known Mynes species have a general coloration

similar to syntopic Delias species, at least in the female (Table 1). Where

both sexes are mimetic, the male and female Mynes exhibit the same

upperside coloration as the corresponding sex of the putative Delias model.

Striking parallels occur - for example the underside of the hindwing of

Delias madetes (Godman & Salvin) from New Britain is extensively yellow

with a bright red costal streak and this pattern is mirrored in Mynes

eucosmetos Godman & Salvin from the same locality. Table 1 is by no

means comprehensive in its listing of Mynes forms or potential models and it

is clear that field observations are required to support the suggested

associations (see Parsons 1992, 1999).

Australian Entomologist, 1999, 26 (2)

Table 1. Possible Batesian associations.

Putative model putative mimic distribution

Delias nigrina (Fabricius) Mynes geoffroyi guerini Wallace Eastern Australia

(female/ male) (female/ male dark form)

Delias ornytion (Godman Mynes geoffroyi ogulina Fruhstorfer New Guinea lowlands

& Salvin) (female/ male dark form)

Delias ladas Grose-Smith Mynes geoffroyi ogulina Fruhstorfer New Guinea highlands

(female/ male dark form)

no obvious model

Delias duris Hewitson

Delias funerea

Rothschild ?

No obvious model

Mynes geoffroyi (Guérin-Méneville)

(male pale form)

Mynes doubledayi Wallace

Mynes plateni Staudinger

Mynes halli Joicey & Talbot

Australia, NG

lowlands & highlands

Ceram

Halmahera

New Guinea highlands

No obvious model Mynes websteri Grose-Smith New Guinea highlands

No obvious model Mynes woodfordi Godman & Salvin Solomon Is

Tellervo Kirby sp. Mynes anemone Vane-Wright New Guinea lowlands

Tellervo Kirby sp. Mynes katherina Ribbe (male) New Britain

Delias totilla Heller Mynes katherina Ribbe (female New Britain

yellow form)

Delias madetes (Godman Mynes eucosmetos Godman & Salvin New Britain / New

& Salvin) ? or Hannover

Delias salvini Butler

Delias eschatia Joicey & Mynes talboti Juriaanse & Volbreda Buru

Talbot/ Delias vidua

Joicey & Talbot

Much closer resemblances occur also between certain syntopic Cepora,

Prioneris Wallace and Delias species in New Guinea, Indonesia and

Malaysia. Many examples of strong geographic correspondence in pattern

and colour, apparently centred on a Delias model, are figured by Dixey

(1920). But as the Capparidaceae feeding Cepora and Prioneris may also be

to some extent distasteful, it is uncertain if we can ascribe such convergences

to Batesian mimicry, or to a weak Mullerian association (see Turner 1984).

Both genera are mostly somewhat faster flying than Delias, but the

distinction is not a strong one and in my experience similarly patterned

species of all three genera are easily mistaken.

Certain montane Satyridae in New Guinea appear to mimic Delias species.

In particular Erycinidia virgo (Rothschild & Jordan) and E. dulcis (Jordan)

are slow flying generalized mimics of the Delias aroae-cunningputi group

(Parsons 1988). It should again be noted that these are not very good mimics

in terms of wing pattern, but they have evolved a general coloration and

mode of flight which makes the similarity striking when seen in nature.

50 Australian Entomologist, 1999, 26 (2)

Other species of Erycinidia Rothschild & Jordan, such as E. hemileuca

Jordan, resemble syntopic Delias species on the upperside but not on the

underside and as they are very rapid in flight they are probably not mimetic.

In Southeast Asia both sexes of the satyrids Elymnias vasudeva Moore and

females of E. esaca (Westwood) mimic Delias pasithoe (Linnaeus) or

D. ninus (Wallace) which they strongly resemble in flight (Corbet and

Pendlebury 1992). Most other species of the genus mimic Danaidae.

Evidence of Mullerian mimicry

Apparently Delias are only rarely involved in clear Mullerian associations. It

is probable that certain species of Southeast Asian Cyclosia Hübner

(Zygaenidae) are Mullerian mimics of Delias. Zygaenidae typically feed on

cyanogenic plants and are considered unpalatable (Scoble 1992). There is no

evidence of Mullerian assemblages forming within the Delias. In New

Guinea a few convergences do occur, but this appears to be an incidental

consequence of intense speciation among closely allied forms. There is no

reason to expect the distinctive patterns on the underside to be maintained by

sexual selection, which might be suggested as a source of diversification of

colour patterns opposing the formation of Mullerian rings. Firstly, they are

not generally sexually dimorphic on the undersides; secondly, courting males

expose the upperside of their wings to the female (Orr 1988), presumably

wafting a species-specific androconial secretion over her; and thirdly, closely

allied species with nearly identical underside wing markings, such as

D. isocharis Rothschild & Jordan, D. ligata Rothschild and D. kummeri

Ribbe, fly in the same localities on Mt Kiandi, without apparent

hybridisation. There is a definite tendency for all Delias species occurring at

high altitudes to be very dark, but this is surely a developmental effect due to

low temperatures or an adaptation to allow maximum absorption of the

scarce sunlight. Despite this lack of convergence in pattern, Delias in any

given locality in the New Guinea highlands are very difficult to tell apart on

the wing and are generally perceived in general terms as light coloured

species, yellowish species, dark species and so on. Perhaps with their

fluttering flight they are sufficiently similar to one another for a Mullerian

selective advantage to be operating without the development of obvious

Mullerian resemblances which are apparent to us in the pinned specimen.

Discussion

Although it seems likely that Delias species sequester toxic compounds, the

chemical nature of these must remain uncertain. The mistletoes, at least

those of the family Viscaceae, are known to contain potent cardioactive

compounds (Samuelsson 1962). This is apparently the source of the

supposed magical properties attributed to the plant by Celtic druids. Active

compounds isolated include choline and y-amino butyric acid, but most

significant toxic substances are complex polypeptides (viscotoxin). In

Australia Delias species feed naturally on members of both families of

Australian Entomologist, 1999, 26 (2) 51

mistletoe, with the Loranthaceae dominating. Samuelsson (1966, 1969) did

not find viscotoxins in European members of this family, but as our

understanding of mistletoe chemistry is clearly incomplete, this should not be

taken as evidence of a lack of toxicity in Australian region species.

However, even if we assume that complex toxic compounds confer toxicity

on the insects which feed on them, it is still unclear if these are sequestered

directly, or are in some way modified to form secondary protective toxic

substances.

As the Delias appear to be strongly aposematic, questions must be asked such

as why do they not attract high quality Batesian mimics, why are so few

Delias species involved as models and why is there no obvious development

of well defined Mullerian rings? The answers to these questions are probably

twofold. Firstly, as discussed above, since Delias constantly flutter an

approximate mimetic resemblance in coloration is probably effective as long

as the correct flight pattern is adopted. The efficacy of an approximate

resemblance will also tend to prevent the formation of obvious Mullerian

tings. Secondly, the Delias are very probably of Gondwanan origin, adapted

largely to temperate or montane rainforest. Only a handful of other

butterflies which belong to potentially mimetic groups are of similar origin

and currently occur in the same types of habitats as Delias. Mynes is an

Australian region genus of uncertain affinities which does overlap broadly

with some Delias in habitat preferences and the two genera have probably

existed together for a very long time. The montane satyrid genus Erycinidia

may also be of southern origin. The remainder of the common Batesian

mimetic groups, such as Hypolimnas Hiibner and Elymnias Hiibner, are of

Oriental origin and may simply not have coexisted for long enough with the

few species of Delias which have established themselves in the tropical

lowlands for associations to develop, especially as most species in these

genera were already Batesian mimics of the very differently patterned

danaine genera. Moreover, these same danaiines are already largely involved

in Mullerian associations with each other (Ackery and Vane-Wright 1984)

and convergence toward the vastly different pattern and mode of flight of

Delias is unlikely. This postulated historical effect could be tested by

examining other probable mistletoe feeding Pieridae, Mylothris Hübner in

Africa and Archonias Hiibner (?), Catasticta Butler, Pereute Herrich-

Schaeffer, Charonius Röber (?) and Leodonta Butler (?) in South America

(see DeVries 1987) to determine if they are more closely involved in mimetic

associations than are Delias. A perusal of D’Abrera (1978, 1981-94)

suggests this may be the case.

Acknowledgments

I thank Roger Kitching for playing devil’s advocate on many occasions on

this subject and David Hancock for helpful comments on the manuscript.

52 Australian Entomologist, 1999, 26 (2)

References

ACKERY, P.R. and VANE-WRIGHT, R.I. 1984. Milkweed butterflies: their cladistics and

biology. British Museum (Natural History), London.

BROWN, K.S. 1984. Adult-obtained pyrrolizidine alkaloids defend ithomiine butterflies

against a spider predator. Nature (London) 309: 707-709.

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

Robertson, Sydney.

CORBET, A.S. and PENDLEBURY, H.M. 1992. The butterflies of the Malay Peninsula.

Fourth edition revised by J.N. Eliot. Malay Nature Society, Kuala Lumpur.

D'ABRERA, B.A. 1978. Butterflies of the Afrotropical region. Landsdowne, Melbourne.

D'ABRERA, B.A. 1981-94. Butterflies of the Neoptropical region, Parts I-VII. Landsdowne

and Hill House, Melbourne.

DeVRIES, P.J. 1987. The butterflies of Costa Rica and their natural history. Princeton

University Press, Princeton, NJ.

DIXEY, F.A. 1918. On mimicry in certain butterflies of New Guinea. Transactions of the

Entomological Society of London 1918: 118-129.

DIXEY, F.A. 1920. The geographical factor in mimicry. Transactions of the Entomological

Society of London 1920: 208-211.

ORR, A.G. 1988. Mate conflict and the evolution of the sphragis in butterflies. PhD thesis,

Griffith University, Brisbane.

ORR, A.G., TRIGO, J.R., WITTE, L. and HARTMANN, T. 1996. Sequestration of

pyrrolizidine alkaloids by larvae of Tellervo zoilus (Lepidoptera: Ithomiinae) and their role in

the chemical protection of adults against the spider Nephila maculata (Araneidae).

Chemoecology 7: 68-73.

PARSONS, M.J. 1991. Butterflies of the Bulolo-Wau valley. Bishop Museum, Honolulu.

PARSONS, M.J. 1999. Butterflies of Papua New Guinea. Academic Press, New York.

ROBBINS, R.K. 1980. The lycaenid “false head” hypothesis: historical review and quantitative

analysis. Journal of the Lepidopterists' Society 34: 194-208.

SAMUELSSON, G. 1962. Phytochemical and pharmacological studies on Viscum album L. VI

Studies on the homogeneity of Viscotoxin A. Svensk farmakologie tidskrift 66: 201-211

SAMUELSSON, G. 1966. Screening of plants of the family Loranthaceae for toxic proteins.

Acta Pharmacologica Suecica 3: 353-362.

SAMUELSSON, G. 1969. Screening of plants of the family Loranthaceae for toxic proteins.

Part II. Acta Pharmacologica Suecica 6: 441-446..

SCOBLE, M.J. 1992. The Lepidoptera, form, function and diversity. Oxford University Press,

Oxford.

TURNER, J.R.G. 1984. Mimicry: the palatbility spectrum and its consequences. Pp 141-161

in: Vane-Wright, R.I and Ackery, P.R. (eds). The biology of butterflies. Symposium of the

Royal Entomological Society of London 11.

Australian Entomologist, 1999, 26 (2): 53-55 53

NEW AUSTRALIAN RECORDS OF XEROPHILIC ACARIFORM

MITES (ORIBATIDA AND PROSTIGMATA)

Roy A. Norton' and Adrianne Kinnear?

"State University of New York, College of Environmental Science & Forestry, I Forestry Drive,

Syracuse, New York, USA 13210

"School of Natural Sciences, Edith Cowan University, Bradford Street, Mount Lawley,

WA 6050, Australia

Abstract

The first Australian records of four xerophilic mite taxa are reported. Amnemochthonius

taeniophorus Grandjean (Oribatida: Haplochthoniidae) was collected from soil in southwestern

Western Australia. Gordialycus tuzetae Coineau et al., two undescribed species of

Nematalycus Strenzke (all Nematalycidae) and an undetermined species of Stigmocheylus

Berlese (Stigmocheylidae) were represented in a collection from a coastal sand dune near Perth.

Neither of the latter two families has been reported previously from Australia.

Introduction

Xerophilic mites — ones found predominantly in dry environments, or in dry

microhabitats within mesic environments - are found in most major taxa of

the mite order Acariformes and even some entire families may be so

characterized. One example is the oribatid mite family Haplochthoniidae,

which with four other xerophilic families comprise the Protoplophoroidea

(Norton et al. 1983). Another is Nematalycidae, the three known genera of

which inhabit fine sands (Thibaud and Coineau 1998). This latter family is

usually included in a paraphyletic group of early-derivative acariform mites,

the Endeostigmata (Krantz 1978, Evans 1992), but is considered by others

(e.g. Kethley 1982) to be closely related to Tydeoidea (Prostigmata). Our

purpose is to report the first Australian records for several taxa in these

families, as well as another taxon whose habitat requirements are poorly

known.

Discussion

Haplochthoniidae has few known species but one, Haplochthonius simplex

Willmann, is widely distributed and is a common human associate

(Grandjean 1946). Two species in the genus are known from Australia,

though neither has been identified (Colloff and Halliday 1998). The second

genus in the family, Amnemochthonius Grandjean, was known only from the

type species, A. taeniophorus Grandjean. The latter was described from 18

specimens from xeric habitats in southern and western France (Grandjean

1948) and no collections have been reported since in the literature. Based on

our comparisons with Grandjean's (1948) meticulous description, the same

species is represented at two xeric locations in Western Australia.

One adult, one tritonymph, one deutonymph and one larva of

A. taeniophorus were collected from ‘good condition’ swale sites on

Boolethana Station (24°39’S, 113°42’E), 50km north of Carnarvon, Western

54 Australian Entomologist, 1999, 26 (2)

Australia, in soils of the Sable Land System, in August 1994. One adult was

collected from Widgemooltha (31°30’S 121°32’B), in relict laterite soils of

the Greenstone belt region of the Eastern Goldfields, within the former CRA

(Conzinc Rio Tinto) prospecting site, Widgiemooltha No. 3 (described in

Kinnear 1991), in November 1985.

This mite has a number of curious morphological traits. For example,

Grandjean (1948) noted that the dorsal setae in rows ps and h bend to the left

on both sides in his French specimens. He questioned whether this

asymmetry would prove to be a variable trait, but the same distinct bend is

present in the Australian specimens. Grandjean (1948) did not study a larva

of this species but he predicted the absence of Claparédes organ, the papilla-

like structure found between the bases of leg I and II. His rationale was that

other instars of this mite lack genital papillae and there is a close correlation

between the form of these two types of organs in other oribatid mites.

Indeed, the larva we studied lacks this organ.

Specimens of three species of Nematalycidae, representing the first records

of the family from Australia (cf Halliday 1998), were collected by us from

Mindare Keys Beach, north of Perth, Western Australia (31°57’S 115°31’E)

on 18 April 1998. The mites were washed from calcareous, unconsolidated

fine sand on the vegetated lee side of a large, coastal foredune, in the

Quindalup dune system (see description in Seddon 1972). One, Gordialycus

tuzetae Coineau et al. (1967), is a greatly elongated, vermiform species that

was first recorded from beach sand on the coast of southern France. As

recently summarized by Thibaud and Coineau (1998), it has since been

reported from inland sites in South Africa, Namibia and Turkmenistan. The

latter authors included new records of the genus (it is currently monotypic)

from Venezuela, Cuba, Mauritania and New Caledonia. Our specimens,

collected at a depth of 0-30 cm, closely match the original description of this

species and we have no reservation about its identity. At deeper levels in this

same location (60-90 cm), we found several specimens of what appear to be

two undescribed species of Nematalycus Strenzke. Both are most similar to

N. strezkei Cunliffe, especially in having a much richer hysterosomal

chaetome, relative to the only other described species, N. nematoides

Strenzke. However, each has distinctly different appendage structures.

Two specimens of an undetermined species of Stigmocheylus Berlese were

collected from 60-90 cm depth in the same sand dune. This genus, currently

monotypic and previously unreported from Australia, is usually included in

the anystine family Pseudochelidae (other genera of which are known from

Australia: Halliday 1998). Kethley (1990) transferred this genus to the

monogeneric Stigmocheylidae. The general habitat requirements of these

mites are not yet well characterized.

Australian Entomologist, 1999, 26 (2) 55

Voucher specimens of all five mites are in the collection of the second

author. Vouchers of the nematalycids and Stigmocheylus sp. are in the

Western Australian Museum.

Acknowledgments

We thank Drs M. Judson (Paris) and J. B. Kethley (Chicago) for helpful

information.

References

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France des Acariens Nematalycidae Strenzke à l'occasion d'aménagement du Languedoc-

Rousillon. Comptes Rendu d’Académie des Sciences, Paris 265: 685-688.

COLLOFF, M.J. and HALLIDAY, R.B. 1998. Oribatid mites: a catalogue of the Australian

genera and species. CSIRO Publishing, Melbourne.

EVANS, G.O. 1992. Principles of acarology. CAB International, Cambridge; 563 pp.

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Naturelle, Zoologie 8:213-248.

GRANDJEAN, F. 1948. Les Enarthronota (Acariens) (2e série). Annales des Science

Naturelle, Zoologie 10: 29-58.

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Melbourne.

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classification of living organisms, Vol. 2. McGraw-Hill Inc., New York; pp 117-145.

KETHLEY, J.B. 1990. Prostigmata. Jn: Dindal, D.L. (ed.), Soil biology guide. John Wiley &

Sons, New York; pp 667-778.

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region of Western Australia. Pedobiologia 35:273-283.

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the. Pediculochelidae (Acari: Acariformes). Proceedings of the Entomological Society of

Washington 85: 493-512.

SEDDON, G. 1972. Sense of place. University of Western Australia Press, Nedlands, WA.

THIBAUD, J.-M. and COINEAU, Y. 1998. Nouvelles stations pour le genre Gordialycus

(Acarien: Nematalycidae). Biogeographica 74: 91-94.

56 Australian Entomologist, 1999, 26 (2)

BOOK NOTICES

The Amazing World of Stick and Leaf Insects. By Paul D. Brock (1999). Amateur

Entomologists' Society, Orpington, Kent, UK. 182 pp, 46 figs, 26 B/W plates, 40

colour plates. ISBN 0-900054-63-8. Price 16.20 pounds sterling (incl. postage).

Rearing and Studying Stick and Leaf Insects. By Paul D. Brock (1992). Amateur

Entomologists' Society, Feltham, Middlesex, UK. 79 pp, 40 figs, 7 B/W plates. ISBN

0-900054-54-9. Price 5.50 pounds sterling (incl. postage).

Phasmids include some of the largest (certainly the longest) and most spectacular

insects on earth. Australia has its share of the striking species. Among these are the

large green and yellow Eurycnema goliath (probably our heaviest), the longer (but

lighter), grey and purple Acrophylla titan, and the bizarre, flanged, green or mottled

Extatosoma tiaratum. The weird appearance of the last makes it one of the most

photographed of all insects.

All three species mentioned are easily reared on fresh foliage, breed easily in

captivity, and have great potential as pets for children and home hobbyists. They

certainly have more exciting personalities than most goldfish! We keep them as a

permanent live display at the Queensland Museum and have given thousands of eggs

and nymphs away to wide-eyed children over the years. But there is little evidence of

their popular acceptance as hobbyist animals in Australia and I have never seen

phasmids available in an Australian petshop. It is thus astonishing to visit England

and find hundreds of amateur enthusiasts rearing phasmids in their spare rooms,

avidly swapping eggs of prized species, and checking out the latest species in stock at

the petshop down the road. And even more astonishing to find that all three

Australian species mentioned above are popular members of this hobbyist trade.

Much of the hobbyist interest in phasmids in England (and Europe) arises from the

activities of Paul Brock who lives in suburban Slough, on the western outskirts of

London, and leads a double life as a bank auditor by day and a phasmid enthusiast

extraordinaire by night (and every moment of leave). Paul is a leading light in the

Phasmid Study Group, a group with some 500 members, mostly devoted rearing stick

and leaf insects. He also travels widely around the world, studying and

photographing phasmids, and has been to Australia several times.

The two books of Paul's cited above are de rigeur for anyone wanting to take up

rearing phasmids. The 1992 handbook gives a comprehensive and well-illustrated

account of techniques for the task. It includes detailed notes on many individual

species including a number from Australia. The just-released 1999 volume gives a

world tour of phasmids including an overview of the world classification and

numerous colour photographs, mostly taken of exotic species from Paul's own travels.

Many Australian species are covered, including the first publication of the

photographs of the mystery giant species of Ctenomorpha from North Queensland

which at approximately 30 cm head/tail length will prove to be Australia's longest

species (when we get a specimen for verification!).

These useful, attractive and informative books will open the door to a fascinating

world for anyone in Australia wanting to breed and study these wonderful beasts.

Geoff Monteith, Queensland Museum, Box 3300, South Brisbane, Qld, 4101.

Australian Entomologist, 1999, 26 (2): 57-63 57

THE LIFE HISTORY OF

PRAETAXILA SEGECIA PUNCTARIA (FRUHSTORFER)

(LEPIDOPTERA: LYCAENIDAE: RIODININAE)

P.R. SAMSON’, S.J. JOHNSON? and P.R. WILSON’

'Bureau of Sugar Experiment Stations, PMB 57, Mackay Mail Centre, Qld 4741

*Oonoonba Veterinary Laboratory, PO Box 1085, Townsville, Qld 4810

"Department of Natural Resources, PO Box 1143, Bundaberg, Qld 4670

Abstract

The immature stages of Praetaxila segecia punctaria (Fruhstorfer) are described from the

Rocky River, Cape York Peninsula, Australia. The larval food plant is Rapanea porosa

(Myrsinaceae). Eggs are laid in small groups beneath the leaves. There are five larval instars.

Larvae feed at night, mostly on young leaves. They remain on the plant for the first three

instars, but from the fourth instar onwards they shelter beneath debris on the ground during

daylight hours. The egg, larval and pupal stages occupied about 1, 6 and 2-3 weeks,

respectively, when reared at Mackay during winter.

Introduction

Praetaxila segecia punctaria (Fruhstorfer) (Australian harlequin) is the sole

Australian representative of the lycaenid subfamily Riodininae. Some

authors consider this subfamily to be a family in its own right (e.g.

McCubbin 1971). P. s. punctaria is known from Cape York Peninsula in

northern Queensland, as far south as the McIlwraith Range (Moulds 1991),

with other subspecies known from New Guinea and the Aru Islands

(Common and Waterhouse 1981). Its life history has not been recorded

previously. Here we describe the immature stages collected at the Rocky

River, to the east of the McIlwraith Range.

Immature stages

Egg (Fig. 1). Barrel-shaped with rounded sides, flattened base and top, with

sunken micropyle; smooth apart from a band of fine white ‘hairs’ around the

sides a small distance below the upper rim; purple when freshly laid, the first

instar later becoming visible through the clear egg shell. Diameter 0.9 mm,

height 0.7 mm.

First instar (Fig. 4). Elongate, with extremely long finely-branched hairs

mostly greyish brown in colour; prothorax (segment 1) with hairs directed

forward over head; segments 2-11 each with lateral lobes bearing a group of

hairs, and with two pairs of dorsal hairs, the outer pair on segments 4-10

splitting at the base into two long hairs of similar length, the outer pair on

segments 2, 3 and 11 not splitting and extremely long; anal segment with

short and long hairs; body initially whitish, becoming green after feeding;

head light brown. Larvae appear to have only 12 thoracic and abdominal

segments; as noted by DeVries (1997), abdominal segments 9 and 10 are

fused in riodinids.

Australian Entomologist, 1999, 26 (2)

EEB EVE NDR

Figs 1-3. Praetaxila segecia punctaria: (1) egg; (2) fifth (final) instar larva, head at

top; (3) pupa, head at top. Scale bars (1) = 0.5 mm, (2, 3) = 5 mm.

Australian Entomologist, 1999, 26 (2)

Figs 4-5. Praetaxila segecia punctaria: (4) first instar consuming eggshell; (5) second

instar, head at left. Scale bars (4) = 1 mm, (5) = 2 mm.

60 Australian Entomologist, 1999, 26 (2)

Second instar (Fig. 5). Prothorax with pale brown hairs around anterior

margin; segments 2-11 each with lateral lobes bearing many white hairs and

with a pair of subdorsal tubercles bearing very long greyish erect hairs or

slightly shorter whitish hairs; segments 2, 3 and 11 each with a pair of

extremely long white subdorsal hairs; anal segment with greyish and white

hairs; anterior prothorax and anal segment orange, rest of body green, a pair

of large black subdorsal patches on segment 2 and a pair of black dorsolateral

spots on each of segments 4-10, spiracles white, head light brown.

Third instar. Body form and hairs similar to second instar, but with

subdorsal tubercles enlarged on segments 2-3 and with a pair of small

dorsolateral raised patches with white hairs on each of segments 4-11; a pair

of extremely long white hairs on each of segments 2 (directed forwards), 3

and 11, with several pairs directed backwards from anal segment; prothorax

and anal segment orange, other segments greenish grey with a green

middorsal line, large black subdorsal patches on segment 2 covering

tubercles and almost touching medially, segments 3-11 each with two pairs

of black subdorsal spots forming an intermittent line, segments 4-10 each

with a pair of black dorsolateral spots joined by a grey line, an orange patch

around each abdominal spiracle, head brown.

Fourth instar. Form and markings similar to third instar, but colour of

segments 2-11 pale grey with a greenish grey middorsal line that is edged

whitish.

Fifth (final) instar (Fig. 2). Similar to fourth instar but colour duller and

black markings less defined, without black patches on segment 2 and without

defined dorsolateral spots.

Pupa (Fig. 3). Attached by cremaster and girdle. Prothorax slightly

flattened, rounded anteriorly with median indentation, lateral swellings at

posterior margin; metathorax with lateral swellings, abdominal segment 1

with a pair of dorsal knobs holding girdle, abdominal segments 2-4 very

swollen laterally; dorsal surface covered with coarse brown hairs and fine

light brown hairs, abdomen with fine long white hairs ventrally; colour

usually purplish brown, a black dorsal patch on abdominal segment 1, two

pairs of black subdorsal spots on mesothorax and one pair each on

metathorax and abdominal segments 2-8, two pairs of black dorsolateral

spots on each of abdominal segments 2-7 and a single pair on 8, prothorax

with black middorsal line and a pair of faint black subdorsal spots,

mesothorax and metathorax with obscure black dorsolateral spots. Above

description applies to pupae attached to dead leaves; colour of two pupae

attached to living green leaves was light brown with similar markings to

those described above.

Australian Entomologist, 1999, 26 (2) 61

Life history

The larval food plant at the Rocky River was Rapanea porosa (R. Br.) Mez

(Myrsinaceae). This plant was very common at the Rocky River, forming

part of the understorey within the rainforest. Immature stages of

P. s. punctaria were found on both large and small specimens.

Our first encounter with the life history occurred on 11 May 1998 at about

1300 hrs. A female was seen resting on top of a leaf about 1 m from the

ground, deep in the rainforest. She soon flew to the upper surface of a leaf at

a similar height on an adjacent specimen of R. porosa and oviposited

underneath, curling her abdomen around the leaf margin. Eggs were not laid

continuously; the female withdrew her abdomen at one stage before

recommencing laying. Four eggs were subsequently found under the leaf.

The female was caged on a branch of the same plant, and by 1600 hrs of the

following day had laid another 28 eggs.

Our second encounter happened the next day. While one of us (SJJ) was

crouching in the rainforest shortly after midday, a female alighted on a

nearby leaf. She soon moved to the edge of the leaf and oviposited beneath.

Again the female rested during the oviposition sequence, and again four eggs

were laid. The same day, a third group of four eggs was found beneath a

leaf, this time with first instars just hatching. In all instances the eggs were

under older leaves, and were close together but not touching.

We also found several small larvae singly beneath leaves, but could find no

larger larvae.

The eggs and larvae were subsequently reared on potted specimens of the

food plant or on the exotic plant Ardisia humilis (Myrsinaceae), which is

available in nurseries. This plant was readily accepted by larvae. However,

another species, A. crenata, proved unsuitable. The following notes were

made during rearing.

First instars ate most of the egg shell after hatching, except for the base of the

shell attached to the leaf (Fig. 4). They then ate small patches part-way

through the underside of the leaves on which the eggs were laid. The larvae

remained in a group, and this gregarious habit was retained throughout

development. By the second instar, larvae had moved to younger leaves,

from which they ate circular holes (< 1 cm diameter) by feeding from the

undersurface. Third instars rested beneath older leaves during the day and

fed on younger leaves at night. Subsequent instars left the plant during the

day, sheltering singly or more often in groups among dead leaves on the soil

surface. They moved onto the plant early in the evening and returned to the

ground in the early morning. Fourth instars ate pieces from the margin of

young leaves, starting as linear cuts which they gradually enlarged through

the night. Final instars chewed away the ends and margins of young leaves.

62 Australian Entomologist, 1999, 26 (2)

Larvae also fed on mature leaves in later instars. Most larvae pupated among

dead leaves on the soil surface, but two pupae were attached to older living

leaves near the base of the stem. Eggs hatched in about 1 week, and the

duration of the larval and pupal stages was about 6 weeks and 2-3 weeks,

respectively, at Mackay during May-July.

The Rocky River was revisited on 16 June 1998. With our knowledge of the

nocturnal habits of fourth and fifth instar larvae, a search was made of plants

by torch-light during the evening. Several large larvae were found feeding.

Debris on the ground was searched during daylight, and large larvae were

found sheltering beneath dead leaves within 20 cm of the base of the food

plants. No pupae were found, and it is possible that larvae may wander

further before pupation. Most larvae were found singly, as they had been

during our previous visit in May. However, on one occasion two fourth

instar larvae were found together beneath a dead leaf and a third instar,

presumably from the same cohort, was found beneath a leaf of the host plant.

Both larger larvae were subsequently found to be parasitised by an

unidentified braconid wasp. In our field observations of solitary larvae, it is

possible that other members of larval cohorts had died or were in other stages

of development.

Adults have been observed by us at the Rocky River and elsewhere on Cape

York Peninsula over several years. They prefer shaded areas and commonly

fly on to tracks or clearings and alight on leaf litter or feed at blossom. In all

instances, they have been active in late morning and early afternoon, and we

have seen no evidence that they are crepuscular as suggested by Moulds

(1991).

Plants in the family Myrsinaceae are host to some other genera of

Riodininae, including Zemeros and Dodona (Corbet and Pendlebury 1978),

Abisara (Corbet and Pendlebury 1978; DeVries et al. 1992) and Saribea

(DeVries et al. 1992). In his higher classification of the riodinids, Harvey

(1987) placed all of these genera in the subfamily Hamaerinae (DeVries,

pers. comm.), treated as the tribe Hamaerini in the present paper. As is

predicted for all members of the Hamaerini (Harvey 1987), the larvae of P. s.

punctaria do not appear to have ant-attracting organs or associate with ants

as do some neotropical groups (DeVries 1997).

Acknowledgments

We are grateful to the Queensland Herbarium for identifying the larval food

plant, and to P. DeVries of the University of Oregon for advice on

relationships within the Riodininae.

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Australian Entomologist, 1999, 26 (2) 63

CORBET, A.S. and PENDLEBURY, H.M. 1978. The butterflies of the Malay Peninsula, 3rd

edn. revised by J.N. Eliot. Malayan Nature Society, Kuala Lumpur, xiv + 578 pp.

DeVRIES, P.J. 1997. The butterflies of Costa Rica and their natural history, Volume II,

Riodinidae. Princeton University Press, Princeton; xxv + 288 pp.

DeVRIES, P.J., CHACON, LA. and MURRAY, D. 1992. Toward a better understanding of

host use and biodiversity in riodinid butterflies (Lepidoptera). Journal of Research on the

Lepidoptera 31: 103-126.

HARVEY, D.J. 1987. The higher classification of the Riodinidae (Lepidoptera). PhD. thesis,

University of Texas, Austin.

McCUBBIN, C. 1971. Australian butterflies. Thomas Nelson, Melbourne; xxx + 206 pp.

MOULDS, M.S. 1991. Notes on the distribution and adult behaviour of Praetaxila segecia

punctaria (Fruhstorfer) (Lepidoptera: Lycaenidae: Riodininae). Australian Entomological

Magazine 18: 113-114.

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AN ACCUMULATIVE BIBLIOGRAPHY OF

AUSTRALIAN ENTOMOLOGY

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Backhuys Publishers: Leiden.

ANDRESS, R.

1998 Description of the larva of Petalura ingentissima Tillyard, 1907 (Anisoptera: Petaluridae). Odonatologica 27:

353-359.

BICKEL, D.J.

1998 Australian, Melanesian and Micronesian Acropsilus Mik (Diptera: Dolichopodidae). Tijdschr. Ent. 141: 1-17.

1998 Cunomyia, a distinctive new hilarine fly genus from the Tasmanian World Heritage Area (Diptera: Empididae).

Pap. Proc. R. Soc. Tasm. 132: 59-63.

1999 Australian Sympycninae II: Synrormon Loew and Nothorhaphium, gen. nov., with a treatment of the Western

Pacific fauna, and notes on the subfamily Rhaphiinae and Dactylonotus Parent (Diptera: Dolichopodidae).

Invert. Taxon. 13: 179-206.

BRAILOVSKY, H. and CASSIS, G.

1999 New genus and new species of Amorbini (Heteroptera : Coreidae) from Australia. Proc. ent. Soc. Wash. 101:

69-74

1999 Revision of the tribe Agriopocorini (Hemiptera : Coreidae : Coreinae). Can. Ent. 131: 293-321.

BRUST, R.A., BALLARD, J.W.O., DRIVER, F.,HARTLEY, D.M., GALWAY, N.J. and CURRAN, J.

1998 Molecular systematics, morphological analysis, and hybrid crossing identify a third taxon, Aedes (Halaedes)

wardangensis sp.nov., of the Aedes (Halaedes) australis species-group (Diptera : Culicidae). Can. J. Zool. 76:

1236-1246.

DAVIES, D.A.L.

1998 The genus Peralura: Field observations, habits and conservation status (Anisoptera: Petaluridae).

Odonatologica 27: 287-305.

EASTWOOD, R. and FRASER, A.M.

1999 Associations between lycaenid butterflies and ants in Australia. Aust. J. Ecol. 24: 503-537.

GESS, F.W. ;

1995 Descriptions of the male of Riekia nocatunga Richards, the male and two strikingly distinct sympatric colour

forms of Riekia confluens (Snelling) and the male of Rolandia angulata (Richards) (Hymenoptera: Vespidae:

Masarinae) from Australia. J. hym. Res. 4: 33-40.

GESS, F.W., GESS, S.K.'and GESS, R.W.

1995 An Australian masarine, Rolandia angulata (Richards) (Hymenoptera: Vespidae): nesting and evaluation of

association with Goodenia (Goodeniaceae). J. hym. Res. 4: 25-32.

GREY, EJ.

1999 A bizarre ant, Victorian Nat. 116: 82.

1999 The beetle Gondwanennebous minutissimus Kaszab (Coleoptera: Archeocrypticidae) - a first record for

Victoria., Victorian Nat. 116: 91.

HORAK, M.

1998 A reassessment and review of the Australian genus Ctenomeristis Meyrick (Lepidoptera: Pyralidae: Phycitinae).

Ent. scand. 28: 445-470.

KOJIMA, J.

1999 Male genitalia and antennae in an Old World paper wasp genus Ropalidia Guérin-Méneville, 1831 (Insecta:

Hymenoptera; Vespidae, Polistinae). Nat. Hist. Bull. Ibaraki Univ. 3: 51-68.

KOJIMA, J. and CARPENTER, J.M. i

1997 Å taxonomic note and nest description of an Australian paper wasp, Polistes variabilis (Fabricius)

(Hymenoptera, Vespidae, Polistinae). Japan. J. syst. Ent. 3: 117-122.

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1994 Thirty-five new Hydrochus species from the Old and the New World (Coleoptera: Hydrophilidae). Annls hist.-

nat. Mus. natn. hung. 86: 29-42.

1995 Descriptions of ten new species of Hydrochus from different parts of the world (Coleoptera: Hydrochidae).

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1998 Polymorphism and kleptoparasitism in thrips (Thysanoptera: Phlaeothripidae) from woody galls on Casuarina

trees. Aust. J. Ent. 37: 8-16

NEVILLE, T., SCHWARZ, M.P. and TIERNEY, S.M.

1998 Biology of a weakly social bee, Exoneura (Exoneurella) setosa (Hymenoptera: Apidae) and implications for

social evolution in Australian allodapine bees. Aust. J. Zool. 46: 221-234.

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1998 Male genital musculature of Therevidae and Scenopinidae (Diptera: Asiloidea): structure, homology and

phylogenetic implications. Aust. J. Ent. 37: 27-33.

ENTOMOLOGICAL NOTICES

Items for insertion should be sent to the editor who reserves the right to alter, reject or

charge for notices.

FOR SALE: Butterflies from all parts of the world. Papua New Guinea,

Peru, Indonesia, Thailand, China, Africa, Brazil, Colombia, etc.

Papilionidae inc. Parnassius, Delias, Charaxes etc. Free catalogue. David

Hall, 6 Rule St, Cambridge Park, N.S.W., 2747. Ph. 02 4731 2410.

ENTOMOLOGICAL BOOKS. Pemberley Books are specialist suppliers of

entomological literature across the world. Send for our free catalogue

which lists a wide range of antiquarian, second-hand and new natural

history titles. Pemberley Books, lan Johnson, 34 Melrose Close, Hayes,

Middlesex, UB4 OAZ, England. Tel/Fax: +44 181 561 5494. E-mail:

ij @pembooks.demon.co.uk

WANTED. Any information regarding Rhytiphora macleayi (Coleoptera:

Cerambycidae), particularly from private collections. Mark Hura, 111

Oleander Drive, Parafield Gardens, S.A., 5107.

NOTES FOR AUTHORS

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THE AUSTRALIAN

Entomologist

Volume 26, Part 2, 19 November 1999

CONTENTS

LACHLAN, R.B.

A new species of Delias Hübner (Lepidoptera: Pieridae) from the Star Mountains,

Papua New Guinea. 33

MOULDS, M.S. and LANE, D.A.

A new hawk moth from northern Australia with notes on its life history

(Lepidoptera: Sphingidae). 37

NORTON, R.A. and KINNEAR, A.

New Australian records of xerophilic acariform mites (Oribatida and Prostigmata). 53

ORR, A.G.

Evidence for unpalatability in the genus Delias Hübner (Lepidoptera: Pieridae)

and its role in mimetic assemblages. 45

SAMSON, P.R., JOHNSON, S.J. and WILSON, P.R.

The life history of Praetaxila segecia punctaria (Fruhstorfer) (Lepidoptera:

Lycaenidae: Riodininae). 57

BOOK REVIEWS

Australian Ants: Their Biology and Identification. 36

The Amazing World of Stick and Leaf Insects. 56

Rearing and Studying Stick and Leaf Insects. 56

RECENT LITERATURE

An accumulative bibliography of Australian entomology 64

ENTOMOLOGICAL NOTICES Inside back cover.

ISSN 1320 6133

OLN

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