THE AUSTRALIAN ENTOMOLOGIST

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Cover: Our cover illustration for 2013 shows a species of the pebble-mimicking

genus Raniliella (Orthoptera: Acrididae). There are two species in the genus, both

restricted to desert environments of the northern parts of South Australia. They are

commonly seen in stony areas around the Flinders Ranges. Raniliella is one of

several genera of Australian grasshoppers which closely resemble the stones of our

extensive gibber deserts. The illustration is reproduced by permission from CSIRO's

Insects of Australia and is by the late Frank Nanninga, a Dutch-born artist who was

the leading insect illustrator of the 1960s in Australia. His work reached its peak in

the eight magnificent colour plates which grace the pages of Insects of Australia.

Australian Entomologist, 2013, 40 (4): 173-174 173

THE FIRST RECORD OF PARADUBA METRIODES (BETHUNE-

BAKER) (LEPIDOPTERA: LYCAENIDAE) WITHIN AUSTRALIAN

LIMITS

STEPHEN J. JOHNSON! and IAN R. JOHNSON?

‘Queensland Museum, PO Box 3300, South Bank, Qld 4101

2260 Grassdale Road, Gumdale, Qld 4154

Abstract

A single male of Paraduba metriodes (Bethune-Baker) was collected on Dauan Island, Torres

Strait, Queensland. Illustrations are provided of the adult and the characteristic male valvae and

aedeagus that separate Paraduba Bethune-Baker from Nacaduba Moore.

Introduction

The lycaenid genus Paraduba Bethune-Baker has three species confined to

mainland Papua New Guinea and Normanby Island (Parsons 1998). It is

separated from Nacaduba Moore and other allied genera by the simple,

clavate, densely-haired valvae and a strongly cornute aedeagus (Tite 1963).

On a recent trip to Dauan Island in early March 2013, a single male was

collected and presumed at the time to be a male Catopyrops ancyra (Felder).

However, subsequent closer examination showed that it was not C. ancyra

but likely to be Paraduba metriodes (Bethune-Baker, 1911) (Figs 1-2). This

was confirmed by examination of the genitalia (Figs 3-4). Paraduba

metriodes is common and widespread in lowland Papua New Guinea

(Parsons 1998, Chris Müller pers. comm.), with females less often

encountered than males.

Discussion

The male of P. metriodes is easily confused with C. ancyra but the lack of a

long hindwing tail separates it from that species. Placement within Paraduba

is confirmed by the presence of densely haired valvae (Fig. 3) and the cornute

aedeagus (Fig. 4). The forewing upperside is lilac-blue with very fine black

margins. The hindwing upperside is similar but with 2 black spots on tornal

margin and a short, stubby tail at the end of vein CuA>. The underside is

grey-brown with darker bands faintly edged white and the tornal spots edged

orange medially. Eyes hirsute. Forewing length 13.4 mm; wingspan 24 mm.

The specimen was taken in an area of remnant vine thicket growing on a sand

ridge close to the beach on the northern side of the island. It was engaged in

apparent lekking behaviour around a plant of Caesalpinia bonduc (L.)

(Caesalpiniaceae) growing on a sand dune adjacent to the beach. Despite

being disturbed on several occasions, it continually returned to resume its

position on the plant. This behaviour suggests that the plant may be a host.

Although the life history of P. metriodes is unknown, Caesalpinia bonduc is

known to be a host for the closely related Catopyrops ancyra elsewhere in

Torres Strait (Orr and Kitching 2010, S. Johnson pers. obs.) and for C.

florinda (Butler) on mainland Australia (Braby 2000).

174 Australian Entomologist, 2013, 40 (4)

Figs 1-4. Paraduba metriodes (Bethune-Baker) from Dauan Island: (1-2) male

upperside and underside. (3-4) Male genitalia: (3) lateral view of valva and aedeagus;

(4) dorsal view including aedeagus.

Acknowledgements

The authors thank the Chairman of Dauan Island Council for permission to

collect on the island, Geoff Thompson for photography and other staff at

Queensland Museum for assistance in various ways, and Peter Valentine and

Geoff Monteith for editorial assistance.

References

BRABY, M.F. 2000. Butterflies of Australia; their identification, biology and distribution.

CSIRO Publishing, Collingwood; xx + 976 pp.

ORR, А.С. and KITCHING, R.L. 2010. The butterflies of Australia. Allen & Unwin, Sydney;

viii + 328 pp.

PARSONS, M. 1998. The butterflies of Papua New Guinea: their systematics and biology.

Academic Press, London; xvi + 736 pp, 162 pls.

TITE, G.E. 1963. A synonymic list of the genus Nacaduba and allied genera (Lepidoptera:

Lycaenidae). Bulletin of the British Museum (Natural History), Entomology 13(4): 67-116.

Australian Entomologist, 2013, 40 (4): 175-176 " 175

AN OBSERVATION OF POLYURA SEMPRONIUS (FABRICIUS)

(LEPIDOPTERA: NYMPHALIDAE) FEEDING ON BANDICOOT

DROPPINGS IN SOUTH-EAST QUEENSLAND

T.J. SHAKESPEARE, Z.J. SHAKESPEARE and Т.Р. SHAKESPEARE

52 Serene Close, Mons, Qld 4556

Abstract

A specimen of Polyura sempronius (Fabricius, 1793) is recorded feeding on the droppings of a

long-nosed bandicoot, Perameles nasuta Geoffrey, 1804.

Discussion

Braby (2000) noted the occurrence of coprophagy in species of the

nymphalid subfamily Charaxinae and this behaviour is well known overseas,

particularly in Africa, where lion, leopard and otter droppings are widely

used. However, apart from a report of Polyura sempronius (Fabricius, 1793)

attracted to *animal refuse' (Burns and Rotherham 1969), there appears to be

no published documentation of coprophagy in Australian species.

ге

Fig. 1. Polyura sempronius feeding оп bandicoot dung.

At 1015h on 21 March 2013, we observed a specimen of Polyura sempronius

(Fig. 1) that had alighted on the droppings of a long-nosed bandicoot,

Perameles nasuta Geoffrey, 1804 (Mammalia: Peramelidae), on the lawn of a

residence in Mons, Queensland. It is estimated that the droppings were

between 6 and 12 hours old and the butterfly was observed on the same

dropping for 35 minutes, reinserting its proboscis every few minutes. The

176 Australian Entomologist, 2013, 40 (4)

proboscis tested the surface of the dropping by tapping and probing it for 1-8

seconds, before moving 1-5 millimetres and testing a different part of the

surface. This continued until a satisfactory part of the surface was identified

(perhaps a sufficiently soft or moist area), at which point the proboscis was

inserted into the dropping. Liquid was seen to be ingested via the proboscis.

Occasionally, the butterfly readjusted its position by walking slowly to

another part of the dropping and recommencing its exploration of the surface.

It has been previously noted that, when P. sempronius has been found at sap,

rotting fruit and animal refuse, it is easy to approach and capture (Burns and

Rotherham 1969). During the 35 minute period of observation, the present

specimen was not disturbed by the close proximity of a camera held within 5

cm of it, despite the flash being used on at least a dozen occasions. Touching

the butterfly also did not appear to disturb it in any way. Once feeding was

completed, it flew away and began hilltopping behaviour within a 20 metre

radius of where it had been feeding.

References

BRABY, M.F. 2000. Butterflies of Australia: their identification, biology and distribution.

CSIRO Publishing, Collingwood; xx + 976 pp.

BURNS, A. and ROTHERHAM, E.R. 1969. Australian butterflies in colour. A.H and A.W.

Reed, Sydney; 112 pp.

Australian Entomologist, 2013, 40 (4): 177-186 177

A NEW SPECIES OF DICALLANEURA BUTLER (LEPIDOPTERA:

LYCAENIDAE: RIODININAE) FROM PAPUA NEW GUINEA

R.B. LACHLAN! and CHRIS J. MÜLLER?

'Entomology Department, Australian Museum, 6 College St, Sydney, NSW 2010

?Papua Mining PLC, 5" Floor, 17 Hanover Square, London W1S 1HU, England (Address for

correspondence: PO Box 3228, Dural, NSW 2158: chrismuller999@gmail.com)

Abstract

Dicallaneura tennenti sp. n., from Western Province, Papua New Guinea, is described and

illustrated. Characters are provided to distinguish it from the sympatric D. ribbei Róber, D.

hyacinthus Toxopeus and D. pulchra (Guérin-Méneville).

Introduction

Parsons (1998) recorded nine species of Dicallaneura Butler from Papua

New Guinea and a further five species from Indonesian West Papua.

Toxopeus (1944) described additional species and subspecies from

Indonesian West Papua, although some of these are regarded as synonyms

(G. Lamas unpublished checklist). Parsons (1998) noted that *members of

Dicallaneura usually exhibit marked sexual dimorphism in their uppersides,

although their undersides may be used to effectively associate the two sexes.’

Both present authors have examined numerous specimens of all the species of

Dicallaneura from Papua New Guinea and concur with Parsons’

observations. We collected a single male and a single female of an

undescribed species from two different localities but, given their facies and

the fact that the two localities are within 15 km of each other, they are clearly

the same species. This new taxon is described here.

Dicallaneura tennenti sp. n.

(Figs 1-7)

Types. Holotype д, PAPUA NEW GUINEA: Matkomrae (Matkomnai) Catholic

Mission, 50 km NNW of Kiunga, W.P., 5?49'S 140?9'E, Alt. 60 m, 6.x.1993, К.В.

Lachlan (in Australian National Insect Collection, Canberra — Reg. No. 31-023121).

Paratype 9, PAPUA NEW GUINEA: 15 km NNE of Rumginae, Western Province,

80 m, 7.11.2008, C.J. Müller (in Natural History Museum, London).

Description. Male (Figs 1-2). Forewing length 21 mm. Antenna with dorsal

surface black, ventral surface with thin, white segmental bands, club

elongate, tip orange-brown. Dorsal surface of thorax and abdomen dark

brown, ventral surface creamy pale brown. Hindwing shape typical of

Dicallaneura, pronounced between veins CuA, and CuA;. Upperside ground

colour unicolourous very dark brown; forewing slightly darker with four

white, slightly elongated spots suffused with dark grey-brown scales in

subapical region in spaces between veins Rs and GOA, largest spot below

vein Rs, smallest above vein CuA;; basal area of inner margin of hind wing

pale creamy-brown. Forewing underside ground colour dark brown on

anterior half and terminal areas, with slight bluish tinge in central area, an

178 Australian Entomologist, 2013, 40 (4)

Figs 1-2. Dicallaneura tennenti sp. n., holotype male: (1) upperside; (2) underside.

orange-brown patch mostly below vein CuA, to mid point of cell above vein

1A+2A, two whitish spots at distal edge of patch, the uppermost small and

triangular, the lower and larger spot between veins CuA; and CuA, almost

rhomboid in shape; below this patch to dorsum light greyish-brown. Basal

Australian Entomologist, 2013, 40 (4) 179

Figs 3-4. Dicallaneura tennenti sp. n., paratype female: (3) upperside; (4) underside.

area of costa with thin, pale cream and orange-brown streak, close and

parallel a second whitish streak of similar length; from mid point of costa a

small, prominent, cream, basally curved thin bar crosses the discal cell to

cubitus, distally inside this curved bar is a small, thin, orange-brown mark

180 Australian Entomologist, 2013, 40 (4)

Figs 5-7. Dicallaneura tennenti, genitalia of holotype male: (5) dorsal view of

tegumen and uncus; (6) lateral view with aedeagus removed; (7) lateral view of

aedeagus.

then a larger, cream elongated spot curved inwardly on its basal edge; in the

dark central patch are four small spots, three in an oblique line from near the

costa to the subterminal region of the cell between vein М; and CuA;, the

Australian Entomologist, 2013, 40 (4) 181

costal spot whitish, the remaining three spots bluish-white; a thin, slightly

curved outwardly, subapical and subterminal line of whitish spots, absent

between veins М; and CuA;, from costa to inner margin; near the apex

between the veins are two small, slightly elongate bluish-white spots,

inwardly acute, rounded outwardly, with a black spot in the middle of the

uppermost spot, both surrounded by a thin line of orange-brown scales more

pronounced at their distal edge.

Hindwing underside ground colour orange-brown; base with prominent

cream streak; a small crescent-shaped marking near costa, curved basally

across discal cell; a small, less distinct cream spot distal of crescent marking;

above this spot a small streak at and parallel to costa; an irregular, broken,

cream marking from mid point of costa to vein M3; below cell four thin,

elongated creamy-orange streaks, partly along veins М; down to 1А+2А in

discal region; a darker brown, distally curved, irregular, thin subtornal band

between veins CuA, and 3A that is vestigial between veins CuA, and М»; the

brown band edged distally by a wider cream band between veins CuA, and

3A and basally by a similar width cream band from veins M; to 3A that

spikes sharply basally in the cell between veins 1A+2A and CuA;; two black

terminal spots in the cells between veins M; and М,» and veins CuA, and

CuA;, both with silver-blue scaling on their outer edges; two thin, silver-blue

bars between veins M; and CuA, and two more between veins CuA; and ЗА,

all edged lightly with black scales; two small, parallel, silver-blue streaks, the

larger adjacent to the apical black spot and the smaller in the cell between

veins M; and M3.

Male genitalia (Figs 5-7). In dorsal view, tegumen and uncus combined twice

as long as broad; uncus setose, in lateral view with sharp, apical, ventral hook

directed ventrally at 45°; brachia relatively long, gently curved upwards along

whole length, slightly wider basally then tapering to a fine point; saccus not

prominent, angle between saccus and vinculum noticeably obtuse; valvae not

large, relatively short, distal half narrow and clearly hirsute with some setae

relatively long at distal margin, very slightly concave at mid point of ventral

margin, dorsal margin slightly curved upwards, distally rounded; aedeagus in

lateral view long, slender, almost parallel sided, slightly sinuate along middle

two-thirds, distally tapering to a long, thin, slightly dorsally directed sharp

point at apex; ductus seminalis close to rounded end of coecum.

Female (Figs 3-4). Forewing length 25 mm. Antenna similar to male, length

14.5 mm. Head above and anteriorly light orange-brown, beneath cream; eye

ringed with white. Thorax and abdomen dorsally light orange-brown, ventral

surface and legs similar to male. Wings similar in shape to male except

termen more convex (tornal section of hind wing missing due to wing

damage). Forewing upperside ground colour uniform light orange, apical area

broadly dark brown, extending from midway along costa to junction of vein

Мз with cell and across to tornus, boundary with ground colour

182 Australian Entomologist, 2013, 40 (4)

approximately parallel to termen but distinctly stepped at veins M3 and CuA,

and dark border reduced to 1.5 mm between veins CuA; and 1A+2A.

Forewing underside very similar to male. Hindwing upperside ground colour

as for forewing, costa and inner margin slightly paler orange, tornus along

inner margin with light brown dusting, two subapical dark brown spots

between veins near termen. Hindwing underside as in male.

Etymology. This new species is named after John Tennent, a long-standing

scientific associate of The Natural History Museum, London. During his long

and outstanding career in the field of entomology, he has published a large

number of books and scientific papers and is regarded as a world authority on

the butterflies of the Pacific Islands.

Distribution. At present, D. tennenti is known from only two specimens taken

at two localities, approximately 15 km apart, north of Kiunga in Western

Province, Papua New Guinea.

Discussion

Although D. tennenti exhibits no sign of blue on its upperside, it most closely

resembles three species of Dicallaneura recorded from Papua New Guinea:

D. ribbei Röber, D. hyacinthus Toxopeus and D. pulchra (Guérin-Méneville).

However, D. tennenti can be readily distinguished from D. hyacinthus as the

latter species has a plain dark blue upperside and a darker underside pattern.

Similarly, D. ribbei also has a plain upperside with an obvious blue sheen and

constant differences in its underside maculation when compared with D.

tennenti. The third species, D. pulchra (Figs 8-11), has a distinctive whitish-

blue, oblique bar across the forewing in males. More than 70 males of D.

pulchra were examined and none showed any significant variation in the

width, length and shape of this character. Dicallaneura tennenti clearly

differs by having four white, slightly elongated spots in the subapical region

of the forewing, a character not seen in any other species of Dicallaneura.

Comparison of the underside maculation of D. tennenti with that of D.

pulchra revealed several differences. In D. tennenti the forewing termen is

slightly straighter and the small, nearly straight, oblique, pale orange-brown

bar between the two basally curved cream markings near the costa is more

prominent. The cubitus vein at the base of the discal cell is without the clear

streak seen in the vast majority of D. pulchra specimens examined. The small

triangular spot seen in D. pulchra, just above the cubitus vein in the discal

cell, is absent in D. tennenti but the small, whitish postmedian spot between

veins М; and CuA, seen in D. tennenti is not evident in D. pulchra. On the

hind wing, the median, cream, broken oblique line from the costa to vein М;

is reduced in D. tennenti but larger and clearly crescent shaped in D. pulchra.

The dark brown, thin subtornal band in D. tennenti largely terminates at vein

CuA, but in D. pulchra it clearly extends to vein M3. The underside cream

markings of D. tennenti are noticeably whiter in D. pulchra.

Australian Entomologist, 2013, 40 (4) 183

Figs 8-9. Dicallaneura pulchra male: (8) upperside; (9) underside.

Dicallaneura pulchra was described from a single female (Guérin-Méneville

1830-1838), although Parsons (1998) referred to a ‘holotype male’. The

location of this type is unknown but the diagnostic characters of the species

are clearly evident in the original illustrations [pl. 16, figs 2-3].

184 Australian Entomologist, 2013, 40 (4)

Figs 10-11. Dicallaneura pulchra female: (10) upperside; (11) underside.

The male genitalia of D. tennenti (Figs 5-7) differ from those of D. pulchra

(Figs 12-14) and D. ribbei in dorsal view, in having a much longer and

narrower tegumen and uncus combined, as seen in Fig. 5. In D. pulchra the

uncus is shorter with a more protruding tip (Fig. 12). In lateral view, the tip

Australian Entomologist, 2013, 40 (4) 185

Figs 12-14. Dicallaneura pulchra male genitalia: (12) dorsal view of tegumen and

uncus; (13) lateral view with aedeagus removed; (14) lateral view of aedeagus.

of the uncus is clearly more ventrally directed than in D. tennenti. A

significant character seen in D. pulchra is the unusually dense, long setae

covering the uncus and valvae; this is also evident on D. tennenti but the

186 Australian Entomologist, 2013, 40 (4)

setae are shorter and less dense. The brachia in D. tennenti are longer than in

D. pulchra and the saccus appears less developed with a more sloping angle

between the saccus and vinculum than seen in D. pulchra. The valvae in D.

pulchra are basally more bulbous than in D. tennenti, while in the aedeagus

the ductus seminalus is slightly further away anteriorly from the coecum than

in D. tennenti and tapers to a shorter, thicker point.

The female of D. tennenti is almost identical to that of the male on the

underside. The upperside is similar to that of females of several other

Dicallaneura species, being light orange with a broad apical border on the

forewing. When compared with the female of D. pulchra, D. tennenti has

slightly more extensive orange on the forewing and is a paler orange.

A clear difference seen in both males and females of D. tennenti is found in

the postmedian cream band between veins M; and 3A, basad of the thin dark

brown line. Between veins 1A+2A and CuA, this cream band spikes sharply

basally, a condition not seen in either D. pulchra or D. ribbei. The male

forewing of D. tennenti (21 mm: n = 1) is slightly larger than either D. ribbei

(19.5-20.5 mm: n = 6) or D. pulchra (18.5-19 mm: n = 4).

In the vicinity of Kiunga, including the site for the paratype female of D.

tennenti, several other Dicallaneura taxa were recorded by the authors,

namely D. decorata (Hewitson), D. kirschi Róber, D. hyacinthinus, D. ribbei

and D. pulchra. The female specimen of D. tennenti was taken as it alighted

on foliage above a steep, shallow ravine about four metres above the ground.

Several Dicallaneura taxa are known from just a few specimens from widely

disjunct localities. It is therefore likely that the new species has a wider

distribution in mainland New Guinea.

Acknowledgements

The authors thank Ted Edwards (ANIC, Canberra) for his dissection and

preparation of the male genitalia and helpful comments, You Ning Su (ANIC,

Canberra) for the genitalia digital images and John Tennent for supplying

obscure literature, numerous photographs and helpful advice.

References

GUÉRIN-MÉNEVILLE, F.E. 1830-1838. Insectes. In Duperry, L.I. (ей), Voyage autour du

monde: exécuté par ordre du roi, sur la corvette de Sa Majesté, la Coquille, pendant les années

1822, 1823, 1824, et 1825: sous le Ministére et conformément aux instructions de S.E.M. le

Marquis de Clermont-Tonnerre, ministre de la marine; et publié sous les auspices de son

excellence mgr le cte de Chabrol, ministre de la marine et des colonies (1825).

PARSONS, M.J. 1998. The butterflies of Papua New Guinea: their systematics and biology.

Academic Press, London; 736 pp, xxvi + 104 pls.

TOXOPEUS, L.J. 1944. Results of the Archbold Expedition to New Guinea. Iho (Treubia) 1-1

2604: 156-192.

Australian Entomologist, 2013, 40 (4): 187-218 187

A REVISION OF THE EUPLOEA BATESII C. & R. FELDER, 1865

‘COMPLEX’ (LEPIDOPTERA: NYMPHALIDAE: DANAINAE) IN

MAINLAND PAPUA NEW GUINEA AND AUSTRALIA, INCLUDING

ITS BIOLOGY AND BIOGEOGRAPHY

TREVOR A. LAMBKIN

School of Biological Sciences, University of Queensland, St Lucia, Qld 4072

Abstract

The biogeography, biology and taxonomy of the Euploea batesii C. & В. Felder, 1865 ‘complex’

from mainland Papua New Guinea and Torres Strait were studied and all specimens examined

were grouped into one of three forms or morphological types. Immature stages of the immaculate

form (Morph type 1) are described and illustrated from material collected on Mer Island, Torres

Strait, where the larval host plant was Parsonsia velutina R.Br. (Apocynaceae). On Mer Island,

adults of several types were found in large numbers imbibing pyrrolizidine alkaloids from dead

and damaged leaves and flowers of Heliotropium foertherianum Diane & Hilger (Boraginaceae).

The three morphological types were found to have distinctive sympatric and allopatric

distributions, which might be determined by larval host plant specificity. Broadly, all three types

occur in eastern Papua New Guinea and on Mer Island in Torres Strait, while only the

immaculate type is known elsewhere in New Guinea and on other Torres Strait islands. On Mer

Island, captive mothers of the immaculate Morph type 1 produced final instar larvae that were

charcoal black in colour and resulted in only immaculate Morph type 1 (F1) offspring; these F1

were not discernibly different from nominotypical E. batesii from Indonesian West Papua. The

distributions of the maculated types (Morph types 2 and 3) and several other butterfly species

which are only known to occur in Torres Strait on Mer Island, Support a probable link in the

geological history of Mer Island and eastern Papua New Guinea. In addition, the distribution of

the immaculate form indicates a link in the geological history of the islands in the west of Torres

Strait to the southern region of mainland Papua New Guinea. The E. batesii ‘complex’ likely

includes more than one species, with perhaps more species occurring through the major islands

and archipelagos along the Solomon chain east of New Guinea. Based on the distinct

distributional and wing morphological differences occurring in populations in Papua New

Guinea and Torres Strait, plus life history evidence, it is proposed that E. b. resarta Butler be

revised to species status (E. resarta Butler, 1876, stat. rev.) and that E. b. squalida (Butler) be

revised to a form of E. resarta (E. resarta f. squalida (Butler, 1878), stat. rev.), both taxa

occurring together exclusively in eastern mainland Papua New Guinea and on Mer Island. Also,

the immaculate monotypic populations (Morph type 1) occurring on mainland New Guinea and

its outlying islands and in Torres Strait are proposed as new synonyms of E. b. batesii.

Introduction

Euploea batesii C. & R. Felder, 1865 is currently classed as a polytypic

species found principally in the New Guinea region, i.e. from Buru, Seram

and Ambon in the west, across New Guinea and its outlying islands, to the

Solomons in the east (Ackery and Vane-Wright 1984, Parsons 1998, Tennent

20022). South of New Guinea it extends to the Aru Islands in the Arafura Sea

(Ackery and Vane-Wright 1984) and into Torres Strait in Queensland,

Australia (Braby 2000). In Papua New Guinea it is widespread, occurring in

marginal secondary vegetation and forests up to 1200 m (Parsons 1998). Its

distribution across the bulk of New Guinea is poorly known and there are few

specimens from this region in Australian collections. In Torres Strait, Е.

batesii occurs mostly in marginal, dense primary or secondary monsoon vine

forest, but can frequent overgrown and abandoned village gardens.

188 Australian Entomologist, 2013, 40 (4)

The life history of E. batesii was partially described from material collected

near Bulolo in Papua New Guinea by Parsons (1998), where he recorded

Parsonsia lata Markgr. and Hoya R. Br. sp. (Apocynaceae) as larval hosts,

while Szent-Ivany and Carver (1967) reported larvae occurring commonly on

the ornamental Nerium oleander L. (Apocynaceae) at Port Moresby, Papua

New Guinea.

Felder and Felder (1865) described the syntypes of E. batesii from Dodinga,

Halmaheira, in the Northern Moluccas (Edwards et al. 2001); several authors

since have expressed doubts concerning the authenticity of this type location

(Ackery and Vane-Wright 1984, Parsons 1998). This uncertainty was

primarily based on the fact that ‘type’ labels are not attached to any Е. batesii

specimens in the Felder collection (Talbot 1943) and, moreover, the E.

batesii ‘complex’ is otherwise not known from Halmaheira. In addition,

Ackery and Vane-Wright (1984) provided circumstantial evidence that a

female syntype (in the Felder collection) was, with another syntype of Е. b.

batesii, in the same batch of Felder specimens from Ambon in the Southern

Moluccas. Ambon is just west of New Guinea, from which the current

designated distribution of the E. batesii ‘complex’ extends eastward to the

Solomon Islands (Fruhstorfer 1910, Corbet 1942, Ackery and Vane-Wright

1984, Parsons 1998, Tennent 20022).

Ackery and Vane-Wright (1984) listed 24 names that have been applied to Е.

batesii at various times, while D’Abrera (1978) listed 13 subspecies from the

region, with four subspecies purportedly occurring on mainland Papua New

Guinea (Parsons 1998). Tennent (2002a) recorded an additional five

subspecies from the Solomon Islands. Corbet (1943) placed E. batesii in the

E. climena (Stoll) species group, indicating that members of this group are

characterised by the presence of a recurrent vein in the forewing cell and, in

males, the absence of a pale raised patch in the cell of the hindwing upperside

and the absence of an androconial brand on the forewing upperside. Corbet

(1943) and Carpenter (1953) also pointed out that males of E. batesii could

then be separated from their congeners within the climena group by

possessing, on the forewing underside, an elongate white stripe in the anterior

half of space 1b and a narrower white stripe below it. These stripes may be

pale or dark and are also used to distinguish female E. batesii. Conversely,

Ackery and Vane-Wright (1984), in their cladistic study of the Danainae,

placed E. batesii in a clade separate from E. climena but admitted that all of

the species within the same clade containing E. batesii were difficult to

characterise. Equally, Parsons (1998) believed that the taxonomy of Е. batesii

was formidable and in need of revision, especially due to the high degree of

variation recorded in wing pattern across its range and the possibility that Е.

batesii might comprise a species complex similar to other suspected

complexes within Euploea Fabricius (Ackery and Vane-Wright 1984, De

Baar 1991).

Australian Entomologist, 2013, 40 (4) 189

Here I review the distribution and polymorphism of the E. batesii ‘complex’

on mainland Papua New Guinea (PNG) and in Torres Strait, Australia,

describe and illustrate the life history of the immaculate morph (Morph type

1) from Mer I. in eastern Torres Strait, determine the larval host plant on Mer

I. and discuss its general ecology. I also document the species’ known

distribution on the island of New Guinea, the archipelagos east of New

Guinea and in Torres Strait, and analyse and discuss the polymorphism

(degree of wing spotting) of the complex from Torres Strait, mainland PNG

and its outlying islands. The results of these analyses, together with life

history data, are used to propose the reinstatement of E. resarta Butler, 1876

(stat. rev.), which occurs exclusively in eastern mainland PNG, on several of

its outlying islands and on Mer I. In addition, in eastern mainland PNG and

on Mer I. it is proposed that this species occurs sympatrically with E. batesii

batesii, with the latter taxon also occurring across mainland New Guinea and

several of its outlying western islands and on several islands in Torres Strait.

Material and methods

Life history and ecology studies on Mer Island, Torres Strait

An extensive search was undertaken for immature stages in January 2011

along the edges of monsoon vine forest on Mer I. (9°55°S, 144?30'E) (Fig.

61), the largest of three basaltic islands collectively called the Murray Island

group, which are the remains of volcanic cones (Willmott 1972). Natural

vegetation consists of tropical deciduous monsoon forest. Immature stages of

E. batesii were found only on Parsonsia velutina R.Br. (Apocynaceae),

following which seven unspotted, dark brown to black adult females were

collected and released in an abandoned shade house (2 m wide, 4.m long, 2m

high), which served as a makeshift flight cage. Long stems of the larval host

plant, plus sprigs of flowering lantana (Lantana camara L. [Verbenaceae]),

which were used as a nectar source by adults, were placed in long-necked jars

of water suspended from the ceiling of the shade house using wire. All early

stages collected were reared at ambient conditions in round, clear plastic food

containers (volume 280 mL, height 75 mm, bottom radius 42.4 mm, top

radius 55 mm). Larvae were fed on fresh host plant introduced daily. Three

adults were reared from field-collected material and 14 from eggs laid in the

improvised flight cage.

Analysis of specimens from mainland Papua New Guinea and Torres Strait

Specimens of the E. batesii ‘complex’, predominantly from mainland Papua

New Guinea and its nearby islands (n = 92) and Torres Strait (n = 154), were

examined and grouped into three morphological types, which were then

further grouped relative to their collection sites. In addition, photographs of

the genital armature of four dark brown to black males of E. batesii (from

Dauan I., Torres Strait x 2; Brown R. near Port Moresby, Papua New Guinea;

and Yapen, Indonesian West Papua) were compared with those of four well-

spotted males (from Mer I. x 2; and Bulolo, Papua New Guinea x 2), using a

190 Australian Entomologist, 2013, 40 (4)

stereo microscope and image stacking software (Helicon Focus 5.3®, 2012),

in order to determine if morphological differences in wing pattern

corresponded with any differences in genital structures, with nomenclature

used following Ackery and Vane-Wright (1984) and Monastyrskii (2011).

Abbreviations of collectors’ names appearing on specimen labels and

collection depositories are: A&GD — A. and G. Daniels; AIK — A.I. Knight;

AM — Australian Museum, Sydney; ANIC — Australian National Insect

Collection, Canberra; AT — A. Tubb; CWWTC – C.W. Wyatt theft collection

(AM); DKM - D.K. McAlpine; EJLH — E.J.L. Hallstrom; EM — E. Mann;

GAWC — G.A. Waterhouse collection (AM); GLC — С. Lyell collection

(AM); GMC - G.M. Carson; GRF - G.R. Forbes; GRFC - G.R. Forbes

collection, Brisbane; GW — С. Wood; GW (ANIC) — С. Wood (Long I., -

.1.1956 locality); HE — H. Elgner; HR — H. Rauber; IFTA - Insect Farming

and Trading Agency, Bulolo, Morobe Province; ТОС — J.G. Grimshaw; JOC

— J.O. Campbell; MDB ~ M. De Baar; MDBC - M. De Baar collection,

Brisbane; MM — Macleay Museum, Sydney; MTQ — Museum of Tropical

Queensland, Townsville; MW — M. Willows jnr; N&SHC — N. and S. Hunter

collection (AM); OM — O. McCaw; PSH - P.S. Hanahan; QDAFFC —

Queensland Department of Agriculture, Fisheries and Forestry collection,

Brisbane; QM — Queensland Museum, Brisbane; RC — К. Carver; SGC - S.

Ginn collection, Sydney; SJJ&IRJ — S.J. and LR Johnson; TAL – Т.А.

Lambkin; TLFC — T.L. Fenner collection, Brisbane; TLIKC — Joint collection

of T.A. Lambkin and A.I. Knight, Brisbane; VRC — van Raalte collection

(AM); WIP — W.I. Potter; WWB — W.W. Brandt.

Life history and biological studies on Mer Island, Torres Strait

Host plant: Parsonsia velutina R.Br. (Apocynaceae).

Egg (Fig. 1): (n = 10); typical of Euploea spp; bullet-shaped; whitish-yellow;

basal half of egg relatively smooth with outlines of square recessed dimples,

6 dimples high, each dimple outlined with vertical columns and diagonal

rows; apical half of egg with hexagonal dimples, 4-5 dimples high, hexagons

becoming smaller at apex; within a few days becoming yellow.

First instar larva (Fig. 2): (n = 16); head black; body cylindrical, smooth,

glossy and semitranslucent; lime-green, abdominal segment 8 orange; a pair

of slightly raised carmine protuberances on mesothorax, metathorax and

abdominal segments 2 and 8; bases of legs and prolegs same colour as body,

legs and prolegs black.

Second instar larva (Fig. 3): (n = 20); head and body similar to first instar,

except body not translucent; yellow-orange; spiracles on prothorax and

abdominal segment 8 black; a pair of blunt, black filaments shorter than

width of body on mesothorax, metathorax and abdominal segments 2 and 8,

with pair on mesothorax slightly longer than others; bases of legs and prolegs

same colour as body, legs and prolegs black.

Australian Entomologist, 2013, 40 (4) 191

Figs 1-8. Early stages of the Euploea batesii ‘complex’ from Mer Island, Torres Strait

(Morph type 1 = Е. b. batesii): (1) egg (height 1.8 mm); (2) Ist instar larva (length 6.5

mm); (3) 2nd instar larva (14 mm); (4) 3rd instar larva (25 mm); (5) 4th instar larva

(30 mm); (6) 5th instar larva (45 mm); (7) 5th instar showing prominent orange collar

(45 mm); (8) pupa (height 22 mm).

192 Australian Entomologist, 2013, 40 (4)

Third instar larva (Fig. 4):(n — 20); similar to second instar except whole

body and head tomentose; body dull in colour; filaments longer, roughly the

width of the body; anal segment black; all spiracles black with those on

prothorax and abdominal segment 8 larger than others.

Fourth instar larva (Fig. 5): (n = 20); similar to third instar except body

glossy and semi-translucent; filaments longer, roughly 1.5 x width of body.

Fifth instar larva (Figs 6-7): (n = 18); similar to fourth instar except entire

body and head tomentose; body, including filaments, dull charcoal-black

except for a semi-circular bright orange dorsal collar on prothorax; head,

legs, prolegs and spiracles glossy black; all filaments roughly twice the width

of the body.

Pupa (Fig. 8): (n = 18); typical of Euploea spp; entirely amber at first; after 2

days abdomen, wing cases, thorax, eyes and antennae change to shining

silver; buff markings on abdomen and wing-cases; spiracles pale brown.

The early stages of the E. batesii ‘complex’, specifically of the black

immaculate form, were discovered on P. velutina, as per Stanley and Ross

(1986) and Harden et al. (2007). Parsonsia velutina was the only Parsonsia

species found on the island. Overall, the early stages were difficult to find as

the host plant predominantly grew out of reach, high in the forest canopy

(Fig. 9). Eventually, eggs and early instar larvae were found on two large

vines, only on the soft growing tips of the host plant, but larger larvae were

found on, and fed freely on, the harder, more mature leaves and, when

ultimate, fed on flower buds and racemes (Figs 7, 9). Interestingly, final

instar larvae superficially resemble, in both colour and form, final instar

larvae of Ornithoptera priamus poseidon (Doubleday, 1847) (Lepidoptera:

Papilionidae), which also occurs on the island.

During the wet season in Torres Strait (normally between December and

April), adults of E. batesii are frequently observed on Mer I., where they

regularly nectar throughout the day from Lantana camara, often with E.

algea amycus Miskin, 1890 and E. tulliolus tulliolus (Fabricius, 1793). In

addition, males of all three Euploea spp, as well as males of Tirumala hamata

hamata (W.S. Macleay, 1826), were found throughout the day, frequently in

large numbers, imbibing pyrrolizidine alkaloids from dead and damaged

leaves, and flowers, of Heliotropium foertherianum Diane & Hilger

(Boraginaceae) (known also as the heliotrope tree, Tournefortia argentea)

(Figs 10-11) growing in sand on beaches and behind dunes (Fig. 12).

Similarly, adults of T. ^. hamata were also found to be attracted to flowers

and leaves of Crotalaria retusa L. (Fabaceae). Females of E. batesii were

often observed flying along margins of vine thickets, in undergrowth and

through abandoned gardens, but were generally secretive and rarely observed

when ovipositing. Overall, the adult butterflies have a slow, gliding flight

with long wing beats.

Australian Entomologist, 2013, 40 (4) 193

Ur n 7

Figs 9-10. (9) Parsonsia velutina (Apocynaceae): host plant of the Euploea batesii

‘complex’ (Morph type 1 = £. b. batesii) on Mer Island, Torres Strait; (10) adults of

the Euploea batesii ‘complex’ (Morph type 1 = E. b. batesii) clustered with E. algea

amycus, E. tulliolus tulliolus and Tirumala hamata hamata imbibing pyrrolizidine

alkaloids from stems and dead leaves of Heliotropium foertherianum (Boraginaceae).

194 Australian Entomologist, 2013, 40 (4)

Figs 11-12. (11) Damaged leaves and immature fruit of Heliotropium foertherianum

(Boraginaceae), Mer Island, Torres Strait; (12) mature H. foertherianum growing on

sand with swarming adults of Euploea spp and T. tirumala imbibing pyrrolizidine

alkaloids.

Australian Entomologist, 2013, 40 (4) 195

Caged females commenced ovipositing 24 hr after being released into the

flight cage. All mature larvae that were reared (i.e. collected directly from the

field off P. velutina and from caged mothers) were as illustrated in Figs 6-7.

These larvae showed no variation in morphology or colour. In addition, all

successfully reared offspring (F1) (n = 17) (e.g. Figs 13-14) originating from

these larvae were similar (n = 14) in markings to their mothers (n = 7) (e.g.

Fig. 15) that were released into the flight cage, ie. dark brown to black,

devoid of white spotting (immaculate, Morph type 1 form). Overall, mothers

and reared F1 were not discernibly different from what could be considered

nominotypical E. batesii from Yapen in Indonesian West Papua (Fig. 16).

Taxonomy

Material examined or reviewed

QUEENSLAND: 15, Cape York (Lectotype of E. b. belia Waterhouse & Lyell, 1914;

illustrated in Waterhouse and Lyell [1914] Fig. 10); 19, Cape York, ex GAWC (AM);

13, 19, Australia (MM); QUEENSLAND (TORRES STRAIT): 14, Mer (or Murray)

Island, 5.ix.1907 ex GAWC, passed through CWWTC, 1946-47 (AM); 1555, 799,

same data except 29.iii-4.iv.1986, TAL (TLIKC); 19, same data except (QM); 959,

699, same data except 22-25.iv.1989 (TLIKC); 2955, 1399, same data except

13.1.1994 (288), 14.1.1994 (254), 15.1.1994 (288), 12.v.1994 (18), 9.iii.1995

(633, 699), 28.1.2011 (4509, 19), 29.1.2011 (734, 399), larva coll. 29.1.2011

(16), em. 6.11.2011 (19), em. 20.11.2011 (19), em. 1.11.2011 (18), em. 4.11.2011

(19), em. 5.11.2011 (16), em. 11.11.2011 (15), em. 18.11.2011 (15) TAL (TLIKC);

638, 799, same data except TAL & AIK 25.1.2011 (344, 19), 26.1.2011 (254,

599), 29.12011 (15, 19); 755, 299, same data except AIK 25.iv-5.v.1999 (655,

29 9), 27.11.2000 (14); 19, same data except 30.11.1986 (QM); 1359, 129 9, same

data except 29.iii-4.iv.1986, MDB (MDBC); 15, 19, same data except (TLFC); 359,

same data except (QM); 244, 19, same data except 26.iv.1984 (19), 30.iv.1984

(14), 3.v.1984 (15), GW (QM); 19, Erub (or Darnley) Island, ex GAWC, passed

through CWWTC, 1946-47 (AM); 16, Iama (or Yam or Turtleback) Island,

7.iv.1987, MDB (TLIKC); 19, same data except 11-12.vi.1992, AIK; 484, 19,

Dauan Island, 20.11.2004 (14), 6.1.2006 (355, 19) TAL (TLIKC); 6595, 19, same

data except AIK 18.xii.2005 (14), 21.xii.2005 (18), 24.xii.2005 (14), 29.xii.2009

(1d), 16.1.2004 (18), 2.1.2010 (14), 16.1.2011 (19); 19, same data except 09.228,

142.39E, 2-4.111.2013, SJJ&IRJ (МТО). ;

PAPUA NEW GUINEA [Province names abbreviated as in Table 1]: 19, Aitape

[SP], 26.viii.1944, JOC (VRC), (AM); 13, Angoram [ESP], 20 ft, 28.iv.1950, WB &

EJLH (ANIC); 14, Balamuh [WP], 9.v.1992 JGG (QDAFFC); 24d, 399, Brown

River, nr Port Moresby [CP], 30.vii.1968 (15), 18.x.1968 (19), 20.x.1968 (19),

18.1.1976 (19), 28.1.1976 (15) СКЕ (GRFC); 659, 299, same data except

17.xii.1967, 26.1.1968, 7.iv.1968, 17.vi.1968, 15.viii.1968, 23.ix.1968, 1.xi.1967

(19), 11.11.1968 (19) HR (QM); 8 3, 699, Bulolo [MP], -.х.1984 (14, 19),

-xii.1984 (299), 1.1985 (14, 19), -.111.1985 (16), -.v.1985 (488, 19), -.11.1987

(19), -.vii.1987 (15) IFTA (MDBC); 209, same data except 11.viii.1980, -.vi.1991

(SGC); 24 4, Erora Creek, nr Oro Bay [OP], -.v.1943, -.vi.1943 AT (AM); 19, Kapa

Kapa [CP], 16.iv.1972 GRF (GRFC); 255, same data except 20.xi.1967, 7.iv.1968

HR (QM); 14 Lae, [MP], 1.xi.1973 СКЕ (GRFC); 15, same data except 24.xii.1963

196 Australian Entomologist, 2013, 40 (4)

(DKM); 1144, 19, same data except 4.vi.1951 (14), 12.vi.1951 (14), 17.ix.1951

(14), 18.1х.1951 (13), 20.ix.1951 (13), 30.ix.1951 (19), 2.x.1951 (15), 3.x.1951

(18), 4.x.1951 (13), 10.x.1951 (259), 21.x.1951 (14), (WB & EJLH) (ANIC);

29 9, Langemak [MP], 20.iii.1944, 24.11.1944, JOC (VRC), (AM); 19, Laloki River,

nr Port Moresby [CP], -.vi.1943, AT (AM); 15, 19, Long I., [MBP], -.i.1956 GW

(ANIC); 12 Morobe G [MP], WIP (AM); 1d, Mt. Lamington Dist, Northern

Division [OP], -.vii.1927, (AM); 399, Munum Waters, 15 ml W of Lae [MP]

18.1.1971, A&GD (AM); 14, Paga Hill, Port Moresby [CP], 15.viii.1966 EM (QM);

233, 299 Port Moresby, [CP], 20.1.1927 (18), 27.111.1927 (18), 28.1.1927 (29 9)

(AM); 2594, same data except no date RC (AM); 14, same data except ‘purchased for

Gerrard’ (AM); 19, same data except 20.11.1966 RC (AM); 254, same data except

28.11.1964, 23.v.1965 HR (QM); 15, same data except 7.vi.1964 HR (QM); 19, same

data except no date HR (QM); 15, Rigo [CP], 28.xi.1965 HR (QM); 455, 399,

Sambio Mumeng [MP], -.i.1983 (8), -.xii.1984 (13), -.1.1985 (4, 399), -.111.1985

(d) IFTA (MDBC); 14, Sambio [MP] PNG, 20.xii.1980 (SGC); 14, Sogeri [CP],

12.1.1968 СКЕ (GRFC); 26d, 299, Subitana, 1800 ft [CP], 11.xii.1949 (14),

12.xii.1949 (15, 19), 14.xii.1949 (19) WWB & EJLH (ANIC); 355, Wakaiuna,

Sewa Bay, Normanby І. [MBP] 23.х.1956-11.1.1957 WWB (ANIC); 14, Wau [MP],

1200m, -.iv.1974, collection PSH (MDBC); 14 (Е. batesii resarta) Nivani Island,

Louisiade Archipelago [MBP], GMC, 5.iv.1912, illustrated in: Carpenter (1953), Plate

5, Fig. 5.

SOLOMON ISLANDS: 14 (Е. batesii kunggana Holotype), Kunganna Bay, Rennell

Island [Rennell and Bellona Province], Templeton-Crocker expedition, MW,

6.vi.1933, illustrated in Carpenter (1953), Plate 5, Fig. 1; 19 (E. batesii kunggana

paratype), same data except illustrated in Carpenter (1953), Plate 5, Fig. 4.

INDONESIAN WEST PAPUA: 255, Yapen Island, Irian Bay, -.х.1995, -.x.1997

(MDBC); 15 Wangaar, Irian Jaya, illustrated іп: Ackery and Vane-Wright (1984),

Fig. 272, p 384.

MOLUCCAS: 19, Buano Island, W. of Seram, Jan-Feb.1992 (MDBC).

Analysis of morphological types

For analyses, all specimens were grouped into three morphological types,

illustrated in monochrome in Figs 13-36.

Morph type 1: Taxon batesii

Euploea batesii C. & R. Felder, 1865: 331.

E. b. batesii (Fruhstorfer 1910: 232); E. b. batesii (D’Abrera 1978: 179); E. b.

batesii (Ackery and Vane-Wright 1984: 234); Е. b. batesii (Parsons 1998: 520); Е.

b. batesii (Edwards, Newland and Regan 2001: 311).

Euploea batesii belia Waterhouse & Lyell, 1914: 22.

E. b. belia (Waterhouse 1932: 78); E. b. belia (Carpenter 1953: 88); E. b. belia

(Common 1964: 58); E. b. belia (McCubbin 1971: 14); E. b. belia (Common and

Waterhouse 1972: 225); E. b. belia (D' Abrera 1978: 178); E. b. belia (Common

and Waterhouse 1981: 308); E. b. belia (Ackery and Vane-Wright 1984: 234); Е.

E ed (Scheermeyer 1999: 192); E. b. belia (Edwards, Newland and Regan

1: 311).

Australian Entomologist, 2013, 40 (4) 197

Figs 13-20. Euploea batesii ‘complex’, Morph type 1 =. b. batesii (all figures not to

scale, upper side left, under side right): (13, 16, 17) males: (13) Mer I., Torres Strait,

em. 11.11.2011, TAL, F1 [forewing length 40 mm]; (16) Yapen, Irian Bay, 1995, [44

mm]; (17) Murray (Mer) I., Torres Strait, 22-25.iv.1989, TAL, [43 mm]; (14, 15, 18-

20) females: (14) Mer I., em. 4.11.2011, TAL, ЕІ [39 mm]; (15) Mer I. (mother),

29.1.2011, TAL, [46 mm]; (18) Mer I., em. 6.11.2011, TAL, [43 mm]; (19) Murray 1.,

9.11.1995, TAL, [46 mm]; (20) Mer I., 29.1.2011, TAL, [46 mm].

198 Australian Entomologist, 2013, 40 (4)

Wings, upper and underside, dark brown to black ground colour; all wings

with varying degrees of pale outer margins, especially on underside; upper

and undersides of wings almost always devoid of marginal white spotting

(Figs 13-20).

Morph type 2: Taxon resarta

Euploea resarta Butler, 1876: 241.

Crastia resarta (Butler 1878: 298); E. resarta (Fruhstorfer 1910: 234); E. batesii

resarta (Carpenter 1953: 88); E. b. resarta (D'Abrera 1978: 178); E. b. resarta

(Ackery and Vane-Wright 1984: 234); E. b. resarta (Wood 1987a: 39); E. b.

resarta (Lambkin and Knight 1990: 108); E. b. resarta (Parsons 1998: 520); E. b.

resarta (Braby 2000: 611); E. b. resarta (Edwards, Newland and Regan 2001:

311); C. resarta (Edwards, Newland and Regan 2001: 311).

As for Morph type 1, except, not all individuals with pale outer margins and

upper and underside with bright, white subterminal bars, with or without

white terminal spots, including bright, white subapical forewing bars, all

variable in size; spotting ranging from predominantly on hind wing only and

sometimes forewing only, to being present on all wings (Figs 21-28).

Morph type 3: Taxon squalida

Crastia squalida Butler, 1878: 298.

Euploea squalida (Fruhstorfer 1910: 235); E. batesii squalida (Carpenter 1953:

88); C. squalida (Edwards, Newland and Regan 2001: 311).

As for Morph type 2, except spots and bars overlaid variably with dark

ground-colour scales, giving them a smoky appearance; Corbet (1942)

referred to this as ‘obsolescence of the white spots’ (Figs 29-32).

Carpenter (1953) grouped the E. batesii ‘complex’ in table format,

assembling each taxon based on essentially similar layouts of spots as in this

revision (Table 1), ie. groupings based on absence (Morph type 1), or

presence of ‘admarginals’ (terminal spots) and ‘submarginals’ (submarginal

and subterminal spots and bars) (Morph type 2). In addition, Carpenter

(1953) categorised the other group (Morph type 3) as having ‘obsolescence of

the white spots’ and having ‘faint indications of submarginals’.

The review of specimens and their data (Table 1) showed that all those from

Australia were from Torres Strait, except for a pair of doubtful origin

purportedly collected at Cape York (Table 1). Of the Torres Strait specimens

examined, the majority were from Mer I. (138 of 154), where it occurs as a

polymorphic population ranging from dark brown or black, predominantly

immaculate individuals without white spotting [Morph type 1 (E. b. batesii-

like: Figs 13-15, 17-20) (61 of 138; 44.2% of specimens examined)], to

specimens with variable amounts of bright white spots [Morph type 2 (E. b.

resarta-like: Figs 21-28) (55 of 138; 39.9% of specimens examined)],

Australian Entomologist, 2013, 40 (4) 199

АШ

Figs 21-28. Euploea batesii ‘complex’ from Torres Strait, Morph type 2 = Е. resarta

stat. rev. (all figures not to scale, upper side left, under side right): (21-23, 25, 27)

males: (21) Mer I., 25.1.2011, TAL & AIK, [forewing length 42 mm]; (22) Mer I.,

29.1.2011, TAL & AIK, [44 mm]; (23) Murray (Mer) I., 29.iii-4.iv.1986, TAL, [44

mm]; (25) Murray I., 25.iv-5.v.1999, AIK, [46 mm]; (27) Murray I, 9.11.1995, TAL,

[45 mm]; (24, 26, 28) females: (24) Murray I., 29.iii-4.iv.1986, TAL, [43 mm]; (26)

Murray I., 9.iii.1995, TAL, [46 mm]; (28) Murray I., 9.11.1995, TAL, [46 mm].

200 Australian Entomologist, 2013, 40 (4)

Figs 29-36. Euploea batesii ‘complex’ from Torres Strait. (29-32) Morph type 3 = E.

resarta Butler, form squalida stat. rev. (all figures not to scale, upper side left, under

side right): (29, 31) males: (29) Mer I., 26.1.2011, TAL & AIK, [forewing length 42

mm]; (31) Murray (Mer) L, 22-25.iv.1989 TAL, [43 mm]; (30, 32) females: (30)

Murray I., 29.iii-4.iv.1986, TAL, [41 mm]; (32) Mer I., 26.1.2011, TAL & AIK, [42

mm]. (33-36) Morph type 1 = Е. b. batesii: (33, 35) males: (33) Dauan I., 8.1.2006,

TAL, [45 mm]; (35) Yam (Iama) I., 7.iv.1987, MDB, [43 mm]; (34, 36) females: (34)

Dauan I., 20.11.2004, TAL, [40 mm]; (36) Yam I., 11-12.vi.1992, AIK, [43 mm].

Australian Entomologist, 2013, 40 (4) 201

Table 1. Specimens of the E. batesii ‘complex’ collected from Torres Strait,

Queensland and mainland Papua New Guinea (including Long and Normanby

Islands), grouped into one of three morphological categories (Morph types 1-3),

relative to collection locations (Papua New Guinea collection locations listed roughly

west to east in descending order).

Specimens in each morphological category

Collection location data :

Morph type 1 Morph type 2 Morph type 3

* Australia" 15, 19* (MM) - -

Cape York, Qld 19 (AM) > 14 (AM)**

Erub (Darnley) I., TS 19 (AM) - -

Mer (Murray) I., TS 18 (AM) - -

Mer (Murray) I., TS 4143, 1999 3543, 2099 1463, 899

Dauan I., TS 933,429 = €

Iama (Yam) I., TS 18,19 e :

Balamuh, WP 14 = т

Orokolo, GP W&L - -

Aitape, SP 19 - =

Angoram, ESP 18 2 =

Long I., MP - 268 -

Langemak, MP 299 - -

Lae, MP 399 1366,19 5

Bulolo, MP 13,19 733, 699 233

Wau, MP - 15 -

Sambio, MP - 359,399 233

Port Moresby, CP 343,429 633 -

Brown R, Port Moresby 433,19 284,299 288,299

Кара Кара, Port Moresby 16,19 18 -

Ѕорегі, Port Moresby 1d ^ -

Laloki R., nr Port Moresby 1d = -

Subitana, CP 288,229 n =

Rigo, СР 14 :

Mt Lamington, OP - 14 e

Erora Ck, nr Oro Bay, OP - 20d e

Normanby I., MBP - 333 -

Papua New Guinea provinces are: CP = Central; ESP = East Sepik; GP = Gulf; MBP = Milne

Bay; MP = Morobe; OP = Oro; SP = Sandaun; WP = Western.

AM = Australian Museum, Sydney; MM = Macleay Museum, Sydney; Qld = Queensland; TS =

Torres Strait; W&L = citation from Waterhouse and Lyell (1914).

*Lectotype of E. b. belia Waterhouse & Lyell; "illustrated in Waterhouse & Lyell (1914).

202 Australian Entomologist, 2013, 40 (4)

including an intermediate morph with ‘obsolescent white spots’ [Morph type

3 (closest to E. b. squalida-like: Figs 29-32) (22 of 138; 15.9% of specimens

examined)]. In addition to Mer I., specimens of the E. batesii ‘complex’ are

known in Torres Strait in the east from one historical specimen collected

from Erub Island (Morph type 1) (Table 1); elsewhere in Torres Strait from

Dauan Island (Fig. 62) in the north-west (13 specimens known, all of which

are Morph type 1) (Figs 33-34); and just two specimens known further south

in the central western part of the Strait from Iama Island (Fig. 63) (also

Morph type 1) (Figs 35-36). Moreover, one of the two historical specimens

purportedly from Cape York is Morph type 1, while the other is Morph type

3. This latter specimen, a male, is believed to be the Lectotype of E. b. belia

(Peters 1971) and was illustrated in Waterhouse and Lyell (1914: fig. 10)

(Table 1).

The review of specimens from mainland Papua New Guinea (including

specimens from Long [MP] and Normanby [MBP] Islands: Fig. 37; Table 1),

together with the Orokolo locality reference from Waterhouse and Lyell

(1914) (92 records in total), plus the eastern Papua New Guinea locality

references in Carpenter (1953) and Ackery and Vane-Wright (1984), showed

that populations from the east of mainland Papua New Guinea (85 specimens

examined) exhibited a high degree of polymorphism, similar to that seen in

the population from Mer I.. Specifically, this eastern zone of Papua New

Guinea, containing these mixed populations, forms a broad, almost

longitudinal area running from the Astrolabe Bay and Long Island region

(Madang Province) in the north to Oro Bay (Oro Province) and Rigo (Central

Province) in the south, including the D'Entrecasteaux Islands (Goodenough,

Fergusson and Normanby Islands: Milne Bay Province), which then stretches

south-east along the Louisiade Archipelago in the south of the Solomon Sea

(Fig. 37). Morph type 2 is known to the east of the Louisiade Archipelago

solely on Rennell I., the southernmost island of the Solomon Archipelago

(Fig. 37).

Specimens from eastern mainland Papua New Guinea therefore included all

three morph types, ie. Morph type 1 (30 of 85: 35.3% of specimens

examined), Morph type 2 (47 of 85: 55.396 of specimens examined) and

Morph type 3 (8 of 85: 9.496 of specimens examined), similar to the

proportions of each type observed on Mer I. Interestingly, all Morph type 3

specimens examined from Australia and mainland Papua New Guinea, except

the Lectotype of E. b. belia [allegedly collected on Cape York], were either

from Mer I. or the zone of polymorphism in the far east and south-east of

mainland Papua New Guinea (Fig. 37). The small number of specimens (4)

recorded from the western and southern parts of Papua New Guinea (Aitape,

Angoram, Balamuh and Orokolo: Table 1) (Fig. 37) were all Morph type 1,

i.e. immaculate, brown to black in colour and devoid of white spotting on the

uppersides of the wings.

Australian Entomologist, 2013, 40 (4) 203

Morph types

Pr N га

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| PAPUA NEW ў

GUINEA NEW Ser

bitza Rex

Balamuh i SK a NC ОРКЕ det 2 OP

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Fig. 37. Map of Papua New Guinea, Solomon Islands and northern Queensland,

Australia, showing the known distributions of Е. b. batesii (Morph type 1), E. resarta

stat. rev. (Morph type 2) and E. resarta form squalida stat. rev. (Morph type 3).

Morphological characters of populations in the area bounded in eastern mainland

Papua New Guinea (except Goodenough and Fergusson Islands [Ackery and Vane-

Wright 1984]) and Torres Strait were derived from museum specimens.

Morphological characters of populations from western mainland Papua New Guinea

were based on museum specimens plus Fruhstorfer (1910), Barrett and Burns (1951),

Carpenter (1953), Ackery and Vane-Wright (1984) and Parsons (1998). Other

arrowed island locations were based on Carpenter (1953), Ackery and Vane-Wright

(1984), Parsons (1998) and Tennent (2001, 2002a).

Examination of the genital armature, in particular the phallus (Figs 38-41),

saccus (Figs 42-46) and left valva (Figs 47-54) of each of the eight voucher

specimens (4 x Morph type 1 and 4 x Morph type 2), showed that these

structures were variable and, therefore, could not be used to discern morph

types within the eight specimens.

The phalli (Figs 38-41) (i.e. two Morph type 1 and two Morph type 2) were

overall similar in shape but the degree of variation was dependant on how far

the cornuti were extended (Figs 38-41). The shapes of the tips of the sacci in

particular were variable including their length, but the variation in length

might be relative to the size of the butterfly (Figs 42-46). In addition, overall

shapes of the sacci were comparable only when viewed at the same angle and

this was difficult to standardise when viewed microscopically. The valvae

examined were mostly similar in overall shape (Figs 47-54) but their apical

tips (claspers) were variable in their acuteness, with some more rounded

(Figs 47, 50, 53) than others.

204 Australian Entomologist, 2013, 40 (4)

Figs 38-46. Euploea batesii ‘complex’, male genital armature: (38-41) ventral view of

‘phallus to scale, all phalli approximately 4.5 mm long: (38, 40) Morph type 1 = E. b.

batesii: (38) Brown R., nr Port Moresby, [Central Province); (40) Yapen, Irian Bay,

1995; (39, 41) Morph type 2 — E. resarta stat. rev.: (39) Sambio Mumeng, [Morobe

Province], -.i.1985; (41) Bulolo, [Morobe Province], -.v.1985. (42) ventral view of

genitalia including saccus, total length approximately 5 mm, Morph type 1 = Е. b.

batesii, Brown R. nr Port Moresby. (43-46) dorsal view of saccus to scale: (43, 45)

Morph type 1 = E. b. batesii: (43) Brown R., nr Port Moresby; (45) Yapen, Irian Bay,

1995; (44, 46) Morph type 2 = Е. resarta stat. rev.: (44) Sambio Mumeng, [Morobe

Province], -.1.1985; (46) Bulolo, -.v.1985.

Australian Entomologist, 2013, 40 (4) 205

Figs 47-54. Euploea batesii ‘complex’, male genital armature (lateral left view of left

valva [clasper] with tegumen and uncus removed [dorsal surface uppermost]), all

valvae approximately 2 mm high: (47, 49, 51, 53) Morph type 1 = E. b. batesii: (47)

Dauan L, Torres Strait, 2.1.2010; (49) Dauan I., 8.1.2010; (51) Brown R., nr Port

Moresby [Central Province]; (53) Yapen, Irian Bay, 1995; (48, 50, 52, 54) Morph

type 2 = Е. resarta stat. rev.: (48) Murray (Mer) L, Torres Strait, 29.iii.-4.iv.1986;

(50) Murray I., 29.iii.-4.iv.1986; (52) Sambio Mumeng [Morobe Province], -.i.1985;

(54) Bulolo [Morobe Province], -.v.1985.

206 Australian Entomologist, 2013, 40 (4)

MAA

У | P

Figs 55-60. Euploea batesii ‘complex’, proportions of antennal length fulvous in

colour (figures not to scale): (55, 56) fulvous colour <10% of antennal length: (55)

Murray (Mer) I., 9.11.1995, 4, Morph type 2 = E resarta stat. rev.; (56) Mer I., em.

11.11.2011, $, Morph type 1 = E. b. batesii. (57, 58) fulvous colour 15-25% of

antennal length: (57) Murray I., 29.iii.-4.iv.1986, 9, Morph type 3 = £. resarta form

squalida stat. rev.; (58) Mer I., em. 6.111.2011, 9, Morph type 1 = Е. b. batesii. (59,

60) fulvous colour circa 40% of antennal length: (59) Dauan I., 16.1.2011, 9, Morph

type 1 = Е. b. batesii, (60) Murray I., 22-25.iv.1989, 4, Morph type 1 = Е. b. batesii.

In addition, examination of specimens showed that the distal flagellomeres of |

the antennae, in both older and relatively fresh specimens and inclusive of the

three morph types, were fulvous in colour (Figs 55-60), not black as in all but

one other Euploea spp from Torres Strait. The extent of this fulvous

colouring varied from being just the antennal tips (Figs 55-56) to the

approximate terminal 35-4596 of the antennal lengths (Figs 59-60).

Discussion

Precis of the biogeography of Torres Strait and eastern Papua New Guinea

The geological histories of the Torres Strait islands, Papua New Guinea and

the Solomon archipelago are complex (Parsons 1998). Willmott (1972)

indicated that northern Queensland (ie. Cape York), the southern Torres

Strait islands (e.g. Muralug [Prince of Wales] and Moa Is) and the top

western Torres Strait islands to the north, i.e. Тата, Gabba and Dauan, have a

similar geology, being composed of Carboniferous granite and acid volcanic

rocks. This same granite rock ‘basement’ appears again as a granite boulder

Australian Entomologist, 2013, 40 (4) 207

cluster at Mabaduan on the southern coastline of Papua New Guinea just

north-east of Dauan I., then reappears further north several kilometres inland

as the Oriomo Plateau, a low plateau mostly less than 50 m above sea level.

Since around 10,000 years ago, the intervening areas between Cape York and

the Oriomo Plateau have been inundated by Torres Strait and, in southern

Papua New Guinea, covered by an alluvial terrace which is low and swampy

(Willmott 1972). Prior to this, a prolonged Quaternary connection existed

between New Guinea and Australia (Cranston and Naumann 1991).

Therefore, based on the evidence of Willmott (1972) that Papua New Guinea

and Cape York were once joined along this granite ‘basement’, it is not

surprising that the butterfly fauna of the Torres Strait islands contains a

mixture of Australian and Papua New Guinea faunal elements (Kitching and

Dunn 1999, Braby 2000, unpublished data).

In contrast, the geology of Mer and its two associated islands Dauar and

Waier (constituting the Murray Island group), in eastern Torres Strait, are

thought to be the remnants of old volcanic activity of the Upper

Carboniferous Period (Willmott 1972). While Mer I. is presently well

isolated from the western Torres Strait islands and Papua New Guinea,

components of the butterfly fauna recorded from the island indicate a link to

eastern mainland Papua New Guinea, away from the granite based Oriomo

Plateau (which lies directly north of Dauan Island).

Of the Mer I. butterfly fauna, several New Guinea taxa that occur on the

island have not been recorded elsewhere in Torres Strait, including Mer’s

neighbouring volcanic islands of Erub (Darnley) and Ugar (Stephens). In

addition to Morph types 2 and 3 of the E. batesii ‘complex’, which are only

known to occur in Torres Strait on Mer I., several other butterfly taxa also

appear to occur in Torres Strait only on Mer I. The occurrence of these taxa

suggests that the island's butterfly fauna might be derived from eastern Papua

New Guinea.

These other butterfly taxa are: Nothodanis schaeffera caesius (Grose-Smith,

1894) (Lycaenidae) (Lambkin and Knight 1990, Meyer et al. 2005), which

occurs sporadically throughout mainland Papua New Guinea (Parsons 1998);

Hypolimnas antilope mela Fruhstorfer, 1903 (Nymphalidae) (with one female

specimen also recorded from Masig (Yorke) I., Torres Strait: Wood 19872, b,

Lambkin and Knight 1990) is rarely encountered in Papua New Guinea

(Parsons 1998); Melanitis constantia (Cramer, 1777) (Nymphalidae)

(Johnson et al. 1994), which is again widespread throughout mainland Papua

New Guinea but rarely observed (Parsons 1998); and, in particular, Euploea

modesta lugens Butler, 1876, (a single male recorded by Meyer et al. (2004)

on Mer I.), whose distribution is otherwise restricted to the hinterlands of

Port Moresby, Central Province (Parsons 1998, Meyer et al. 2004).

Curiously, Tellervo zoilus digulica Hulstaert, 1924 (Nymphalidae), another

butterfly taxon known only in Torres Strait from Mer I. (Johnson et al. 1994,

208 Australian Entomologist, 2013, 40 (4)

Braby 2000) but, enigmatically, outside Torres Strait appears restricted to the

Bensbach and Morehead River regions in the Western Province of Papua

New Guinea (Parsons 1998), both locations not far from Dauan I.

With regard to the E. batesii ‘complex’, it extends east of the New Guinea

mainland throughout the Solomon archipelago, where observed trends with

its distribution and polymorphism are difficult to interpret, although different

island populations are phenotypically constant (M. De Baar pers comm.).

Similarly Carpenter (1953) found this in his review of Euploea in

Micronesia, Melanesia, Polynesia and Australia, where he found conundrums

when attempting to elucidate the biogeography of the Solomons.

Interestingly, in the east of the Solomons, E. b. ackeryi Tennent, 2001, which

occurs exclusively on Ulawa Island (Fig. 37), resembles nominotypical Ё.

batesii, i.e. Morph type 1 from western New Guinea (Tennent 2001). In

addition, E. b. kunggana Carpenter, 1953 from Rennell Island (Fig. 37),

which is the most southerly island in the Solomon archipelago, is noticeably

different from all other populations occurring northward through the

archipelago but, in general, most resembles the well-spotted E. b. resarta

(Morph type 2) from eastern mainland Papua New Guinea (including the

Louisiade and D'Entrecasteaux island groups) (Fig. 37). When discussing

this, Carpenter (1953) postulated that the well-spotted butterflies generally

occurring on Rennell I., including Е. b. kunggana, show a closer affinity with

the butterflies of the eastern tip of mainland Papua New Guinea (i.e. Central,

Oro and Milne Bay Provinces) (Fig. 37) and may have originated from this

region, rather than via the main Solomon archipelago north of Rennell I.

Alternatively, Ackery and Vane-Wright (1984) and Tennent (2002) suggested

that the well-spotted butterfly species occurring on Rennell Island, including

E. b. kunggana, might be part of a Müllerian mimicry complex that might

play a role in the phenotypic appearance of danaines on the island.

Additionally, Ackery and Vane-Wright (1984) reported on the danaid

populations occurring on the islands in the Louisiade Archipelago, south-east

of Milne Bay, and listed E. b. rotunda (Morph type 1) on the Conflict Is

group and Е. b. resarta (Morph type 2) on Nivani, Misima and Tagula Is

(Fig. 37). This archipelago stretches partly across the southern extremity of

the Solomon Sea almost directly in line with Rennell I. (Fig. 37). In a similar

vein, Tennent (2001) commented that Е. b. ackeryi resembled dark, unspotted

E. b. batesii (Morph type 1) from New Guinea and that it was quite different

from all the races of E. batesii proximal to it along the Solomon archipelago.

Therefore, in Torres Strait, a biogeographical pattern exists with island

populations of the butterfly on the western islands, Dauan and Iama, being

unspotted, as with populations occurring across most of New Guinea. This

pattern supports the possible link in the geology of Dauan (Fig. 62) and Iama

(Fig. 63) Islands with those of the Oriomo Plateau directly north in the

Australian Entomologist, 2013, 40 (4) 209

Western Province of southern Papua New Guinea. Similarly, Mer I. (Fig. 61)

in the east has a mix of morph types similar to populations from the east of

mainland Papua New Guinea. Occurrence of these morph types on Mer I.,

including those of other butterfly species recorded only in Torres Strait on

Mer L, also indicates a possible geological link between Mer I. and the

provinces to the east and north of Port Moresby in Papua New Guinea.

Further studies of the butterfly fauna of the northwestern Torres Strait

islands, including Gabba I. (which lies between Dauan and Iama Is) (Fig. 64),

and the fauna of the southern and eastern provinces of Papua New Guinea,

may offer further empirical evidence to support this link.

Figs. 61-64. Torres Strait islands: (61) Mer (northeastern aspect); (62) Duaun

(southeastern aspect); (63) Iama (southern aspect); (64) Gabba (eastern aspect).

Life history and biology

The life history and host plant of Morph type 1 of the E. batesii ‘complex’ in

Torres Strait were previously unknown. The larval host plant, Parsonsia

velutina, is widely distributed throughout Torres Strait and southern and

eastern Papua New Guinea (Atlas of Living Australia 2013, G. Sankowsky

unpublished data), so it is possible that this species may be the host plant for

Morph type 1 of the E. batesii ‘complex’ throughout these regions. In Papua

New Guinea, Szent-Ivany and Carver (1967) reported E. b. funerea Butler,

1878 (which resembles Е. b. squalida) from Port Moresby to be a frequently

observed taxon whose larvae were found commonly on Nerium oleander, but

they did not provide a description of the larva. In Torres Strait, immature

210 Australian Entomologist, 2013, 40 (4)

stages of the E. batesii ‘complex’ have not been observed on N. oleandey

despite careful surveillance of plants over several decades (unpublished data),

The only reference to N. oleander as a Euploea larval host plant in Torres

Strait is by Johnson and Valentine (1997), who reared adults of E. alcathog

misenus Miskin, 1890 (as E. a. monilifera [Moore, 1883]) from Pupae

collected on N. oleander on Dauan I.

Parsons (1998) illustrated the fourth and fifth instars of the E bates jj

‘complex’ from material collected on Parsonsia lata from the Bulolo district,

Morobe Province, Papua New Guinea. Although very similar to the two

instars of Morph type 1 described in this work and based on a single record,

there are some subtle differences between the larvae from Bulolo (Parsons

1998) and those from Mer I. Based on the illustration in Parsons (1998) and

the final instar larvae collected and reared on Mer I., it appears that the

lengths of all the filaments of the final instar larvae from Mer I. are

approximately twice the width of the body of the larva (Figs 6-7), Whereas

the filaments of the final instar larva illustrated in Parsons ( 1998) are almost

all less than the width of the body, except the mesothoracic pair which is a

little more than the width of the body. The filaments of the fourth instar larva

illustrated again appear to be shorter overall in the Bulolo specimen (Parsons

1998) than in specimens from Mer I. (Fig 5), while the Bulolo Specimen

illustrated is a much brighter orange (Parsons 1998) than any fourth instar

larva observed from Mer I., which were all dull yellow-orange in colour (Fig

5). Unfortunately, Parsons (1998) did not specify the wing pattern

morphology of the adults that he reared from these larvae and there is only

one illustration (used in Ackery and Vane-Wright 1984 and Parsons 1998)

available for comparison, so it is not known which morph type he reared

from these particular larvae. Considering that Bulolo lies in the area where all

three morph types occur, the subtle differences of the larvae noted between

the two locations, and in view of Parsons (1998) indicating the extreme

variability of E. batesii from the Bulolo area, it is possible that the adult

butterflies reared from these larvae might have been Morph type 2 or 3, ie.

resembling Е. b. resarta or E. b. squalida, with white or ‘obsolescent’ Spots,

but further information is required before any inference can be made. In any

case, the distinctive appearance of final instar larvae from Mer I. and Bulolo,

especially their charcoal black colour devoid of any colour banding, has not

been observed elsewhere in final instar larvae of Euploea spp occurring in

New Guinea and Australia (Saguru and Fukuda 1997, 2000, Parsons 1998,

Braby 2000) and might be characteristic of the E. batesii ‘complex’.

Ackery and Vane-Wright (1984) documented the plant species known to

attract adult danaines, predominantly as a source of pyrrolizidine alkaloids.

Here, I record for the first time a pyrrolizidine alkaloid source for danaines in

Torres Strait, viz. Heliotropium foertherianum, for the E. batesii ‘complex’

(three morph types), E. a. amycus, E. t. tulliolus and T. h. hamata.

Heliotropium foertherianum is known to contain pyrrolizidine alkaloids and

Australian Entomologist, 2013, 40 (4) 211

is recorded in the literature as readily attracting danaines in the Pacific

(Ackery and Vane-Wright 1984, Tennent 2002b, Patrick and Patrick 2012).

Taxonomy of the E. batesii 'complex'

From the early days, the Northern Moluccas remained the type location and

the types appeared to be the only known specimens of nominotypical Е.

batesii (Edwards et al. 2001). As a result of confusion with the types,

Fruhstorfer (1910) did not illustrate nominotypical E. batesii in his treatise

but chose to illustrate E. b. pinaria Fruhstorfer, 1910 from Waigeo as a

representative of the type. Subsequently, a plethora of names, many

originally described as species, was given to populations from mainland New

Guinea and its adjacent islands. Excluding the far east of mainland Papua

New Guinea, the most recently assigned names are: E. b. arcana Talbot & Le

Cerf, 1925 from Seram, Ambon and Buru; ЕЁ. b. pinaria from Waigeo; E. b.

ebinena Butler, 1866 from the Aru islands; E. b. incerta Joicey & Noakes,

1915 from Biak (including Yapen) (Fig. 16); E. b. gorgonia Hulstaert, 1924

and E. b. mimica Fruhstorfer, 1910 from western New Guinea; and E. b.

publilia Fruhstorfer, 1910 from Astrolabe Bay, Madang Province, Papua

New Guinea (Fruhstorfer 1910, Carpenter 1953, Ackery and Vane-Wright

1984, Parsons 1998). Based on their descriptions, the wing patterns of these

populations from most of mainland New Guinea and its close outlying

western islands are, for the most part, similar, if not essentially the same as

the description of the type of E. batesii (Felder and Felder 1865) (i.e. Morph

type 1) (Fruhstorfer 1910, Barrett and Burns 1951, Carpenter 1953, Ackery

and Vane-Wright 1984, Parsons 1998). In contrast, white-spotted and

‘obsolescent’ populations appear to be restricted to the far east of mainland

Papua New Guinea, its close neighbouring islands and Mer I. in Torres Strait.

Fruhstorfer (1910) was the first to reduce the number of species names within

the complex, indicating that E. batesii occurred as an unspotted species

across most of the New Guinea island but at the same time describing several

subspecies, possibly succumbing to the trend of using the trinomial system of

taxonomy that had just gained popularity by the early 20th Century (Braby et

al. 2012). In addition, Fruhstorfer (1910) synonymised many of the names in

the east of Papua New Guinea, retaining E. resarta and E. fünerea as species.

Following this, Waterhouse and Lyell (1914) and Waterhouse (1932) referred

the Torres Strait population to E. b. belia, which was in accord with

Fruhstorfer (1910), while Corbet (1943) referred all populations of the E.

batesii ‘group’ to one single species occurring from the Moluccas to the

Solomons. Carpenter (1953) later grouped the populations into geographical

races, effectively subspecies, as part of a complex. Most recently, Ackery and

Vane-Wright (1984) followed Corbet (1943) and Carpenter (1953) and

maintained their synonymy, placing them in a single species, i.e. E. batesii.

In the current study, examination of the genital armature of Morph types 1

and 2 showed no consistent differences. Similarly, Corbet (1942) examined

212 Australian Entomologist, 2013, 40 (4)

the genitalia of several species-groups of Euploea and concluded that

differences in genital armature were not discernible between closely related

species. Moreover, Talbot (1943) determined that there were no differences

between the genital armature of E. batesii (he did not specify the origins of

the specimen[s] he examined) and E. honesta Butler, 1882 (E. b. honesta)

from the Solomons, despite there being noticeable and consistent differences

in the adult wing pattern.

Interestingly, the noticeably fulvous colouring of the distal flagellomeres of

the antennae on all specimens examined of the E. batesii ‘complex’ from

New Guinea and Torres Strait (Figs 55-60) provides another character to

assist in distinguishing specimens of female E. batesii from females of

several similar-looking species in Torres Strait, viz. E. sylvester (Fabricius,

1793), E. algea (Godart, [ 1819]) and E. alcathoe (Godart, [1819]).

Furthermore, examination of specimens of all species of Euploea from Torres

Strait (in TLIKC) showed that this antennal character was shared only with E.

netscheri nerana Fruhstorfer, 1910 (31 specimens examined from Dauan

055, 2099] and Saibai [19] Islands). Thus, despite the fulvous distal

flagellomeres of the antennae being a helpful character in distinguishing

female specimens of the E. batesii ‘complex’ from most other similar-looking

Euploea species, this character may make it difficult to separate them from

female E. netscheri Snellen, 1889. This particular character may indicate a

closer relationship between E. batesii and E. netscheri than is currently

thought (Ackery and Vane-Wright 1984). The relationship between these two

taxa might be better understood once the life history and, in particular, the

final instar larva of E. n. nerana are known.

Distribution in Papua New Guinea and Torres Strait

In the far east of mainland Papua New Guinea, numerous names have been

applied to populations of the E. batesii *complex' (with six subspecies

described), in which wing pattern morphology seems to be highly variable

even in discrete locations, i.e. specimens range from unspotted black or dark

brown forms (Morph type 1) to well spotted forms (Morph type 2), often with

both forms and intermediate-looking forms (i.e. Morph type 3) occurring at

the same locality (Table 1). Carpenter (1953) looked at the variation within

these names and accordingly treated several taxa (E. b. funerea and E. b.

squalida from Port Moresby, Central Province; E. b. turbonia Fruhstorfer,

1910 from Simbang, Morobe Province; and E. b. murena Fruhstorfer, 1911

from Yule Island, Central Province) as junior synonyms of E. b. resarta, the

Holotype being from Port Moresby (Butler, 1876). Similarly, as discussed

earlier, E. b. publilia from Astrolabe Bay (a predominantly dark brown to

black form) and E. b. rotunda van Eecke, 1915 (which is also dark brown to

black), are known from Mekeo (type locality) and Yule Island (both Central

Province) respectively (Parsons 1998), with Yule I. being the same type

locality as for E. b. murena (Fruhstorfer, 1911). Both Mekeo and Yule I. are

Australian Entomologist, 2013, 40 (4) 213

close to Port Moresby so, according to the correlation of Carpenter (1953), it

is probable that E. b. rotunda should also be treated as a synonym of E. b.

resarta.

In Australia, E. b. belia was described from three males and five females

originating from ‘Cape York’ (16, 19 AM), Erub (Darnley) I. (19 AM), Mer

(Murray) I. (14 AM) and ‘Australia’ (1d, 19 MM) (Peters 1971) (the

depository of the remaining 299 is unknown). The wing pattern of almost all

of these early Australian specimens is dark brown to black without spots

(Morph type 1), strongly reminiscent of E. b. batesii. As reported earlier, the

exception is the male allegedly collected at Cape York and illustrated in

Waterhouse and Lyell (1914), whose wing pattern morphology is consistent

with Morph type 3, i.e. with white spots and bars overlaid with dark ground

colour scales (as per Figs 29-32). Later, it seems that Waterhouse (1932) had

doubts about the authenticity of Cape York as a location for Е. batesii as he

redefined the species as only occurring within Australia on Darnley (Erub)

and Murray (Mer) Islands, Torres Strait. Perhaps the Cape York records are

similar to those of several other butterfly specimens from this early period

that were also purportedly collected from Cape York and Thursday Island,

but were found later to originate, in all likelihood, from New Guinea or

further afield (Waterhouse and Lyell 1914, Meyer et al. 2004, Lambkin

2005). Thus, in all probability, these two Australian locations were points of

export for these consignments of natural history specimens, not the places of

capture.

In Torres Strait, specifically on Mer I., the E. batesii ‘complex’ occurs as the

three morph types (described as subspecies); i.e. Morph type 1 described as

E. b. belia (Figs 13-15, 17-20), Morph type 2 (known as E. b. resarta) (Wood

1987a, Lambkin and Knight 1990) (Figs 21-28) and Morph type 3 (nearest to

E. b. squalida) (Figs 29-32). On other Torres Strait islands (i.e. Erub, Тата

and Dauan) Е. batesii is only known as Morph type 1. Wood (19872) found

all three subspecies (or morph types) on Mer I. and, based on this, he treated

E. b. belia as a junior synonym of E. b. resarta. Thus, from that time, all

three morph types of the E. batesii ‘complex’ were considered, at least on

Mer I., to be E. b. resarta (Braby 2000). Similarly, this study has shown that

Morph type 3 occurs only in areas where Morph types 1 and 2 occur

sympatrically. These areas appear to be exclusively eastern mainland Papua

New Guinea and Mer I. in Torres Strait.

Morph type 1 of the E. batesii ‘complex’ and Parsonsia velutina

On searching for immature stages of the £. batesii ‘complex’ on Mer I., P.

velutina was the only species of Parsonsia found (and is the only species

recorded: Atlas of Living Australia 2013), and only on this species were

found immature stages, which solely produced adults of Morph type 1.

However, it is possible that either an unrecorded species of Parsonsia or

another species in the Apocynaceae (with four others recorded from the

214 Australian Entomologist, 2013, 40 (4)

island: Atlas of Living Australia 2013) may constitute the larval host of the

maculated forms of the E. batesii ‘complex’ (ie. Morph types 2 and a)

which occur sympatrically with Morph type 1 on the island.

Conclusion

The geographical distribution and wing pattern morphological stasis of

Morph type 1, which occurs in a continuum across New Guinea with ап

abrupt and exclusive modification in the far east of the island where Morph

types 2 and 3 occur with it, is problematic in its interpretation, especially in

terms of differentiating the morph types as subspecies (as per Fruhstorfe,

(1910) and Carpenter (1953)). This is especially the case when attempting to

apply the most fundamental premise of the subspecies model to this unique

distribution, the premise in question being based on allopatric populations

requiring geographic isolation to be regarded as subspecies. In a recent

review of the utility of subspecies as a taxonomic unit, Braby ef al. (2012

concluded that defining species was problematic because of attempts to apply

the species concept in nature. Their review concluded that, when applying the

species concept, multiple lines of evidence are preferable.

In view of the data presented here on the Е. batesii ‘complex’ in the New

Guinea region, this study provides evidence indicating that ‘Æ. batesii’ does

not constitute a single species on the island of New Guinea. The data further

suggest that the E. batesii ‘complex’ occurring through the major islands anq

archipelagos east of New Guinea, in particular the Solomon Islands, also

constitutes more than one species. These lines of evidence are:

Specimens of Morph type 1 (taxon batesii) and Morph type 2 (taxon resarta) can

clearly be differentiated based on wing pattern morphology, i.e. specimens of

batesii are black or dark brown without white spotting (immaculate), while

specimens of resarta are also black or dark brown but have varying degrees of

bright white spots or bars (maculation), these spots sometimes overlayed and

obscured with dark brown ground colour scales (taxon squalida);

The stasis of the dark immaculate taxon (Morph type 1) that occurs across New

Guinea and several islands in Torres Strait;

The high frequency of polymorphism that includes the well-spotted taxon Morph

type 2 (taxon resarta) and Morph type 3 (taxon squalida), which occur uniquely

together with the dark immaculate form (Morph type 1) only in the far east of

mainland Papua New Guinea and on Mer I. in the eastern sector of Torres Strait;

Life history data showing that Morph 1 types only arise from Morph 1 type

mothers;

Morphologically consistent larval characters of material reared from Mer I., which

includes the unique colouring of the final instar larva and larval filament length,

being consistent with Morph 1 type adults; and

Apparently longer filament lengths of larvae, in particular the final instar from

Mer I., than filaments of the same from Bulolo in eastern Papua New Guinea.

Australian Entomologist, 2013, 40 (4) 215

Based on the evidence presented in this revision, it is proposed that E. resarta

stat. rev. be revised to species status. In addition, the results of the study

indicate that E. batesii and E. resarta are closely related in terms of their

genital armature, the elongate stripes in and above space 1b on the underside

of the forewings and the fulvous colour of the distal flagellomeres of the

antennae. Moreover, the distributional data show that these two species form

a sympatric zone in eastern mainland Papua New Guinea (and perhaps also in

the D'Entrecasteaux and Louisiade Archipelagos) and on Mer I. in Torres

Strait. Whereas E. b. batesii occurs across the island of New Guinea and

throughout Torres Strait as a distinctive stand-alone species, E. resarta

(including f. squalida) in contrast appears to exist only in the east, where it

occurs sympatrically with E. b. batesii. These unique distributions might be

determined by larval host plant preferences.

Finally, based on evidence provided here, populations of the Е. batesii

*complex' (Morph type 1) occurring on mainland New Guinea and its

outlying islands and in Torres Strait (subspecies E. b. arcana, E. b. ebinena,

E. b. gorgonia, E. b. incerta, E. b. mimica, E. b. pinaria, E. b. publilia, E. b.

rotunda and E. b. belia) are placed as new synonyms of Е. b. batesii. Until

further research is undertaken on the distribution, life histories and molecular

character of the E. batesii species ‘complex’ in New Guinea, Torres Strait

and beyond, an explanation for the distribution and frequency of the

morphological types observed in mainland Papua New Guinea and Torres

Strait is that of a single monotypic species (£. b. batesii) in New Guinea that

overlaps in distribution with a polytypic species (E. resarta) in eastern Papua

New Guinea and on Mer I. in Torres Strait. Within its distribution, E. resarta

is polymorphic and varies from specimens with a high degree of white

spotting or maculation to relatively dark individuals with vestigial spotting,

or having white spots that are obscured under a layer of dark coloured ground

scales (form squalida stat. rev.).

Acknowledgements

I thank the local community councils of Dauan, Mer and Iama Islands, Torres

Strait for allowing entry into their communities, supplying lodging and

providing much cooperation during the time spent on their islands.

Appreciation is given to С. De Baar (MDBC), 1.5. Bartlett (ODAFFC), C.J.

Burwell (QM), E.D. Edwards (ANIC), T.L. Fenner (TLFC), G. Forbes

(GFC), S. Ginn (AM) and SJ. Johnson (MTQ) for allowing access to

specimens in their care. J.S. Bartlett gave valuable support by formatting and

preparing the map of the Papua New Guinea region. I acknowledge W.

Crowe and J. Walker of the Australian Quarantine and Inspection Service

(AQIS) for providing relevant permits for exporting live insect material out

of Torres Strait. I thank R. Kendall, President of the Butterfly and other

Invertebrates Club, Brisbane for providing use of the club's stereo

microscope and Helicon Focus” image stacking software. I acknowledge С.

216 Australian Entomologist, 2013, 40 (4)

Sankowsky for providing valuable access to his unpublished data including

the identification of Crotalaria retusa; and E.D. Edwards and G. Sankowsky

for confirming the identity of H. foertherianum. Finally, the late Murdoch De

Baar is acknowledged for his expertise and extensive collection of Euploea

spp, which was of extreme value during the preparation of this work. This

work partially fulfills the requirements for a Master of Philosophy degree

undertaken by the author at the University of Queensland, Brisbane.

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Australian Entomologist, 2013, 40 (4): 219-236 219

A REVISED CHECKLIST OF AUSTRALIAN FRUIT FLIES

(DIPTERA: TEPHRITIDAE)

DAVID L. HANCOCK

8/3 McPherson Close, Edge Hill, Cairns, Qld 4870

Abstract

A revised list and classification is provided for the 301 species of Tephritidae recorded from

Australia. Platensina ampla de Meijere is newly recorded and Scedella vaga (Hardy & Drew,

1996) is placed as a new synonym of S. infrequens (Hardy & Drew, 1996).

Introduction

The dipteran family Tephritidae contains more than 300 species in Australia.

The main taxonomic and faunistic works for the family are: Drew (1989) for

tribe Dacini; Permkam and Hancock (1995a, b) for subfamilies Phytalmiinae,

Trypetinae and Dacinae except Dacini; and Hardy and Drew (1996) for

subfamily Tephritinae. Additional species were recorded or described by

Hancock (1995, 1996) and Drew et al. (1999), while Hancock et al. (2000)

provided a host plant listing for the 278 species then recorded.

Further investigation since the review by Hancock et al. (2000) has refined

the classification and added further taxa to the Australian fauna. Tribe

Callistomyiini was proposed by Hancock (2007b), tribe Phascini recorded by

Hancock (2011) and subfamily Tachiniscinae recorded by Korneyev (2012).

The presence of tribe Myopitini has been confirmed (Julien et al. 2012).

Ten names have been removed from the Hancock ef ai. (2000) list: one

eradicated (Bactrocera philippinensis Drew & Hancock), one homonym

(Oedaspis serrata Hardy & Drew), three junior synonyms (Bactrocera

neotigrina Drew & Hancock, ‘Campiglossa’ turneri Hardy & Drew and

Dacus concolor Drew: see Drew and Hancock 2000, Hancock 2006, 20092),

one new synonym (‘Campiglossa’ vaga Hardy & Drew: see below) and four

misidentifications (Bactrocera pseudodistincta (Drew), Bactrocera redunca

(Drew), Trupanea amoena (Frauenfeld) and Platensina amplipennis

(Walker): see Huxham and Hancock 2002, Hancock 2001, 2009b, 2012c).

Additions to the Hancock et al. (2000) list result from previous omissions,

reinstated or replacement names, newly described species and newly

recorded species. Six overlooked, reinstated or replacement names were

noted by Drew and Hancock (2000), Hancock (2001), Huxham and Hancock

(2002) and Julien et al. (2012). Eight new species were described by

Hancock and Drew (2003c), Huxham ef al. (2006), Royer and Hancock

(2012) and Korneyev (2012). Sixteen newly recorded species were noted by

Hancock (2001, 2006, 2012c), Huxham and Hancock (2002), Hancock and

Drew (2003b, c), Huxham et al. (2006) and Julien et al. (2012). An additional

new record is noted below. As a result, 290 described species (plus 11

apparently undescribed: Hardy and Drew 1996, Hancock 2012a, Royer and

Hancock 2012) are now known from Australia. These are listed in Table 1.

220

Australian Entomologist, 2013, 40 (4)

Table 1. List of Australian Tephritidae. Distribution codes: W = Western Australia; N

= Northern Territory; S = South Australia; О = Queensland; E = New South Wales

(including ACT); V = Victoria; T = Tasmania. Comments: * = deliberately introduced

for weed biocontrol; D&H = Drew & Hancock; H&D = Hardy & Drew; R&H =

Royer & Hancock; auct. = misidentified names used by various authors.

Recorded species

Subfamily TACHINISCINAE

Tribe Tachiniscini

Aliasutra australica Korneyev

Subfamily PHYTALMIINAE

Tribe Acanthonevrini

Acanthonevra group (Dacopsis complex)

Austronevra australina (Hendel)

Austronevra bimaculata Permkam & Hancock

Austrorioxa acidiomorpha (Hendel)

Copiolepis colpopteris Permkam & Hancock

Dacopsis flava (Edwards)

Dirioxa group of genera

Dirioxa pornia (Walker)

Lumirioxa araucariae (Tryon)

Micronevrina apicalis Permkam & Hancock

Micronevrina breviseta Permkam & Hancock

Micronevrina gloriosa Permkam & Hancock

Micronevrina hyalina Permkam & Hancock

Micronevrina mediivitta Permkam & Hancock

Micronevrina montana Permkam & Hancock

Micronevrina setosa Permkam & Hancock

Themaroides group of genera

(Clusiosoma subgroup)

Clusiosoma (Clusiosoma) laterale (Walker)

C. (C.) macalpinei Permkam & Hancock

Clusiosoma (C.) semifuscum Malloch

Clusiosoma (Paraclusiosoma) papuaense Hardy

Clusiosomina puncticeps Malloch

Paedohexacinia clusiosomopsis Hardy

Paedohexacinia flavithorax Hardy

Rabaulia nigrotibia Hering

Trypanocentra nigrithorax Malloch

(Neothemara subgroup)

Neothemara formosipennis (Walker)

Pseudacanthoneura sexguttata (de Meijere)

Distribn

WSQE

oo oooogoáoo

Comments

— confusa Hardy

= fascifacies auct.

Australian Entomologist, 2013, 40 (4)

(Themaroides subgroup)

Acanthonevroides basalis (Walker)

Acanthonevroides jarvisi (Tryon)

Acanthonevroides mayi Permkam & Hancock

Acanthonevroides nigriventris (Malloch)

A. variegatus Permkam & Hancock

Aridonevra cunnamullae Permkam & Hancock

Taeniorioxa quinaria Permkam & Hancock

Termitorioxa bicalcarata (Hering)

Termitorioxa cobourgensis Hancock

Termitorioxa exleyae Permkam & Hancock

Termitorioxa inconnexa Permkam & Hancock

Termitorioxa laurae Permkam & Hancock

Termitorioxa termitoxena (Bezzi)

Termitorioxa testacea (Hendel)

Tribe Phascini

Epinettyra setosa Permkam & Hancock

Tribe Phytalmiini

Diplochorda australis Permkam & Hancock

Phytalmia mouldsi McAlpine & Schneider

Subfamily TRYPETINAE

Tribe Acidoxanthini

Acidoxantha quinaria Permkam & Hancock

Tribe Adramini

Adrama biseta Malloch

Adrama selecta Walker

Coelotrypes circumscriptus (Hering)

Euphranta athertonia Permkam & Hancock

Euphranta leichhardtiae Permkam & Hancock

Euphranta lemniscata (Enderlein)

Euphranta linocierae Hardy

Euphranta marina Permkam & Hancock

Euphranta mediofusca (Hering)

Euphranta meringae Permkam & Hancock

Euphranta minor Hendel

Euphranta mulgravea Permkam & Hancock

Euphranta numeralis Permkam & Hancock

Euphranta ternaria Permkam & Hancock

Euphranta variabilis (Kertész)

Hardyadrama alyta Permkam & Hancock

Hardyadrama excoecariae Lee

221

— bicolor Macquart

222

Hardyadrama magister (Lee)

Hardyadrama presignis (Hardy)

Ichneumonosoma consors (Walker)

Piestometopon distinctum (Permkam&Hancock)

Piestometopon luteiceps de Meijere

Soita psiloides Walker

Tribe Callistomyiini

Callistomyia horni Hendel

Tribe Rivelliomimini

Ornithoschema oculatum de Meijere

O. queenslandense Permkam & Hancock

Tribe Trypetini

Aciuropsis pusio Hardy

Calosphenisca unicuneata (Hardy)

Hemiristina pleomeles Permkam & Hancock

Philophylla australina (Hardy)

Philophylla erebia (Hering)

Philophylla fossata (Fabricius)

Philophylla humeralis (Hardy)

Philophylla quadrata (Malloch)

Vidalia dualis Permkam & Hancock

Tribe Xarnutini

Xarnuta confusa Malloch

Xarnuta cribralis Hering

Subfamily DACINAE

Tribe Ceratitidini

Ceratitella amyemae Permkam & Hancock

Ceratitella bifasciata Hardy

Ceratitella loranthi (Froggatt)

Ceratitella recondita Permkam & Hancock

Ceratitella unifasciata Hardy

Ceratitis (Ceratitis) capitata (Wiedemann)

Paraceratitella compta Hardy

Paraceratitella eurycephala Hardy

Paraceratitella oblonga Hardy

Tribe Dacini

Bactrocera (Apodacus) cheesmanae (Perkins)

Bactrocera (Apodacus) visenda (Hardy)

Bactrocera (Austrodacus) cucumis (French)

Bactrocera (Bactrocera) abdonigella (Drew)

Bactrocera (B.) abscondita (Drew & Hancock)

Australian Entomologist, 2013, 40 (4)

Torres Strait only

W Introduced

Q Torres Strait only

Australian Entomologist, 2013, 40 (4)

Bactrocera (B.) abundans Drew

Bactrocera (B.) aeroginosa (Drew & Hancock)

Bactrocera (B.) allwoodi (Drew)

Bactrocera (B.) alyxiae (May)

Bactrocera (B.) amplexiseta (May)

Bactrocera (B.) anfracta Drew

Bactrocera (B.) antigone (Drew & Hancock)

Bactrocera (B.) aquilonis (May)

Bactrocera (B.) aurantiaca (Drew & Hancock)

Bactrocera (B.) bancroftii (Tryon)

Bactrocera (B.) barringtoniae (Tryon)

Bactrocera (B.) batemani Drew

Bactrocera (B.) bidentata (May)

Bactrocera (B.) breviaculeus (Hardy)

Bactrocera (B.) brunnea (Perkins & May)

Bactrocera (B.) bryoniae (Tryon)

Bactrocera (B.) cacuminata (Hering)

Bactrocera (B.) curreyi Drew

Bactrocera (B.) daruensis Drew

Bactrocera (B.) decurtans (May)

Bactrocera (B.) diospyri Drew

Bactrocera (B.) endiandrae (Perkins & May)

Bactrocera (B.) erubescentis (Drew & Hancock)

Bactrocera (B.) fagraea (Tryon)

Bactrocera (B.) frauenfeldi (Schiner)

Bactrocera (B.) fuliginus (Drew & Hancock)

Bactrocera (B.) furvilineatta Drew

Bactrocera (B.) halfordiae (Tryon)

Bactrocera (B.) hispidula (May)

Bactrocera (B.) humilis (Drew & Hancock)

Bactrocera (B.) jarvisi (Tryon)

Bactrocera (B.) kraussi (Hardy)

Bactrocera (B.) laticaudus (Hardy)

Bactrocera (B.) lineata (Perkins)

Bactrocera (B.) manskii (Perkins & May)

Bactrocera (B.) mayi (Hardy)

Bactrocera (B.) melas (Perkins & May)

Bactrocera (B.) mendosa (May)

Bactrocera (B.) moluccensis (Perkins)

Bactrocera (B.) murrayi (Perkins)

Bactrocera (B.) musae (Tryon)

223

Torres Strait only

Probable hybrid

Torres Strait only

224

Bactrocera (B.) mutabilis (May)

Bactrocera (B.) neohumeralis (Hardy)

Bactrocera (B.) nigrovittata Drew

Bactrocera (B.) notatagena (May)

Bactrocera (B.) opiliae (Drew & Hardy)

Bactrocera (B.) pallida (Perkins & May)

Bactrocera (B.) papayae Drew & Hancock

B. (B.) parabarringtoniae Drew & Hancock

Bactrocera (B.) parafrauenfeldi Drew

Bactrocera (B.) peninsularis (Drew & Hancock)

Bactrocera (B.) perkinsi (Drew & Hancock)

Bactrocera (B.) phaleriae (May)

Bactrocera (B.) pulchra Tryon

Bactrocera (B.) quadrata (May)

Bactrocera (B.) recurrens (Hering)

Bactrocera (B.) repanda Drew

Bactrocera (B.) resima (Drew)

Bactrocera (B.) robiginosa (May)

Bactrocera (B.) romigae (Drew & Hancock)

Bactrocera (B.) rufescens (May)

Bactrocera (B.) rufofuscula (Drew & Hancock)

Bactrocera (B.) russeola (Drew & Hancock)

Bactrocera (B.) silvicola (May)

Bactrocera (B.) speewahensis Fay & Hancock

Bactrocera (B.) strigata (Perkins)

Bactrocera (B.) tenuifascia (May)

Bactrocera (B.) torresiae Huxham & Hancock

Bactrocera (B.) trifaria (Drew)

Bactrocera (B.) trivialis (Drew)

Bactrocera (B.) tryoni (Froggatt)

Bactrocera (B.) turneri Drew

Bactrocera (B.) umbrosa (Fabricius)

Bactrocera (B.) vulgaris (Drew)

Bactrocera (B.) yorkensis Drew & Hancock

Bactrocera (B.) sp. near barringtoniae

Bactrocera (Bulladacus) flavinotus (May)

Bactrocera (Bulladacus) tigrina (May)

Bactrocera (Diplodacus) signatifera (Tryon)

B. (Gymnodacus) calophylli (Perkins & May)

Bactrocera (Hemizeugodacus) aglaiae (Hardy)

Bactrocera (Hemizeugodacus) aurea (May)

Australian Entomologist, 2013, 40 (4)

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NQEV

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Torres Strait only

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= neotigrina D&H

Australian Entomologist, 2013, 40 (4)

Bactrocera (H.) ektoalangiae Drew & Hancock

Bactrocera (Javadacus) aberrans (Hardy)

Bactrocera (Javadacus) melanothoracica Drew

Bactrocera (Javadacus) unirufa Drew

Bactrocera (Melanodacus) nigra (Tryon)

Bactrocera (Paratridacus) expandens (Walker)

Bactrocera (Queenslandacus) exigua (May)

Bactrocera (Sinodacus) strigifinis (Walker)

Bactrocera (Zeugodacus) chorista (May)

Bactrocera (Zeugodacus) cucurbitae (Coquillett)

Bactrocera (Zeugodacus) fallacis (Drew)

Bactrocera (Zeugodacus) macrovittata Drew

Dacus (Callantra) axanus (Hering)

Dacus (Mellesis) petioliforma (May)

Dacus (Mellesis) pusillus (May)

Dacus (Neodacus) absonifacies (May)

Dacus (Neodacus) aequalis Coquillett

Dacus (Neodacus) bellulus Drew & Hancock

Dacus (Neodacus) coenensis Royer & Hancock

Dacus (Neodacus) hardyi Drew

Dacus (Neodacus) newmani (Perkins)

Dacus (Neodacus) palmerensis Drew

Dacus (Neodacus) salamander Drew & Hancock

Dacus (Neodacus) secamoneae Drew

Dacus (Neodacus) signatifrons (May)

Tribe Gastrozonini

Carpophthorella nigrifascia (Walker)

Subfamily TEPHRITINAE

Tribe Cecidocharini

Procecidochares alani Steyskal

Procecidochares utilis Stone

Tribe Dithrycini (Subtribe Platensinina)

Oedaspis group of genera

Hyalopeza aristae Hancock & Drew

Hyalopeza schneiderae Hardy & Drew

Liepana apiciclara (Hardy & Drew)

Liepana helichrysii Hardy & Drew

Liepana latifrons Hardy & Drew

Liepana lugubris (Macquart)

Oedaspis apicalis Hardy & Drew

Oedaspis australis (Malloch)

ooooooGgooooooono

225

Torres Strait only

— concolor Drew

Introduced*

226

Australian Entomologist, 2013, 40 (4)

Oedaspis austrina Hardy & Drew

Oedaspis continua Hardy & Drew

Oedaspis escheri (Bezzi)

Oedaspis gallicola Hardy & Drew

Oedaspis goodenia Hardy & Drew

Oedaspis hardyi Norrbom

Oedaspis mouldsi Hardy & Drew

Oedaspis olearia Hardy & Drew

Oedaspis perkinsi Hardy & Drew

Oedaspis semihyalina Hardy & Drew

Oedaspis trifasciata (Malloch)

Oedaspis trimaculata Hardy & Drew

Oedaspis whitei Hardy & Drew

Oedaspis sp. A near mouldsi

Oedaspis sp. B near mouldsi

Oedaspis sp. C

Oedaspis sp. D near continua

Oedaspis sp. E near apicalis

Platensina group of genera

Australasinia sexincisa (Malloch)

Collessomyia setiger Hardy & Drew

Platensina ampla de Meijere

Platensina parvipuncta Malloch

Platensina platyptera Hendel

Platensina trimaculata Hardy & Drew

Platensina zodiacalis (Bezzi)

Tribe Myopitini

Urophora solstitialis (Linnaeus)

Urophora stylata (Fabricius)

Urophora terebrans (Loew)

Tribe Schistopterini

Calloptera queenslandica (Hardy & Drew)

Calloptera wedelia (Hardy & Drew)

Rhabdochaeta pulchella de Meijere

Rhochmopterum venustum (de Meijere)

Tribe Tephrellini

Sphaeniscus group of genera

Dicheniotes ternarius (Loew)

Sphaeniscus atilius (Walker)

Tribe Tephritini

Campiglossa group of genera

— serrata H&D

H&D 1996

Newly recorded

— amplipennis auct.

Introduced*

Introduced

Australian Entomologist, 2013, 40 (4)

Dioxyna brachybasis Hardy

Dioxyna hyalina Hardy & Drew

Dioxyna sororcula (Wiedemann)

Mesoclanis magnipalpis (Bezzi)

Mesoclanis polana (Munro)

Peneparoxyna minuta Hardy & Drew

Scedella infrequens (Hardy & Drew)

Scedella orientalis (de Meijere)

Euaresta group of genera

Euaresta aequalis (Loew)

Euaresta bullans (Wiedemann)

Spathulina group of genera

Paraspathulina apicomaculata Hardy & Drew

Paraspathulina eremostigma Hardy & Drew

Paraspathulina trimacula Hancock & Drew

Spathulina acroleuca (Schiner)

Sphenella group of genera

Sphenella ruficeps (Macquart)

Tephritis group of genera

Austrotephritis brunnea (Hardy & Drew)

Austrotephritis bushi (Hardy & Drew)

Austrotephritis campiglossina (Hering)

Austrotephritis distigmata (Hardy & Drew)

Austrotephritis fuscata (Macquart)

Austrotephritis hesperia (Hardy & Drew)

Austrotephritis pelia (Schiner)

Austrotephritis phaeostigma (Hardy & Drew)

Austrotephritis poenia (Walker)

Austrotephritis protrusa (Hardy & Drew)

Austrotephritis pumila (Hardy & Drew)

Austrotephritis quasiprolixa (Hardy & Drew)

Austrotephritis tasmaniae (Hardy & Drew)

Austrotephritis transversa (Hardy & Drew)

Austrotephritis trupanea (Hardy & Drew)

Austrotephritis whitei (Hardy & Drew)

Austrotephritis sp. A near bushi

Austrotephritis sp. B near phaeostigma

Cooronga mcalpinei Hardy & Drew

Paraactinoptera collessi Hardy & Drew

Paraactinoptera danielsi Hancock & Drew

Paraactinoptera prolixa (Hardy & Drew)

227

SV Introduced*

QE Introduced*

QE — vaga H&D

QE Introduced*

QEV Introduced

W — turneri H&D

H&D Tephritis sp.

Hancock 2012a

228 Australian Entomologist, 2013, 40 (4)

E PM ye == er

Parahyalopeza bushi Hardy & Drew V

Parahyalopeza multipunctata Hancock & Drew E Lord Howe I. only

Parahyalopeza pantosticta (Hardy & Drew) QEVT

Quasicooronga connecta Hardy & Drew V

Quasicooronga disconnecta Hardy & Drew E;

Tephritis furcata Hardy & Drew Q

Trupanea bifida Hardy & Drew E

Trupanea glauca (Thomson) WSQEV

Trupanea heronensis Hardy & Drew Q

Trupanea notata Hardy & Drew QE

Trupanea opprimata Hering NQ = amoena auct.

Trupanea prolata Hardy & Drew WSQEV

Trupanea pusilla Hardy & Drew WQ

Trupanea queenslandensis Hardy & Drew Q

Trupanea sp. A near terryi W H&D 1996

Trupanea sp. B near terryi Q H&D 1996

Trupanea sp. C near mutabilis V H&D 1996

Composition of generic groups in subfamily Tephritinae was discussed by

Hancock (2001, 2007c, 20102) and Hancock and Drew (2003c), while that of

generic groups and subgroups in tribe Acanthonevrini was discussed by

Korneyev (1999) and Hancock and Drew (2003a). Three new Tephritinae

genera were proposed by Hancock (2001: Australasinia), Freidberg (2002:

Calloptera) and Hancock and Drew (2003c: Austrotephritis). Subgeneric

nomenclature in Dacus Fabricius was revised by Hancock and Drew (2006).

Additional new distribution records were noted by Gillespie (2003) and

Hancock (2010b, 2012b). Many of the Bactrocera Macquart species recorded

only from Torres Strait, including the pest species B. papayae Drew &

Hancock, B. trivialis (Drew) and B. cucurbitae (Coquillett), are vagrants

from Papua New Guinea and are not permanently established there.

Indian Ocean Territories

Five primarily SE Asian species of Bactrocera are known from the

Australian Territory of Christmas Island in the Indian Ocean (Bellis ef al.

2004): B. (B.) albistrigata (de Meijere), B. (B.) arecae (Hardy & Adachi), B.

(B.) papayae Drew & Hancock, B. (B.) umbrosa (Fabricius) and B.

(Zeugodacus) cucurbitae (Coquillett). The last three also occur as vagrants in

Torres Strait but none occurs on mainland Australia.

Host plants

Table 2 lists host plants recorded since the listing of Hancock et al. (2000),

including those for six subsequently described or recorded species. Records

of Phytalmiinae, Trypetinae and Dacinae are from fruit unless otherwise

Australian Entomologist, 2013, 40 (4) 229

indicated. Thirteen additional host plants for Ceratitis capitata in Western

Australia were reported by Woods et al. (2005) and are not repeated here.

Table 2. Host plants recorded since Hancock et al. (2000). * = introduced; PNG =

Papua New Guinea. Sources: 1 = Hancock and Drew 2003a; 2 = Hancock 2002, in

oozing resin beneath damaged bark; 3 = Hancock and Drew 2003b; 4 = Leblanc et al.

2012; 5 = Julien et al. 2012; 6 = Hancock and Drew 2003c; 7 = Hancock 2001 and

reared specimens in Department of Agriculture, Forestry and Fisheries, Brisbane; 8 =

Hancock and Drew 2003c, collected on but not reared.

Fly species Host plant family / species Comments

Subfamily Phytalmiinae

Clusiosoma semifuscum MORACEAE: Ficus nodosa 1

Clusiosomina puncticeps | MORACEAE: Ficus coronata confirmed: 1

MORACEAE: Ficus fraseri 1

Termitorioxa termitoxena COMBRETACEAE: Terminalia sp. bark 2

Subfamily Trypetinae

Euphranta numeralis MORACEAE: Maclura cochinchinensis 3

Subfamily Dacinae

Bactrocera cheesmanae CLUSIACEAE: Garcinia cf. hollrungii in PNG: 4

Bactrocera lineata APOCYNACEAE:

Tabernaemontana aurantiaca in PNG: 4

SAPINDACEAE: Pometia pinnata in PNG: 4

Bactrocera strigifinis CUCURBITACEAE: Cucurbita flowers in PNG: 4

Subfamily Tephritinae

Procecidochares alani* ^ ASTERACEAE: Ageratina riparia 5

Urophora solstitialis * ASTERACEAE: Carduus nutans 5

Urophora stylata * ASTERACEAE: Cirsium vulgare 5

Urophora terebrans * ASTERACEAE: Onopordum acanthium 5

ASTERACEAE: Onopordum illyricum 5

Rhabdochaeta pulchella ^^ ASTERACEAE: Blumea lacera SE Asia: 6

Scedella orientalis ASTERACEAE: Wedelia trilobata 7

Paraactinoptera danielsi ^ ASTERACEAE: Pluchea baccharoides not reared:8

Introduced species

Eleven species have been introduced into Australia, eight deliberately for the

biological control of weeds and three accidentally (see Table 1). The former

includes three species of Urophora Robineau-Desvoidy: U. solstitialis (L.),

U. stylata (Fabricius) and U. terebrans (Loew), all released between 1992

and 2000 for the control of Carduus, Cirsium and Onopordum thistles

respectively (Julien et al. 2012, Morley 2012). The latter includes the pest

230 Australian Entomologist, 2013, 40 (4)

species Ceratitis capitata (Wiedemann) [Mediterranean fruit fly or Medfly],

introduced into the Perth and Sydney districts during the 1890s. Still present

in Western Australia, this species disappeared from the eastern States by the

1940s, possibly as a result of improved control techniques and competition

from the endemic Bactrocera tryoni (Froggatt). Interestingly, reports of C.

capitata from India almost certainly refer to Australian specimens donated by

Walter Froggatt during his visit in 1908 (Hancock 2007a).

Two further species, Eutreta xanthochaeta Aldrich (Tephritinae: Eutretini)

and Tephritis postica (Loew) (Tephritinae: Tephritini), were released for the

biological control of lantana and Onopordum thistles respectively but failed

to establish. Mesoclanis magnipalpis (Bezzi) also appears not to have

established but releases are ongoing (Adair et а/. 2012). Bactrocera papayae

and B. philippinensis were present in northern mainland Australia during the

mid-late 1990s but were eradicated. I find no evidence of the release of

Urophora quadrifasciata (Meigen) in Australia (or of U. stylata prior to

1993), with records (1 9 of each) from *Narabeen West, NSW’ [no such

place] (White and Korneyev 1989) (‘NT” [error] in Hardy and Foote 1989)

regarded as mislabelled specimens of European origin. For identification of

the relevant Urophora species, see White and Korneyev (1989).

Cecidochares connexa (Macquart) (Tephritinae: Cecidocharini), has been

released in Papua New Guinea and several other countries for the biological

control of Chromolaena odorata (Asteraceae) and is a potential candidate for

controlling this weed in northern Queensland.

Additional notes

The following morphological and distributional notes are based on specimens

in the Department of Agriculture (Northern Australia Quarantine Strategy),

Cairns (NAQS) and the Queensland Museum, Brisbane (QMB). One species

is newly recorded from Australia and one new synonym is proposed.

Subfamily Phytalmiinae

Acanthonevroides nigriventris (Malloch)

QUEENSLAND: 1 9, Bribie Is, 0.85 km NE of NPHQ, 27.018°S 153.122°E, 10 m,

24.ix.-9.x.2010, G..Monteith, Malaise, wallum (QMB).

This is a widespread species known from SE Queensland to Victoria

(Permkam and Hancock 1995b). Collected in a Malaise trap baited with sawn

saplings (G.B. Monteith pers. comm.).

Austronevra australina (Hendel, 1928)

QUEENSLAND: 1 d, 60 Silver Ash Rd, Cow Bay, 16°14'7"5 145°27'36"E,

16.xi.2011, A.D. Rice, J.A. Walker (NAQS); 2 63, 2 99, Minbin, Mahony Rd,

17°27'13"S 145°35'26"E, 30.х.2011, S. Cowan, from underside of Acronychia acidula

leaves (NAQS); 1 9, Polly Creek, Garradunga, 17.456°S 146.02°E, 2.52 m, 13.i.-

4.11.2010, J. Hasenpusch, Malaise (QMB).

Australian Entomologist, 2013, 40 (4) 231

: The above records include the most northerly locality now known for this NE

Queensland rainforest species, previously known from Daintree to Tully

(Permkam and Hancock 1995b).

Austrorioxa acidiomorpha (Hendel)

QUEENSLAND: 1 9, Blackbutt Range, top, 26.8676°S 152.192°E, 28.iii.-10.iv.2010,

Monteith, RF Malaise; 9 99, 25.757°S 152.697°E, Tallegalla Weir Junction, 35 m,

7.xi.2011-5.1.2012, G. Monteith, Malaise, RF [except 1 OF] (QMB).

These are additional locality records from SE Queensland for this eastern

Australian species (Permkam and Hancock 1995b). Collected in Malaise

traps baited with sawn saplings (G.B. Monteith pers. comm.).

Copiolepis colpopteris Permkam & Hancock, 1995 (Fig. 1)

QUEENSLAND: 2 05, Gordon’s Creek, 12.713°S 143.320°E, 15 m, 9.xii.2010,

Escalona & Will, MV light; 1 5, 2 99, East Claudie River, 12.714°S 143.287°E, 15

m, 8-16.xii.2010, Monteith & Escalona, MV light, rainforest (QMB).

The above females (Fig. 1) are the first recorded for this species. They are

similar to males but the wing is normal in shape and not modified posteriorly,

the stigma is a little shorter and veins Коз and R4+5 are less strongly curved

anteriorly. The pattern is dark brown with paler costal cells and stigma, a

subhyaline to pale brown semicircular marginal spot filling apex of cell ryis, a

broad, subhyaline marginal indentation in cell m and a subhyaline to hyaline

posterior margin covering cell bm, anal lobe and posterior two-thirds of cell

сш. The black medial vitta on the abdomen is thin and indistinct. The

oviscape is black and about as long as terga IV-VI. The species is known

only from Iron Range, Cape York Peninsula.

Figs 1-2. Two species of Phytalmiinae and Dacinae known only from Iron Range: (1)

Copiolepis colpopteris, female; (2) Paraceratitella oblonga, male. Photographs by

Federica Turco (Queensland Museum, Brisbane).

232 Australian Entomologist, 2013, 40 (4)

Lumirioxa araucariae (Tryon, 1927)

QUEENSLAND: 1 d, Yarraman, 2.9 km SSE, 26.865°S 152.995°E, 12-28.iii.2010,

Monteith, RF Malaise; 1 d, Blackbutt Range, top, 26.8676°S 152.192°E, 12-

28.11.2010, Monteith, RF Malaise (QMB).

The above specimens add two more localities to those recorded from SE

Queensland by Permkam and Hancock (1995b). In the Blackbutt Range male

the fore femora and tibiae are extensively blackened. Collected in Malaise

traps baited with sawn saplings (G.B. Monteith pers. comm. ).

Micronevrina hyalina Permkam & Hancock, 1995

QUEENSLAND: 1 9, Mt Mee, 27.077°$ 152.686°E, 530 m, 28.viii.-10.ix.2010, G.

Monteith, Malaise, wet sclerophyll (QMB).

This is the third (and most northern) known locality for this species in SE

Queensland, recorded previously from Mt Glorious and near Caloundra.

Collected in a Malaise trap baited with sawn saplings (G.B. Monteith pers.

comm. ).

Micronevrina montana Permkam & Hancock, 1995

QUEENSLAND: 1 9, Yarraman, 2.2 km SE, 26.857°S 152.994*E, 12-28.111.2010,

Monteith, RF Malaise; 2 9 9. same data except 2.9 km SSE, 26.865°S 152.995°E, 12-

28.11.2010 or 28.11.-10.1у.2010; 7 84, 1 Ф, 25.7578 152.697°E, Tallegalla Weir

Junction, 35 m, 7.xi.201 1-5.1.2012, G. Monteith, Malaise, RF (QMB).

One of the above specimens differs from those recorded by Permkam and

Hancock (1995b) in having tergite III fulvous medially and all have terga V-

VI wholly black but this species is known to be variable. The outer hyaline

streak in cell r4, is small, very faint or absent. All three Yarraman specimens

have a pair of thin, secondary (middle) scutellar setae present. Collected in

Malaise traps baited with sawn saplings (G.B. Monteith pers. comm.).

Subfamily Trypetinae

Adrama selecta Walker, 1859

QUEENSLAND (Torres Strait): 1 $, Lala Hill, Moa Island, 10.10.2°S 142.18.1°E, 80

m, 1-2.iv.2008, K. Aland, dung/mushroom pitfall; 1 Ф, Prince of Wales Island, camp,

10.42.3°S 142.13.5°E, 10 m, 3-7.1.2008, С. Monteith & К. Aland (QMB).

The above records are the first from Torres Strait for this northern species.

Euphranta leichhardtiae Permkam & Hancock, 1995

QUEENSLAND: 1 д, East Claudie River, 12.714°S 143.287°E, 15 m, 8-16.xii.2010,

Monteith & Escalona, MV light, rainforest (QMB).

The above male has the abdomen mostly fulvous with terga IV-V darker,

tending brownish, similar to that of females and not as black as in other males

recorded by Permkam and Hancock (1995b).

Australian Entomologist, 2013, 40 (4) 233

Subfamily Dacinae

Paraceratitella oblonga Hardy, 1967 (Fig. 2)

QUEENSLAND: 4 85, East Claudie River, 12.714°S 143.287°E, 15 m, 8-

16.xii.2010, Monteith & Escalona, MV light, rainforest (QMB).

The above males (Fig. 2) are the first recorded for this species, known only

from Iron Range, Cape York Peninsula. They are similar to females but the

grey-pollinose area on the scutum is more extensive.

Subfamily Tephritinae

Platensina ampla de Meijere, 1914

QUEENSLAND: 1 9, Ina Ct, Rocky Point, Weipa, 12°37'44"S 141°52'40"E,

3.vi.2013, net, A.D. Rice (NAQS).

This distinctive species is now known from Indonesia, Papua New Guinea,

Solomon Islands and northern Australia (new country record). For diagnosis

and illustrations see Hancock (2012c).

Scedella infrequens (Hardy & Drew, 1996) (Fig. 3)

QUEENSLAND: 1 ĝ, Gatton, 6.xi.1933, F.A. Perkins; 1 9, Stanthorpe, 19.x.1925

(QMB).

Previously known only from males, the female (Fig. 3) oviscape is red-brown

and about as long as terga IV-VI. Variation in the wing pattern in this species

is such that Campiglossa vaga Hardy & Drew, 1996, new synonym cannot

be separated and is here considered to be conspecific with Paroxyna

infrequens Hardy & Drew, 1996 (placed in Scedella Munro by Hancock

2001), the latter name given nomenclatural priority under the Principle of

First Reviser (ICZN 1999: Article 24.2).

Figs 3-4. Tephritinae: (3) Scedella infrequens, female; (4) Austrotephritis sp. nr

phaeostigma, male. Photographs by Federica Turco (Queensland Museum, Brisbane).

234 Australian Entomologist, 2013, 40 (4)

Paraspathulina trimacula Hancock & Drew, 2003

QUEENSLAND: 1 d, 1 $, Caloundra, 17.viii.1934, F.A. Perkins; 1 д, Brisbane,

28.x.1964, H.A. Rose; 1 9, Inglewood, 1.ix.1925 (QMB).

Previously known from a single male from Mt Cootha, Brisbane (Hancock

and Drew 2003c), the female oviscape is black. In one specimen, the outer

marginal hyaline spot in cell cu, is absent.

Spathulina acroleuca (Schiner, 1868)

NEW SOUTH WALES: 1 9, Lord Howe Is., Macleay (QMB).

This widespread species is newly recorded fom Lord Howe Island.

Austrotephritis protrusa (Hardy & Drew, 1996)

QUEENSLAND: 1 6, 20.101°S 147.757°E, Mt Abbot camp, 800 m, 22-28.ix.2011,

Monteith, MV light, 34971 (QMB).

There are few records of this distinctive eastern species known from Mt

Finnigan near Cooktown to northern New South Wales.

Austrotephritis sp. nr phaeostigma (Hardy & Drew, 1996) (Fig. 4)

QUEENSLAND: 2 55, Eidsvold, 17.viii.1923, Bancroft (QMB).

The above specimens appear to represent an undescribed species close to A.

phaeostigma (Hardy & Drew) (Hancock 2012a). Hyaline subapical spots in

cell dm are small and a single spot in cell tza forms a triangle with the pair in

cell гу. A second undescribed species, near A. bushi (Hardy & Drew), was

reported from Victoria by Hardy and Drew (1996, as ‘Tephritis sp. A’).

Acknowledgements

I thank Anthony Rice and Sally Cowan (NAQS) and Geoff Monteith, Susan

Wright and Federica Turco (QMB) for the loan of specimens or photography.

References

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BELLIS, G.A., DONALDSON, J.F., CARVER, M., HANCOCK, D.L. and FLETCHER, M.J.

2004. Records of insect pests on Christmas Island and the Cocos (Keeling) Islands, Indian

Ocean. Australian Entomologist 31(3): 93-102.

DREW, R.A.I. 1989. The tropical fruit flies (Diptera: Tephritidae: Dacinae) of the Australasian

and Oceanian Regions. Memoirs of the Queensland Museum 26: 1-521.

DREW, R.A.I. and HANCOCK, D.L. 2000. Synonymy, geographic distributions, lectotype

designations and type depositories of some Australian and South Pacific Dacinae (Diptera:

Tephritidae). Australian Entomologist 27(1): 27-30.

DREW, R.A.I., HANCOCK, D.L. and ROMIG, М.С. 1999. New species and records of fruit

flies (Diptera: Tephritidae: Dacinae) from north Queensland. Australian Entomologist 26: 1-12.

FREIDBERG, A. 2002. Systematics of Schistopterini (Diptera: Tephritidae: Tephritinae), with

descriptions of new genera and species. Systematic Entomology 27: 1-29.

Australian Entomologist, 2013, 40 (4) 235

GILLESPIE, P. 2003. Observations on fruit flies (Diptera: Tephritidae) in New South Wales.

General and Applied Entomology 32: 41-47.

HANCOCK, D.L. 1995. Philophylla humeralis (Hendel) (Diptera: Tephritidae: Trypetinae)

newly recorded from Australia. Australian Entomologist 22(4): 113-114.

HANCOCK, D.L. 1996. A new species and new combination in Australian Trypetinae (Diptera:

Tephritidae). Australian Entomologist 23(3): 87-90.

HANCOCK, D.L. 2001. Systematic notes on the genera of Australian and some non-Australian

Tephritinae (Diptera: Tephritidae). Australian Entomologist 28(4): 111-116.

HANCOCK, D.L. 2002. A note on the biology of Termitorioxa termitoxena (Bezzi) (Diptera:

Tephritidae). Australian Entomologist 29(3): 96.

HANCOCK, D.L. 2006. The taxonomic placement of Campiglossa vaga Hardy & Drew and

Mesoclanis campiglossina Hering (Diptera: Tephritidae: Tephritinae). Australian Entomologist

33(3): 142.

HANCOCK, D.L. 20072. Book review. Australian Entomologist 34(2): 61-62.

HANCOCK, D.L. 2007b. A review of Callistomyia Bezzi and related genera (Diptera:

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HANCOCK, D.L. 2007c. Notes on the genus-group placement of Peneparoxyna Hardy & Drew

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HANCOCK, D.L. 2009a. Additions and amendments to a recent classification of Dacus

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HANCOCK, D.L. 2009b. A note on Trupanea opprimata Hering (Diptera: Tephritidae).

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HANCOCK, D.L. 2010a. A review of the fruit fly tribe Tephrellini (Diptera: Tephritidae:

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HANCOCK, D.L. 2010b. Supplementary additions to a recent classification of Dacus Fabricius

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HANCOCK, D.L. 2011. Epinettyra setosa Permkam & Hancock, an Australian representative of

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species of Platensina Enderlein and Dicheniotes Munro (Diptera: Tephritidae: Tephritinae), with

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stylata and thistle receptacle weevil Rhinocyllus conicus in south eastern Australia. Proceedings

of the Eighteenth Australasian Weeds Conference, Melbourne, Victoria, pp 351-352. Available

at: http://www.caws.org.au/awc/2012/awe201213511. paf

PERMKAM, S. and HANCOCK, D.L. 1995a. Australian Ceratitinae (Diptera: Tephritidae),

Invertebrate Taxonomy 8: 1325-1341.

PERMKAM, S. and HANCOCK, D.L. 1995b. Australian Trypetinae (Diptera: Tephritidae),

Invertebrate Taxonomy 9: 1047-1209.

ROYER, J.E. and HANCOCK, D.L. 2012. New distribution and lure records of Dacinae

(Diptera: Tephritidae) from Queensland, Australia, and a description of a new species of Dacus

Fabricius. Australian Journal of Entomology 51: 239-247.

WHITE, I.M. and KORNEYEV, V.A. 1989. A revision of the western Palaearctic species of

Urophora Robineau-Desvoidy (Diptera: Tephritidae). Systematic Entomology 14: 327-374.

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Australian Entomologist, 2013, 40 (4): 237-242 237

THE FIRST RECORD AND DESCRIPTION OF A MALE IMAGO OF

AUSTREMERELLA PICTA RIEK (EPHEMEROPTERA:

EPHEMERELLIDAE)

PHILLIP J. SUTER and JULIA H. MYNOTT

Department of Environmental Management and Ecology, La Trobe University, PO Box 821,

Wodonga, Vic 3689

Abstract

Austremerella picta Riek was originally described in 1963 from nymphs and female imagos from

Lamington National Park in Queensland. A male subimago and a male imago were collected at

light traps in 2011 and 2012 respectively. Association of the nymphs with the adults was made

using a fragment of the COI gene. The subimago and imago are here described to complete the

life series for the species.

Introduction

The only species of the mayfly family Ephemerellidae known from Australia

is Austremerella picta Riek, 1963, which was described from nymphal and

adult female material from a small stream in Lamington National Park

[Queensland] (Riek 1963). No further locality information exists for the type

series, which consists of a nymph (designated the holotype) and three female

paratypes: two imagos and a damaged subimago. The nymph is clearly

distinctive and differs from all other known Australian mayfly nymphs.

The distribution of this species was extended into New South Wales, where

the nymphs were found to occupy leaf packs in flowing water, by Chessman

and Boulton (1999). Male adults have not been recorded even though

additional sites have been recognised. However, recent collections from

Lamington National Park included a male subimago and an imago, both

clearly belonging in the Ephemerellidae and from the same area as the type

series. Nymphs and a male subimago collected from the same stream as the

male imago were associated using a fragment of the СО1 gene. The male

imago is described here to complete the descriptions of all life stages.

Methods

The subimago and imago were collected using a black light and nymphs were

collected from leaf packs in the stream. Nymphs and adults were associated

using a fragment of the COI gene (assigned GenBank accession numbers

KF740665-KF740670) following the methods of Mynott et al. (2011).

Association of nymphs and adults

Genetic material was extracted from the subimago and five nymphs of

Austremerella picta (see *material examined' below) to associate the life

stages and assess the species delimitation. Specimens from the northern and

southern ends of the known distribution range (Chessman and Boulton 1999)

were sequenced. The nymphs and subimago sequenced from. Lamington

National Park formed a distinct genetic clade (Fig. 1) that confirmed the

association of the life stages. Additionally, the genetic data supported

238 Australian Entomologist, 2013, 40 (4)

Austremerella picta as a single species, with less than 196 sequence

divergence between the southern (JWA2904, JWA2905, JWA2711) and

northern (JWA2710, JWA2712, JWA2713) populations.

57 | JWA2904 Austremerella picta

51 |! JWA2711 Austremerella picta

100 | 'JWA2905 Austremerella picta

JWA2710 Austremerella picta

JWA2712 Austremerella picta

JWA2713 Austremerella picta

JQ661631.1 Penelomax, septentrionalis

JN200395.1 Penelomax, septentrionalis

0.02

Fig. 1. Neighbour-joining tree with bootstrap supports shown (2000 replications).

Distance between Austremerella clades «196 (p-distance). Outgroup data were

sourced from GenBank (Accession numbers: JN200395.1 and JQ561531.1, both

Penelomax septentrionalis).

Austremerella picta Riek

(Figs 2-7)

Austremerella picta Riek, 1963: Riek 1970; McCafferty and Wang 1997; Chessman

and Boulton 1999; Hubbard 2002; Kluge 2004.

Ephemerellina (Austremerella) picta: Allen 1965; Campbell 1990; Peters and

Campbell 1991.

Material examined. QUEENSLAND: 1 3 imago, Morans Creek above Morans Falls,

Lamington National Park (O’Reillys), -28.2318°S 153.125°E, 16.xi.2012, J. Mynott

and D. Black; 1 $ subimago, same location as imago, 17.xi.2011, J. Mynott and M.

Shackleton (Accession number: JWA2710). Deposited in the Australian National

Insect Collection (ANIC Registration No. 06-000001), along with the original type

series.

Nymphs: QUEENSLAND: Morans Creek above Morans Falls, Lamington National

Park (O'Reillys), -28.2318°S 153.125°E, 17.xi.2011, J. Mynott and M. Shackleton

(Accession numbers: JWA2712; JWA2713). NEW SOUTH WALES: Coppernook

Creek crossing Dorrigo-Coffs Harbour Road, Dorrigo National Park, -30.2925°S

152.8178°E, 10.xi2010, J. Mynott and М. Shackleton (Accession numbers:

JWA2904; JWA2905); Allyn River (upper) Allyn River Road, Barrington Tops

National Park, -32.1292°S 151.4733°E, 14.xi.2011, J. Mynott and M. Shackleton

(Accession number: JWA2711).

Description of male imago (Fig. 2). Body length 10 mm. Fore and hind wings

hyaline with distal area of costal and sub-costal region white/opaque;

forewing length 11.25 mm, width 3.75 mm; hindwing length 2.20 mm, width

1.10 mm.

Australian Entomologist, 2013, 40 (4) ` 239

Figs 2-6. Austremerella picta, male: (2) dorsal view of imago. Scale line = 1 mm; (3)

dorsal view of (a) head and thorax; (b) enlarged mesothoracic filaments indicated by

arrow. Scale line = 1 mm; (4) tarsal claws: (a) middle leg, (b) hind leg. Scale line =

0.2 mm; (5) lateral colour pattern of imago. Scale line = 1 mm; (6) ventral colour

pattern of imago. Scale line = 1 mm.

240 Australian Entomologist, 2013, 40 (4)

Eyes grey dorsally, meeting on meson of head, ventral eyes black. Pronotyum

white with central brown stripe and lateral brown marking. Mesonotuna

striped, brown and white, white anteriorly and laterally, hind margins white

with central black stripe, brown laterally; long fine filament (0.52 mm)

present near hind margin (Fig. 3). Metanotum brown.

Wing venation with costal crossveins 8 distal of bullae, 1 proximal,

pterostigma region with fine net-like veinlets, Sc crossveins 7 distal of bullae

and 2 proximal, Rs forked in proximal third of wing, MA forked in proxima]

half, МР1 not forked; MP2 joins МР1 in proximal quarter, CuA joins Cup

near base of wing, intercalaries single and not attached to main veins.

Hind wing: C meets Sc almost at apex of wing slightly concave, costal region

with 8 cross veins in distal half, no cross veins in proximal half, subcostal

region with 5 cross veins. Wing venation as shown by Riek (1963).

Foreleg lacking tarsal segments, buff coloured, brown at joints; leg segment

ratios 1.0 (2.00 mm):1.2: - . Middle and hind legs buff brown at joints, first

tarsal segment partially fused in middle leg, fused in hind leg; middle leg

segment ratios 1.0 (1.9 mm): 1.2: - : 0.01: 0.01: 0.05: 0.18; hind leg segment

ratios 1.0 (2.4 mm): 0.96: - : 0.075: 0.063: 0.042: 0.104. Tarsal claws paired,

dissimilar, one pad-like and one a short narrow blunt claw (Fig. 4).

Abdomen: dorsal colour pattern white centrally with thick brown latera]

markings (Fig. 2); segments 1-8 with fine brown line medially, broader on

segment 9 and only in anterior half of segment 10. Lateral pattern (Fig. 5)

red-brown on tergites, white dorsally. When alive the white areas were

distinctly yellow, which was lost after preservation in ethanol. Ventral colour

pattern (Fig. 6); similar to the dorsal pattern.

Male genitalia (Fig. 7): penes fused over almost entire length, bulbous

distally with slight indentation medially, no ornamentation present. Forceps

3-segmented with basal segment rectangular, length 2x width, second

segment elongate, 1.2x basal segment length and third segment short and

elongate, triangular, 0.4x length of basal segment.

Terminal filaments and cerci broken.

Male subimago. Body length 8 mm. Fore and hind wings dark grey and

opaque, forewing length 10 mm, width 3.6 mm; hindwing length 2.25 mm,

width 1.27 mm. Wing venation as for imago. Mesothorax with a pair of

posterior projections. Foreleg 5.0 mm long, buff coloured but with tarsi and

apex of tibia brown; leg segment ratios - 1.0 (1.5 mm): 1.4 : 0.05 : 0.34 : 0.27

: 0.20 : 0.15. Middle and hind legs buff without markings, first tarsal segment

partially fused in middle leg, fused in hind leg; middle leg segment ratios —

1.0 (1.7 mm): 1.18 : 0.08 : 0.09 : 0.08 : 0.04 : 0.19; hind leg segment ratios —

1.0 (1.6 mm): 1.21 : - : 0.06 : 0.06 : 0.05 : 0.2.

Australian Entomologist, 2013, 40 (4) 241

Fig. 7. Austremerella picta, male genitalia: (a) dorsal view, (b) ventral view, (c)

lateral view. Scale lines = 0.2 mm.

Discussion

Riek (1963) described the nymph and female imago of Austremerella picta

but illustrated only the fore and hind wings of the imago. Riek (1963, p. 50)

noted that the venation was similar to Ephemerella and the *wings and body

pinkish with some brownish hues'. He also noted the presence of long fine

filaments on the mesothorax. The male imago described here lacks this

coloration but when alive is a distinct yellow colour, although the wings are

hyaline. The original diagnosis for the genus was based on the nymphs and

recorded gills on segments 2-6 (Riek 1963). McCafferty and Wang (1997)

illustrated the mesothoracic filaments of the female and noted that the

nymphs had gills present on abdominal segments 2-7 and female imagos also

242 Australian Entomologist, 2013, 40 (4)

had gill socket vestiges on abdominal segments 2-7. Kluge (2004) also noted

the presence of the small vestigial gills that are covered by the gill on

abdominal segment 6.

The male imago described above is consistent with the wing characteristics

and also has the long fine mesothoracic filaments. Association of the life

stages was confirmed using genetic data. Additionally, the genetic data

supported Austremerella picta as a single species ranging from southern

Queensland to north of the Hunter River in New South Wales.

Austremerella picta can be distinguished from all other Australian adult

mayflies by the presence of the thoracic filaments, dissimilar tarsal claws and

forewing with single, free marginal intercalaries.

Acknowledgements

Specimens were collected under permit number WITK10277111

(Queensland) and scientific licence SL100434 (New South Wales). Funding

was provided through ABRS grant number CT211-32.

References

ALLEN, R.K. 1965. A review of the subfamilies of Ephemerellidae (Ephemeroptera). Journal of

the Kansas Entomological Society 38: 262-266.

CAMPBELL, I.C. 1988. Ephemeroptera. In: Zoological Catalogue of Australia 1-22. Bureau of

Flora and Fauna, Burwood.

CAMPBELL, LC. 1990. The Australian mayfly fauna: composition, distribution and

convergance. Pp 149-153, in: Campbell, I.C. (ed.), Mayflies and stoneflies: life histories and

biology. Kluwer Academic Publishers, Dordrecht.

CHESSMAN, B.C. and BOULTON, A.J. 1999. Occurrence of the mayfly family Teloganodidae

in northern New South Wales. Australian Journal of Entomology 38: 96-98.

HUBBARD, M.D. 2002. Synonymy and valid name of the families Vietnamellidae and

Austremerellidae (Ephemeroptera: Ephemerelloidea). Florida Entomologist 85: 382.

KLUGE, N. 2004. The phylogenetic system of Ephemeroptera. Kluwer Academic Publishers,

Dordrecht/Boston/London.

McCAFFERTY, W.P. and WANG, T.Q. 1997. Phylogenetic systematics of the family

Teloganodidae (Ephemeroptera: Pannota). Annals of the Cape Provincial Museums of Natural

History 19: 387-437.

MYNOTT, J.H., WEBB, J.M. and SUTER, P.J. 2011. Adult and larval associations of the alpine

stonefly genus Riekoperla McLellan (Plecoptera: Gripopterygidae) using mitochondrial DNA.

Invertebrate Systematics 25: 11-21.

PETERS, W.L. and CAMPBELL, I.C. 1991. Ephemeroptera (Mayflies). Pp 279-293, in: The

Insects of Australia - A textbook for students and research workers. Melbourne University Press,

Melbourne.

RIEK, E.F. 1963. An Australian mayfly of the family Ephemerellidae (Ephemeroptera). Journal

of the Entomological Society of Queensland 2: 48-50.

RIEK, E.F. 1970. Ephemeroptera (Mayflies). Pp 224-240, in: CSIRO, The Insects of Australia.

Melbourne University Press, Melbourne.

Australian Entomologist, 2013, 40 (4): 243-247 243

THE GENUS CRYPHAEUS KLUG, 1833 IN AUSTRALIA, WITH

DESCRIPTION OF A NEW SPECIES (COLEOPTERA:

TENEBRIONIDAE: TOXICINI)

ROLAND GRIMM

Unterer Sdgerweg 74, 75305 Neuenbürg, Germany (Email: grimm.tenebrio@t-online.de)

Abstract

Cryphaeus wachteli sp. n. (Australia: NSW and Qld) is described and compared with its

Melanesian congeners. The previously suggested occurrence of C. chevrolati (Montrouzier,

1855) in Australia is discussed and regarded as based on mislabelled specimens.

Introduction

The tribe Toxicini is subdivided into the subtribes Eudysantina, Nycteropina,

and Toxicina (Bouchard et al. 2005). In Australia the Eudysantina and

Toxicina are represented (Matthews and Bouchard 2008). The subtribe

Toxicina includes the two genera Toxicum Latreille, 1802 and Cryphaeus

Klug, 1833. According to Merkl (1989), in Cryphaeus the eyes are

completely divided by the epistomal canthi and the head of males has two

glabrous supraorbital horns and lacks epistomal horns. In Toxicum the eyes

are not completely divided and males are equipped with two supraorbital

horns fringed with yellow hairs as well as epistomal horns. Recently,

Schawaller (2006) described a Cryphaeus species with epistomal horns from

South Africa and concluded that Cryphaeus is perhaps a junior subjective

synonym of Toxicum but, according to him, the problem needs to be studied

in greater detail.

Toxicum is listed for Australia by Carter (1926), Doyen et al. (1989) and

Matthews and Bouchard (2008) but none of these standard works cite the

genus Cryphaeus from the continent. However, according to Merkl (1989)

Cryphaeus is widely distributed in the Palearctic, Afrotropical and

Indomalayan realms as well as Melanesia and Australia. For Australia, Merkl

(1989) mentioned ‘unidentified specimens belonging probably to new

species’ and cited specimens of C. chevrolati (Montrouzier, 1855) in HNHM

from ‘northern Queensland’. Fieldwork by Franz Wachtel (Egling, Germany)

in New South Wales and Queensland yielded a new species of Cryphaeus

which is described below. Examination of the Australian specimens in

HNHM mentioned by Merkl (1989) showed that they belong to the same

species. Additional specimens were found in the Australian National Insect

Collection and the Queensland Museum.

The acronyms of the following depositories are given in parentheses: AM —

Australian Museum, Sydney, Australia; ANIC — Australian National Insect

Collection, Canberra, Australia; CRG - Collection Roland Grimm,

Neuenbürg, Germany; HNHM - Hungarian Natural History Museum,

Budapest, Hungary; QM — Queensland Museum, Brisbane, Australia; SMNS

— Staatliches Museum für Naturkunde, Stuttgart, Germany.

244 Australian Entomologist, 2013, 40 (4)

Taxonomy

Cryphaeus wachteli sp. n.

(Figs 1-4)

Material examined. Holotype á, NEW SOUTH WALES: Macksville, i.[19]91, [F.]

Wachtel (AM: K.349042). Paratypes: 2 99, same data as holotype (AM: K.349043

and K.349044), 3 99, same data as holotype (CRG), 1 9, same data as holotype

(SMNS); 1 c, Maxville [sic], v.1981, [no collector stated] (HNHM); 1 3, Greta,

ix.1951, J. Sedlacek (HNHM); 1 3, Araluen, 25.xi.1978, J. Е. Lawrence, Lot 78-195,

Osmophorus latus (ANIC); 1 д, Kangaroo Valley, 7.x.1986, C. Reid, at light (ANIC);

1 3, Sandy Creek Road, Braemar SF south of Casino, 16.x.1997, Watkins, 97:567,

mv light in open forest; S.G. Watkins Collection, donated 2001 (ANIC).

QUEENSLAND: 2 99, Mt Maroon, Border Range, i.[19]95, [F.] Wachtel (QM &

CRG); 1 д, Brisbane [no date stated], J. Sedlagek (HNHM); 1 9, South Percy Island

(Lagoon Area), 21°46’S 150°18’E, 25.xi.1992, rainforest, Monteith, Thompson &

Janetzki, pyrethrum (QM); 1 9, Lever’s Plateau, 18.iv.1964, D.F. O'Sullivan (QM); 1

$, N of Maryborough, Salt Water Creek, 29°29’S, 152°43°E, 11.viii.1986, К.

Eastwood (QM); 1 9, Millmerran, 22.v.1966, H. Burton (QM); 1 9, Biggenden

(25.315 152.03E), under bark of blue gum, 13.viii.1972, Н. Frauca (ANIC).

Description. Elongate-oblong, slender, subconvex transversally, dull black.

Body length 8.2-10.9 mm, body width 2.5-3.5 mm. Head coarsely and

densely punctured; distances among punctures somewhat smaller than their

diameters; punctures of genae and epistoma smaller; epistoma nearly straight

anteriorly, with weakly sinuate canthi. Antennal club 3-segmented;

antennomere 9 nearly twice as wide as antennomere 8. Mentum

subcordiform. Pronotum transverse, width/length ratio 1.18-1.25, moderately

convex; sides subparallel, very weakly sinuate in the middle and before

posterior angles; anterior angles rounded, not protruding beyond anterior

margin; pronotal punctures coarse as on head, distances among them usually

equal to puncture diameter. Elytra elongate, with regular rows of punctures.

Ventral surface and legs without any modification.

Male (Figs 1-2). Supraorbital horns long, slightly incurved in frontal view,

pointing forward in lateral view; they may be reduced to rather large

tubercles. Cranial concavity punctures coarse and deep; distances among

them various; shining. Aedeagus as in Fig. 3.

Female. Cranial surface between supraorbital swellings slightly concave, set

with coarse punctures; distances among them various but mostly less than

their diameters.

Differential diagnosis. According to Merkl (1989), five species of the genus

Cryphaeus are known from the Melanesian region, of which C. nudicornis

(Fairmaire, 1883) differs from C. wachteli sp. n. in having a 4-segmented

antennal club. Cryphaeus irregularis (Gebien, 1920) and C. vacca Merkl,

1989 have a rather broad body and the rows of punctures on the elytra are

irregular. Cryphaeus chevrolati has a conspicuous cinnabar-red pronotum

Australian Entomologist, 2013, 40 (4) 245

(often with a black marking along the midline) and C. biroi (Kaszab, 1939) is

deep black with reddish legs and the anterior corners of the pronotum

protrude beyond the anterior margin.

Figs 1—3. Cryphaeus wachteli sp. n.: (1-2) holotype: (1) dorsal; (2) head lateral. (3)

aedeagus: (a) ventral; (b) dorsal.

246 Australian Entomologist, 2013, 40 (4)

Etymology. This species is named in honour of Franz Wachtel (Egling, Germany), one

of the collectors of the type series.

Distribution. Coastal regions of NSW and southern Qld (Fig. 4).

Fig. 4. Distribution of Cryphaeus wachteli sp. n.

Australian Entomologist, 2013, 40 (4) 247

Discussion

As mentioned by Merkl (1989), in HNHM there are two females of C.

chevrolati labelled from Northern Queensland: Endeavour River [no date or

collector stated]. Evidently these specimens are mislabelled. No additional

specimens of this conspicuous, bicoloured Melanesian species could be found

in Australian museum collections, despite Australian museums having done

an enormous amount of field work in North Queensland near the Endeavour

River (Monteith pers. comm.). In discussing erroneous Australian records of

the Melanesian phasmid Eurycantha calcarata Lucas, 1869, Monteith and

Dewhurst (2011) pointed out that the Endeavour River was an early port of

transshipment for exploratory ships carrying specimens back from New

Guinea and other Melanesian islands and shipments sometimes received the

name of the locality of transhipment port rather than their true origin.

Acknowledgements

Cordial thanks are due to Drs Ottó Merkl (Budapest), Geoff Monteith

(Brisbane), Cate Lemann and Tom Weir (Canberra) for the loan of material,

and Johannes Reibnitz (Stuttgart) who produced the figures and assembled

the plates. Two anonymous referees improved the manuscript with their

comments.

References

BOUCHARD, P., LAWRENCE, J.F., DAVIES, A.E. and NEWTON, A.F. 2005. Synoptic

classification of the world Tenebrionidae (Insecta: Coleoptera) with a review of family-group

names. Annales Zoologici 55: 499-530.

CARTER, H.J. 1926. A check list of the Australian Tenebrionidae. Australian Zoology 4: 117-

163, 280, 294, pls XVI-XVII.

DOYEN, T., MATTHEWS, E.G. and LAWRENCE, J.F. 1989. Classification and annotated

checklist of the Australian genera of Tenebrionidae (Coleoptera). Invertebrate Taxonomy 3: 229-

260.

MATTHEWS, E.G. and BOUCHARD, P. 2008. Tenebrionid beetles of Australia — description

of tribes, key to genera, catalogue of species. Australian Biological Resources Study, Canberra;

398 pp.

MERKL, O. 1989. Melanesian representatives of Toxicum and Cryphaeus (Coleoptera,

Tenebrionidae: Toxicini). Acta Zoologica Hungarica 35: 235-254.

MONTEITH, G.B. and DEWHURST, C.F. 2011. Does the phasmid Eurycantha calcarata

Lucas, 1869 (Phasmida: Phasmatidae) occur in Australia? Australian Entomologist 38: 179-196.

SCHAWALLER, W. 2006. A new species of Cryphaeus and new records of other fungus

adapted tenebrionids from South Africa (Coleoptera: Tenebrionidae). Annals of the Transvaal

Museum 43: 69-74.

248 Australian Entomologist, 2013, 40 (4)

BOOK NOTICE

Field Guide to the

damselflies of

New Guinea

Brachytron

Illustrated by Albert Orr

This attractive, newly released book on the damselflies (Odonata) of New Guinea has

been produced as a 128-page supplementary volume to the Dutch Society for

Dragonfly Studies? journal Brachytron and is available from them for €15. For details

see their website: http:/Avww.brachytron.nl/Brachytron/Brachytron] 6supinhoud.html.

It is also available locally for $22 + p&p from Albert Orr at agorr@bigpond.com. The

book is also featured on the website of the Papuan Insects Foundation at

http://www.papua-insects.nl/ and shared the 2013 Whitley Medal awarded by the

Royal Zoological Society of NSW.

With it, the reader will be able to identify all genera and most species of damselflies

recorded from the island of New Guinea, including its satellite islands and the

Bismarck Archipelago. The guide contains nearly 300 colour illustrations and over

250 line drawings by Albert Orr and 16 colour photographs taken in the field by

Stephen Richards, both recent contributors to the Australian Entomologist. Many

genera and most included species have never been depicted in colour before. It is

especially intended to stimulate local interest in the fauna, but is an essential reference

for anyone interested in New Guinea Odonata.

THE

AUSTRALIAN

ENTOMOLOGIST

VOLUME 40

2013

Published by:

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THE AUSTRALIAN ENTOMOLOGIST

Contents: Volume 40, 2013

BARTLETT, J.S.

Thriocerodes anthicides (Newman), comb. n. (Coleoptera: Cleridae): an

Australian, not South African, species

BARTLETT, J.S. and SZITO, A.

Opetiopalpus scutellaris Panzer (Coleoptera: Cleridae: Korynetinae)

established in the Western Australian wheatbelt

DAVID, K.J. and HANCOCK, D.L.

The first record of Ortalotrypeta isshikii (Matsumura) and subfamily

Tachiniscinae (Diptera: Tephritidae) from India, with redescription

of the species

FERGUSON, D.J. and YEATES, D.K.

The courtship behaviour of the bee fly Meomyia vetusta Walker

(Diptera: Bombyliidae)

GERSTMEIER, R. and SEITNER, M.

Kanaliella gen. n., a new genus of Cleridae from the Melanesian Region

(Coleoptera: Cleridae: Clerinae)

GRIMM, R.

The genus Cryphaeus Klug, 1833 in Australia, with description of a new

species (Coleoptera: Tenebrionidae: Toxicini)

HALSEY, M.

Notes on the ecology, phenology and distribution of Pollanisus cyanotus

(Meyrick, 1886) (Lepidoptera: Zygaenidae: Procridinae: Artonini)

HANCOCK, D.L.

Themara maculipennis (Westwood) and Themara hirtipes Rondani

(Diptera: Tephritidae: Acanthonevrini): a case of confused synonymies

A yellow-form male of Delias mysis mysis (Fabricius) (Lepidoptera:

Pieridae) from northern Queensland

A revised checklist of Australian fruit flies (Diptera: Tephritidae)

HOPKINSON, M.

Life history notes and new food plant records for Graphium macfarlanei

macfarlanei (Butler, 1877) (Lepidoptera: Papilionidae: Papilioninae)

HOUSTON, T.F. and PIKE, D.T.

Aerial brood cells constructed by some Australian resin bees

(Hymenoptera: Megachilidae) and a case of gregarious nesting

JOHNSON, S.J. and JOHNSON, LR.

The first record of Paraduba metriodes (Bethune-Baker) (Lepidoptera:

Lycaenidae) within Australian limits

KING, D.

A provisional range extension for Theclinesthes sulpitius (Miskin)

(Lepidoptera: Lycaenidae) in Western Australia

Range extension of Acraea terpsicore (Linnaeus) (Lepidoptera:

Nymphalidae) to Western Australia

iii

131

243

219

101

173

111

118

KOHOUT, R.J.

A review of the Polyrhachis continua species-group of the subgenus

Myrma Billberg (Hymenoptera: Formicidae: Formicinae) with keys

and descriptions of new species

Polyrhachis (Myrmothrinax) nepenthicola, a new species of the thrinax-

group inhabiting pitcher plants (Hymenoptera: Formicidae: Formicinae)

A review of the Polyrhachis aculeata species-group of the subgenus

Myrma Billberg (Hymenoptera: Formicidae: Formicinae), with keys

and descriptions of new species

LACHLAN, R.B. and MULLER, C.J.

A new species of Dicallaneura Butler (Lepidoptera: Lycaenidae:

Riodininae) from Papua New Guinea

LAMBKIN, T.A.

A revision of the Euploea batesii C. & R. Felder, 1865 ‘complex’

(Lepidoptera: Nymphalidae: Danainae) in mainland Papua New

Guinea and Australia, including its biology and biogeography

MARINOV, M. and RICHARDS, S.

Notes on the female colour forms of Synthemis miranda Selys, 1871

(Odonata: Synthemistidae) in New Caledonia

MEYER, C.E., WEIR, R.P and BROWN, S.S.

Some new and interesting butterfly (Lepidoptera) distribution and

temporal records from Queensland and northern Australia

MOUND, L.A. and DANG, L.-H.

New Australian records of Asian or New Zealand Phlaeothripidae

(Thysanoptera)

ORR, A.G.

Predation on butterflies and other insects by breeding Rainbowbirds

(Merops ornatus: Meropidae) in south-east Queensland

SHAKESPEARE, T.J., SHAKESPEARE, Z.J. and SHAKESPEARE, T.P.

An observation of Polyura sempronius (Fabricius) (Lepidoptera:

Nymphalidae) feeding on bandicoot droppings in south-east Queensland 175

SMITH, G. and IRISH, J.

Further Australian locality data for Ctenolepisma rothschildi Silvestri

119

(Zygentoma: Lepismatidae) 99

SUTER, P.J. and MYNOTT, J.H.

The first record and description of a male imago of Austremerella picta

Riek (Ephemeroptera: Ephemerellidae) 237

CORRIGENDA 56

MISCELLANEOUS NOTES 172

BOOK NOTICE 248

Publication dates: Part 1 (pp 1-56) 20 February 2013

Part 2 (pp 57-100) 21 May 2013

Part 3 (pp 101-172) 26 August 2013

Part 4 (pp 173-248) 29 November 2013

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°° THE AUSTRALIANO

der

At 5

Gea 40; Part 4, 29 j November 2013

ЖЖ

CONTENTS

GRIMM, R.

The genus Cryphaeus Klug, 1833 in Australia, with description of a new species

(Coleoptera: Tenebrionidae: Toxicini)

HANCOCK, D.L.

A revised checklist of Australian fruit flies (Diptera: Tephritidae)

JOHNSON, S.J. AND JOHNSON, I.R.

The first record of Paraduba metriodes (Bethune-Baker) (Lepidoptera: Lycaenidae)

within Australian limits

_ _—_—————

LACHLAN, R.B. AND MULLER, C.J.

A new species of Dicallaneura Butler (Lepidoptera: Lycaenidae: Riodininae) from

Papua New Guinea

а ола eee

LAMBKIN, Т.А.

A revision of the Euploea batesii C. & К. Felder, 1865 ‘complex’ (Lepidoptera:

Nymphalidae: Danainae) in mainland Papua New Guinea and Australia,

including its biology and biogeography

SHAKESPEARE, T.J., SHAKESPEARE, Z.J. AND SHAKESPEARE, T.P.

An observation of Polyura sempronius (Fabricius) (Lepidoptera: Nymphalidae)

feeding on bandicoot droppings in South-East Queensland

SUTER, P.J. AND MYNOTT, J.H.

The first record and description ofa male imago of Austremerella picta Riek

(Ephemeroptera: Ephemerellidae)

E a ae lS RUP

BOOK NOTICE

ISSN 1320 6133

DPI