THE AUSTRALIAN

Entomologist

published by

THE ENTOMOLOGICAL SOCIETY OF EENSLAND

Volume 44, Part 2, 30 June 2017

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ISSN 1320 6133

THE AUSTRALIAN ENTOMOLOGIST

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COVER

The giant wingless carabid, Nurus rex Darlington 1961, at the entrance to its burrow

under a rainforest tree root. The species occurs only in a small cap of rainforest

on the summit of the 1000 m Mt Elliot, just south of Townsville, and was first

collected by the noted Harvard biogeographer, Philip Darlington, when he made the

first entomological ascent of the mountain in March 1958. It is the largest and most

northerly of about a dozen species in its genus, all of which are now known to live

in burrows with a cleared entrance court where they ambush passing invertebrates

at night. Pen and ink drawing by Caloundra ESQ member, Dr Albert Orr, whose

illustrated books on dragonflies and butterflies have won awards in Australia and

overseas.

Australian Entomologist, 2017, 44 (2): 57-60 57

POPPLEPSALTA AEROIDES OWEN & MOULDS (HEMIPTERA:

CICADIDAE): DESCRIPTION OF THE FEMALE

D.L. EMERY! and N.J. EMERY?

! School of Veterinary Science, University of Sydney, NSW 2006

(Email: david.emery@sydney.edu.au)

”The Australian Plant Bank, the Royal Botanic Gardens and Domain Trust, the Australian

Botanic Garden, Mount Annan, NSW 2567

Abstract

The female of the cicada species Popplepsalta aeroides Owen & Moulds is described. Features

which differentiate P. aeroides from P. rubristrigata (Goding & Froggatt) are included.

Introduction

In their recent review of the Australian cicada genus Pauropsalta Goding &

Froggatt, Owen and Moulds (2016) erected the genus Popplepsalta Owen &

Moulds to accommodate 16 species, one of which was new. The new species,

Popplepsalta aeroides Owen & Moulds, 2016, was described from eight

males collected from southern Queensland, the Sydney region and south

coastal New South Wales (Owen and Moulds 2016). The female remained

unknown. Here we provide the first description of the female of Popplepsalta

aeroides from four specimens: three from Royal National Park, south of

Sydney and one from Upper Dalrymple Creek in SE Queensland.

Abbreviations. DE — private collection of Prof. D. Emery, Sydney; LP —

private collection of Dr L. Popple, Brisbane; QM — Queensland Museum,

Brisbane.

Popplepsalta aeroides Owen & Moulds

(Figs 1-3)

Material examined. NEW SOUTH WALES: 1 9, Garie Beach, Royal National Park,

34?10'08"S 151?03'56"E, 13.1.2012, S., N. & D. Emery (LP); 1 9, same location,

29.1.2012, N. & D. Emery; 1 9, same location, 24.xii.2016, N. & D. Emery (DE).

QUEENSLAND: 1 9, Upper Dalrymple Ck via Goomburra, 21-22.x1.1987, G.B.

Monteith (QM).

Description of female. Head about as wide as lateral angles of pronotal collar,

predominantly black, with golden pubescence. Vertex with spot at posterior

midline. Postclypeus black with orange-brown markings, a reddish brown

spot on midline around ventral anterior segment, lateral and posterior margins

orange-brown; transverse ridges black, central groove distinct. Lorum black,

gena black. Anteclypeus black along midline, brown laterally. Rostrum

ochraceous, becoming darker posteriorly, reaching the posterior margin of the

mid coxae. Antennae black, brown at base. Supra-antennal plates brown with

reddish anterior margin.

Thorax. Pronotum black, with ochraceous and reddish brown markings,

anterior margin orange-brown, fascia along midline orange-brown centrally,

58 Australian Entomologist, 2017, 44 (2)

ochraceous laterally, expanded laterally at pronotal collar; pronotal collar

reddish brown along posterior margin, broadening at lateral margin.

Mesonotum primarily ochraceous brown, lateral and submedian sigellae

black, scutal depressions black; cruciform elevation reddish brown, a black

fascia along anterior midline. Metanotum black at hindwing base, edged

orange-brown or ochraceous along dorsal ridge. Legs mainly ochraceous

brown, with reddish longitudinal stripes on femora; femoral spines brown

with black tips; fore and mid tibiae reddish brown, hind tibiae pale brown;

tarsi reddish brown, claws brown at base, black at tips. Meracanthus

triangular, black basally, cream to ochraceous over posterior half.

Wings hyaline. Forewing costa orange-red; veins M and CuA fused before

meeting basal cell, venation red to brownish red tending black distally; basal

membrane pale grey to blackish. Hind wing with 6 apical cells, venation red

to brownish red becoming black distally, basal cell translucent, basal

membrane grey; plaga muddy white to pale brown, limited to edges of vein

3A; black infuscation on wing margin at distal end of vein 2A.

Abdomen. Tergites predominantly black; tergites 1 and 2 entirely black;

tergites 3-7 with lateral margins increasingly reddish brown, posterior

intersegmental membranes blue to varying degrees, fading to yellow-brown

on dried specimens; tergite 8 black anteriorly, brownish red centrally,

posterior margin greenish blue. Sternite I ochraceous; sternites II-VII

ochraceous, lateral sides and posterior margin reddish brown and greenish

blue to varying degrees, with black spot on posterior midline of each sternite;

abdominal segment 9 black along anterior margin, black dorsally with brown

stripe along midline, reddish-brown laterally, blue over ventral surface;

dorsal beak black, anal styles red-brown with black posterior margins;

ovipositor sheath black; ovipositor brown, sheath and ovipositor extending

23 mm beyond abdominal segment 9.

Measurements. Range and mean (in mm) for 3 99; includes smallest and

largest of available specimens. Length of body: 21.1-23.1 (22.1). Length of

forewing: 22.6-25.5 (23.8). Width of forewing: 7.6-8.0 (7.8). Width of head

(including eyes): 5.7-6.5 (6.1). Width of pronotum (across lateral angles):

5.3-6.6 (5.8). Abdominal width (across segment II): 4.8-5.8 (5.0). Ovipositor

length: 9.4-10.9 (9.9).

Distinguishing features. In the original description of this species, Owen and

Moulds (2016) noted the similarity between males of Popplepsalta aeroides

and P. rubristrigata (Goding & Froggatt, 1904). The principal distinguishing

feature was the blue coloration of the intersegmental membranes on

abdominal segments 3-7. This also applies to the female but, with drying, the

blue coloration fades and reduces the reliability of this character. Therefore,

P. aeroides can also be distinguished from P. rubristrigata by the following

combination of characters: body length «24.0 mm; ovipositor length «10.0

mm; and forewing width «10.0 mm.

Australian Entomologist, 2017, 44 (2) 59

a

cou es Bite ae Se Re ee

AE uti ET d

pt UR eng

PR

a

Figs 1-3. Popplepsalta aeroides (Garie Beach, NSW): (1) dorsal habitus of female;

(2) ventral habitus of female; (3) live specimens of the male (left) and female (right).

Scale bars = 10 mm.

60 Australian Entomologist, 2017, 44 (2)

Behaviour. Like P. rubristrigata, females of P. aeroides are found within 2-3

m of the ground, while males call from >5 m in taller eucalypts. Oviposition

by females of both species occurs in green branches of young eucalypts

within 5 m of the ground.

Acknowledgements

The authors thank Samantha Emery for assistance in the field. We also thank

Drs Tony Ewart and Lindsay Popple for constructive comments that

improved the manuscript.

Reference

OWEN, C.L. and MOULDS, M.S. 2016. Systematics and phylogeny of the Australian cicada

genus Pauropsalta Goding and Froggatt, 1904 and allied genera (Hemiptera: Cicadidae:

Cicadettini). Records of the Australian Museum 68(4): 117-200. http://dx.doi.org/10.3853/

J.2201-4349.68.2016.1598

Australian Entomologist, 2017, 44 (2): 61-62 61

MESONIRVANA EVANS, 1956, REPLACES EOSCARTOIDES EVANS,

1956 (HEMIPTERA: CICADOMORPHA: PROSBOLOIDEA:

DYSMORPHOPTILIDAE), A HOMONYM OF EOSCARTOIDES

MATSUMURA, 1940 (HEMIPTERA: CERCOPIDAE)

KEVIN J. LAMBKIN

Queensland Museum, South Brisbane, Old 4101 (Email: megapsychops@bigblue.net.au)

Abstract

Mesonirvana Evans, 1956, is the available replacement name for Eoscartoides Evans, 1956,

which is a homonym of Eoscartoides Matsumura, 1940. The replacement means that the names

for the three species formerly ascribed to Eoscartoides Evans are now: Mesonirvana abrupta

Evans, 1956 (= Eoscartoides bryani Evans, 1956), Mesonirvana orthoclada (Tillyard, 1922),

comb. n. and Mesonirvana dmitryi (Lambkin, 2016), comb. n.

Discussion

In his review of Palaeozoic and Mesozoic Hemiptera, Evans (1956)

established the monotypic genera Eoscartoides Evans, with type species £.

bryani Evans, 1956 and Mesonirvana Evans, with type species M. abrupta

Evans, 1956, both from the Late Triassic Mount Crosby Formation of

Queensland. In my revision of the Dysmorphoptilidae of the Queensland

Triassic (Lambkin 2016), I identified the two type species as synonyms. As

both generic names were established in the same paper, as first reviser (ICZN

Article 24), I chose Eoscartoides as the senior generic synonym on the

grounds that its type specimen was better preserved and that all subsequent

published specimens (Evans 1961) were identified as E. bryani. At that time I

also ascribed two additional species to the genus — E. orthocladus (Tillyard,

1922) (transferred from Mesocixiodes Tillyard, 1922) and E. dmitryi

Lambkin, 2016. Eoscartoides Evans, however, is a homonym of the cercopid

Eoscartoides Matsumura, 1940 (now considered a synonym of Eoscarta

Breddin, 1902 — see Liang 1996). As its junior synonym, Mesonirvana is

available as the replacement name for Eoscartoides Evans (ICZN Article

23.3.5). The nomenclatural adjustments associated with this replacement are

summarised as follows:

Mesonirvana Evans, 1956

Mesonirvana Evans, 1956: 191-192. Type species Mesonirvana abrupta Evans, 1956,

by original designation.

Eoscartoides Evans, 1956: 220, junior homonym (non Eoscartoides Matsumura,

1940). Type species Eoscartoides bryani Evans, 1956, by original designation.

Three included species:

Mesonirvana abrupta Evans, 1956: 192, fig. SE (= Eoscartoides bryani

Evans, 1956: 221, fig. 18A; Evans 1961: 20, figs 4A-4C).

Mesonirvana orthoclada (Tillyard, 1922): 463, text-fig. 83. Comb. n.

Mesonirvana dmitryi (Lambkin, 2016): 211-213, figs 7-9. Comb. n. (Fig. 1).

62 Australian Entomologist, 2017, 44 (2)

Fig. 1. Mesonirvana dmitryi (Lambkin): holotype specimen in the Queensland

Museum (QM F58591) from the Late Triassic fossil insect locality at Dinmore, south-

east Queensland.

Acknowledgements

I am very grateful to John MacConnell of North Carolina for drawing my

attention to the homonymy and to Geoff Thompson of the Queensland

Museum for the photograph.

References

EVANS, J.W. 1956. Palaeozoic and Mesozoic Hemiptera (Insecta). Australian Journal of

Zoology 4: 165-258.

EVANS, J.W. 1961. Some Upper Triassic Hemiptera from Queensland. Memoirs of the

Queensland Museum 14: 13-23.

LAMBKIN, K.J. 2016. Revision of the Dysmorphoptilidae (Hemiptera: Cicadomorpha:

Prosboloidea) of the Queensland Triassic—Part 2. Zootaxa 4092: 207-218.

LIANG, A.-P. 1996. The spittlebug genus Eoscarta Breddin of China and adjacent areas

(Homoptera: Cercopidae). Oriental Insects 30: 101-130.

MATSUMURA, S. 1940. New species and genera of Cercopidae in Japan, Korea and Formosa,

with a list of the known species. Journal of the Faculty of Agriculture, Hokkaido Imperial

University 45: 35-82.

TILLYARD, R.J. 1922. Mesozoic insects of Queensland. No. 9. Orthoptera, and additions to the

Protorthoptera, Odonata, Hemiptera and Planipennia. Proceedings of the Linnean Society of New

South Wales 47: 447-470.

Australian Entomologist, 2017, 44 (2): 63-64 63

A TENTATIVE RECORD OF PAPILIO DEMOLEUS MALAYANUS

WALLACE, 1865 (LEPIDOPTERA: PAPILIONIDAE) FROM THE

SOLOMON ISLANDS

JOHN E. NIELSEN

C/- Pathway Surveillance and Operational Science, Department of Agriculture and Water

Resources, GPO Box 858, Canberra, ACT 2602

Abstract

Commercially obtained specimens of Papilio demoleus malayanus Wallace, 1865, ostensibly

from Guadalcanal Province, Solomon Islands, are presented as a tentative new record. Further

specimens are needed for confirmation.

Introduction

Papilio demoleus Linneaus, 1758 has a wide distribution from the Middle

East and throughout southern Asia to southern mainland Papua New Guinea

and Australia (Smith and Vane-Wright 2010). Recently, the Southeast Asian

subspecies P. demoleus malayanus Wallace, 1865 has spread through the

Indonesian archipelago into Papua New Guinea (Tennent et al. 2011, Morgun

and Wiemers 2012).

Specimens of P. d. malayanus obtained from a commercial source,

purporting to originate from Guadalcanal Province of the Solomon Islands,

are presented here as a tentative range extension.

New record

This record is based on three females of P. demoleus malayanus (Figs 1-3)

obtained from a commercial insect dealer in Poland. Data provided with the

specimens were 'Guadalcanal, Solomon Islands, April 2015'. Identity of

these specimens was confirmed using the key in Smith and Vane-Wright

(2010).

Discussion

The record represented by these specimens is considered probably genuine —

but with qualification. It is considered unlikely that they were passed off as a

new distribution record for profit because P. demoleus forms part of the ‘low

value, high volume” insect trade (Collins and Morris 1986) and specimens are

inexpensive. Papilio demoleus specimens obtained for another project also

lacked geographic precision (e.g. simply ‘West Java’ or ‘Ceram’ [Indonesia])

but were still accurate to the resolution provided. For these reasons, it is

concluded that the specimens presented here are evidence that P. d.

malayanus has reached Guadalcanal Province, Solomon Islands, a conclusion

supported by the presence of this taxon in nearby New Britain, New Ireland

and Milne Bay Provinces, Papua New Guinea (Tennent ef al. 2011).

However, confirmation of this record is needed, making it inappropriate at

present to use it to justify biosecurity measures for P. d. malayanus on trade

pathways from the Solomon Islands.

64 Australian Entomologist, 2017, 44 (2)

Figs 1-3. Papilio demoleus malayanus specimens representing a putative record from

Guadalcanal Province, Solomon Islands. Specimen (1) in author's collection; (2) and

(3) in Australian National Insect Collection (ANIC), Canberra.

Acknowledgements

You-Ning Su and Ted Edwards accepted voucher specimens forming the basis of this

tentative record for deposition in the ANIC and John Tennent provided literature.

Caroline Martin and Gertraud Norton assisted with departmental clearance of the

manuscript. I am grateful to my wife Haliz and Albert Orr for encouraging my interest

in the Lepidoptera and to Pawel Lewanthal, Poland, for supplying the specimens.

References

COLLINS, N.M. and MORRIS, M.G. 1985. Threatened swallowtail butterflies of the world: the

IUCN red data book. IUCN, Gland, Switzerland; vii + 401 pp.

MORGUN, D.V. and WIEMERS, M. 2012. First record of the lime swallowtail Papilio

demoleus Linneaus, 1758 (Lepidoptera, Papilionidae) in Europe. Journal of Research on the

Lepidoptera 45: 85-89.

SMITH, C.R. and VANE-WRIGHT, R.I. 2008. Classification, nomenclature and identification

of lime swallowtail butterflies: a post-cladistic analysis (Lepidoptera: Papilionidae). Systematics

and Biodiversity 6: 175-203.

TENNENT, W.J., DEWHURST, C.F. and MÜLLER, C.J. 2011. On the recent spread of Papilio

demoleus Linnaeus, 1758, in eastern Papua New Guinea (Lepidoptera, Papilionidae). Butterflies

(Teinopalpus) 58: 30-33.

Australian Entomologist, 2017, 44 (2): 65-74 65

PAPILIO DEMOLEUS MALAYANUS WALLACE, 1865

(LEPIDOPTERA: PAPILIONIDAE) ON DAUAN ISLAND, TORRES

STRAIT, QUEENSLAND AND RECENT CONFIRMATION OF P. D.

STHENELINUS ROTHSCHILD, 1895 IN THE LESSER SUNDA

ISLANDS

TREVOR A. LAMBKIN

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

(Email: trevor.lambkin(a)uqconnect.edu.au or t.lambkin@hotmail.com)

Abstract

The lime swallowtail, Papilio demoleus malayanus Wallace, 1865, is recorded for the first time

in Australia from Dauan Island, Torres Strait, Queensland in 2010. Subsequent reports from

January and December 2016 and in January 2017 indicate that it is established on the island, with

oviposition observed on Citrus. Originally confined to the Malay Peninsula, P. d. malayanus has

expanded its range eastward since the 1960s. In addition, the current study has confirmed that P.

d. sthenelinus Rothschild, 1895 is still present on Flores in the Lesser Sunda Archipelago

together with P. d. malayanus, suggesting that the two taxa might be separate species. The same

might be true for P. d. malayanus and P. d. sthenelus, as biological data presented here supports

published molecular data confirming a strong separation between the two taxa.

Introduction

The Papilio demoleus Linnaeus, 1758 species group, often referred to as the

‘lime swallowtails', has long been recognised for its distinctiveness within

Papilio L. (Munroe 1961, Hancock 1983, Smith and Vane-Wright 2008). The

group is predominantly tropical with five recognised species: P. demodocus

Esper, 1799 from the Afrotropics; P. erithonioides Grose Smith, 1891, P.

morondavana Grose Smith, 1891 and P. grosesmithi Rothschild, 1926 from

Madagascar; and P. demoleus from the Indo-Australian region (Smith and

Vane-Wright 2008).

Within P. demoleus, five subspecies are recognised (Smith and Vane-Wright

2008, Morgun and Wiemers 2012), historically occurring from the Arabian

Peninsula to the southeastern part of Papua New Guinea and Australia

(Bingham 1905, Corbet and Pendlebury 1978, Common and Waterhouse

1972, Morgun and Weimers 2012, Rothschild 1895). These subspecies form

two disjunct groups. In the western group, subspecies P. d. demoleus

occurred from around the Persian Gulf, through India and Sri Lanka,

subtropical China to southern Japan and Taiwan (Smith and Vane-Wright

2008), while subspecies P. d. malayanus Wallace, 1865 (Figs 1-2, 6)

occurred only in southern Myanmar and the Malay Peninsula (Corbet and

Pendlebury 1978). The eastern group contains subspecies P. d. sthenelinus

Rothschild, 1895 (Figs 3-5) from the Lesser Sunda Islands (viz. Alor,

Komodo, Flores, Sumba and Sumbawa) (Rothschild 1895, Tsukada and

Nishiyama 1982, Smith and Vane-Wright 2008), subspecies P. d. sthenelus

Macleay, 1826 (Figs 7-8) from Australia (Braby 2000) and subspecies P. d.

novoguineensis Rothschild, 1908, which is restricted to the area around Port

Moresby in Papua New Guinea (Parsons 1998).

Australian Entomologist, 2017, 44 (2)

66

Australian Entomologist, 2017, 44 (2) 67

The two subspecies that originally occurred from the Arabian Peninsula to

Indo-China and the Malay Peninsula, P. d. demoleus and P. d. malayanus,

predominantly utilise ornamental Citrus and other Rutaceae as larval hosts,

although Smith and Vane-Wright (2008) and van der Poorten and van der

Poorten (2016) reported P. d. demoleus utilising Psoralea and Cullen

corylifolium (L.) Medik (both Fabaceae) in India and northern Sri Lanka

respectively [almost all Psoralea names were synonymised into Cullen as per

Grimes 1997, but some Psoralea spp still occur]. In eastern Papua New

Guinea and Australia, P. d. novoguineensis and P. d. sthenelus, respectively,

predominantly utilise Cullen spp as larval host plants (Parsons 1998, Braby

2000, Morgan and Weimers 2012). The larval host plant of P. d. sthenelinus

from the Lesser Sunda Is is unknown (Smith and Vane-Wright 2008).

Since the late 1960s, P. d. malayanus has undergone a significant range

expansion. Rothschild (1895), Bingham (1905), Corbet and Pendlebury

(1978) and Common and Waterhouse (1972) indicated that P. d. malayanus

was once much more restricted in its range, initially and chiefly confined to

southern Myanmar and the Malay Peninsula, whereas currently its

distribution encompasses the Philippines, Borneo, Indonesia, Timor, New

Guinea and the Solomon Islands, plus Europe and the New World (Smith and

Vane-Wright 2008, Tennent et al. 2011, Morgun and Wiemers 2012,

Fernández Hernández and Minno 2015, Nielsen 2017) and now Australia.

Smith and Vane-Wright (2008) had speculated on the easterly movement of

P. d. malayanus and its subsequent impact on the other P. demoleus taxa,

particularly P. d. sthenelinus in the Lesser Sunda Islands.

Here I report the first record of P. d. malayanus from Australia, on Dauan

Island in Torres Strait, Queensland (a male collected in 2010) and confirm its

establishment by 2016. In addition, I confirm the coexistence of P. d.

sthenelinus with P. d. malayanus in western Flores, Indonesia. Finally,

differences in final instar larval morphology of P. d. demoleus and P. d.

sthenelus are illustrated and discussed.

Figs 1-8. Papilio demoleus subspecies with some diagnostic features arrowed, as per

Smith and Vane-Wright (2008), for male uppersides of P. d. malayanus, P. d.

sthenelinus and P. d. sthenelus. Arrowed features are: A — large yellow spots in pre-

apical area of forewing discal cell; B — forewing medial yellow spots below vein 1A +

2A; C — hindwing medial yellow colour below eye spot; D — hindwing postmedial

lunules, spots or bars. Images: P. d. malayanus from Australia, P. d. sthenelinus after

invasion of P. d. malayanus; and P. d. malayanus from Timor Leste; all figures not to

scale; uppersides left, undersides right [forewing lengths, in mm, in square brackets]:

(1-2) P. d. malayanus, Dauan I., Torres Strait: (1) £, 22.iv.2012 [45 mm]; (2) 9,

24.iv.2012 [42]; (3-5) P. d. sthenelinus, Labuan Bajo, W. Flores: (3) 3, 25.iii.2011

[41]; (4) 9, 25.iii.2011 [45]; (5) 9, 3.iii.2016 [47]; (6) P. d. malayanus, 3, Saburai

District, Timor Leste, 12.xii.2016 [45]; (7-8) P. d. sthenelus: (7) 3, Wards Hill, E. of

Toowoomba, 7.11.1985 [42]; (8) 9, Long Pocket, Indooroopilly, 10.xi.1977 [45].

68 Australian Entomologist, 2017, 44 (2)

Methods and materials

The following acronyms refer to public institutions and private collections

from which material was examined:

CEMC - C.E. Meyer collection, Brisbane; CGMC — C.G. Miller collection,

Lennox Head; MTQ — Museum of Tropical Queensland, Townsville; PRWC

— P.R. Wilson collection, Bundaberg; SSBC — S.S. Brown collection, Bowral;

TLIKC — T.A. Lambkin and A.I. Knight joint collection, Brisbane.

Abbreviations of collectors’ names appearing on specimen labels are:

CEM - C.E. Meyer; CGM - C.G. Miller; IRJ — I. R. Johnson; RPW — R.P.

Weir; PRW - P. R. Wilson; SSB — S.S. Brown; TAL — T.A. Lambkin.

Over the period from 2004 to 2016, I surveyed and sampled butterflies on

Dauan (seven visits) and Mer Islands (three visits), Torres Strait. In addition,

I sampled butterflies at Labuan Bajo, a coastal town situated on the western

tip of Flores, Indonesia, from 2010 until 2016, during which, in 2011 and

2016, I collected specimens of P. demoleus.

Results

Several of the upperside diagnostic features which can be used to identify and

separate males of the three subspecies P. d. malayanus, P. d. sthenelinus and

P. d. sthenelus are arrowed in Figs 1, 3 and 7 respectively, as per Smith and

Vane-Wright (2008).

Papilio demoleus malayanus Wallace, 1865

Specimens examined. QUEENSLAND: TORRES STRAIT: 14 43,6 99:3 33, I

©, Dauan Island, 9°25'S 142°32'E, (3 63) 6.12010, 22.iv.2012, 24.iv.2012, (1 9)

24.iv.2012, TAL (TLIKC); 1 3, same data except: 22-29.12016, CEM, SSB, RPW,

CGM (CEMC), 4 43, 1 9, same data except: (1 3) 21.1.2016, (2 33) 2212016, (1

d) 26.1.2016, (1 9) 16.1.2017, CGM (CGMC); 3 33, 3 99, same data except: (3

SS, 1 9) 222912016, (2 99) 14-20.12017, SSB (SSBC); 2 24, I 9, same data

except: 3-10.iii.2016, IRJ and PRW (MTQ), 1 3, same data except 9°24'41"S

142°32'12"E, WGS 84, 6.11.2016, PRW (PRWC).

INDONESIA: FLORES: 3 33, Labuan Bajo, W. Flores, Lesser Sunda Islands,

8°29'30"S 119°53'1 1 "E, 3.111.2016, 6.111.2016, 5.x1.2016, TAL (TLIKC).

TIMOR LESTE: 1 3, Saburai District, 900 m, 9 km SSW of Maliana, 12.xii.2016,

TAL (TLIKC).

Papilio demoleus sthenelus Macleay, 1826

Specimens examined. MAINLAND QUEENSLAND: 3 £g, 4 99: 2 do, Cecil

Plains, 22.ix.1977, TAL; 1 3, Wards Hill, E. of Toowoomba, 7.ii.1985, TAL; 1 9, Mt

Crosby, 22.1.1976, TAL; 1 9, Long Pocket, Indooroopilly, 10.xi.1977, TAL, from

larva, Psoralea, 2 99, Jamboree Heights, Brisbane, 20.1.1980, 21.xii.1980, TAL,

from larva (all TLIKC).

Australian Entomologist, 2017, 44 Q) 69

Papilio demoleus sthenelinus Rothschild, 1895

Specimens examined. INDONESIA: FLORES: 4 33, 3 99, Labuan Bajo, W. Flores,

Lesser Sunda Islands, 8°29'30"S 119*53'11"E, (3 43, 1 9) 25.iii.2011, (1 3,1 9)

26.iii.2011, (1 9) 3.iii.2016, TAL (TLIKC).

Field observations: Dauan Island, Torres Strait, Queensland

In January 2010 and April 2012, four specimens of P. d. malayanus (1 3 and

2 33, I 9 respectively) (in TLIKC) (Figs 1-2) were collected at the western

end of the Dauan Island village. The butterflies were collected flying swiftly

about a metre above the ground in open areas, a very similar flight pattern to

that of P. d. sthenelus from mainland Australia (Braby 2000). Four years later

the butterfly was observed again in January and March 2016, when 11 33

and 2 99 were collected by CEM, CGM, IRJ, PRW and SSB (CEMC,

CGMC, MTQ, PRWC and SSBC), and then again in January 2017, when 3

OO (CGMC and SSBC) were collected. In January 2016, adult butterflies

were found to be ‘common’ in the village, with at least two females observed

ovipositing on cultivated Citrus sp. in the grounds of the local school (SSB

and CEM pers. comms), while in December 2016, I. Johnson (pers. comm.)

reported that he observed, on average, four specimens per day and saw

females flying around lemon trees.

Sampling of butterflies by the author on Mer Island, in eastern Torres Strait

(170 km south-east of Dauan Island) in January 2011, February 2015 and

January 2016, failed to locate the species.

Field observations: Labuan Bajo, Flores, Indonesia

In March 2011, 4 33 and 2 99 of P. d. sthenelinus were collected (Figs 3-

4), mostly feeding on blossom of Lantana camara L. (Verbenaceae). In

March and November 2016, 2 33 of P. d. malayanus and 1 9 of P. d.

sthenelinus (Fig. 5) were collected feeding on blossom of Echinacea

purpuria (Asteraceae). In 2011, females of P. demoleus taxa were observed

ovipositing on small ornamental Citrus plants less than a metre high; it is not

known which taxa these were as the adults were not collected.

Discussion

Prior to the late 1960s, P. demoleus occurred as two well defined groups

occurring on the two sides of the Malay Archipelago and separated by

approximately 1900 km (Rothschild 1895, Corbet and Pendlebury 1978).

Corbet and Pendlebury (1978) commented on how curious ‘the hiatus in the

distribution of P. demoleus in the Malay Archipelago' was, despite the

existence of Citrus throughout the region. Matsumoto (2002) and Morgun

and Weimers (2012) proposed that the spread of P. d. malayanus across this

hiatus may have been related to deforestation practices and the concurrent

expansion of Citrus, particularly in the Philippine islands, Borneo, Sumatra

and Java.

70 Australian Entomologist, 2017, 44 (2)

Closer to Australia, Moonen (1999) recorded specimens of P. d. malayanus

collected in 1997 near Nabire in Papua Province, eastern Indonesia. Then,

between 2005 and 2010, P. demoleus (more than likely to be P. d.

malayanus, based on its oviposition on Citrus) was recorded from several

Provinces in mainland Papua New Guinea (Western, Morobe and Milne

Bay) plus New Britain and New Ireland (Tennent e£ a/. 2011). More

recently, P. d. malayanus was reported from the Solomon Islands (Nielsen

2017) and Christmas Island (Braby 2016a) In 2016, P. d. malayanus

appeared to be established on Dauan Island, where it utilises Citrus as a

larval host. How far it has spread to other Torres Strait islands is unknown,

but it is likely to be on neighbouring Saibai Island, which is just 7 km away

and where Citrus is common in village gardens.

On the Australian mainland, P. d. sthenelus has a wide distribution and is

known for its migratory behaviour (Smithers and McArtney 1970, Dell 1977,

Braby 2000, Braby 2016b). Its preferred larval hosts appear to be Cullen spp

(Braby 2000), although Rainbow (1907), Waterhouse (1932) and Braby

(2000) reported P. d. sthenelus also ovipositing on or utilising Citrus.

Rainbow (1907) depicted the first illustration of the larva and pupa of P. d.

sthenelus, on Citrus stems. R. Kendall (pers. comm.) reported that in

Brisbane, Queensland, larvae of P. d. sthenelus are very rarely found on

Citrus but, when females of P. d. sthenelus are confined in a large flight cage,

they readily oviposit on fresh Citrus growth and larvae develop normally on

Citrus.

In addition to documenting the first record for Europe, Morgun and Wiemers

(2012) illustrated a world map indicating the areas invaded by P. demoleus.

They also documented the first records of P. d. malayanus from the Lesser

Sunda Islands and Timor, including the last confirmed records of P. d.

sthenelinus in Alor (in 1988), Timor (in 1992) and Komodo (in 1997).

Certainly, the 2016 record of P. d. sthenelinus reported here from Labuan

Bajo indicates that P. d. sthenelinus appears to be still a viable taxon, with no

evidence of it interbreeding with P. d. malayanus. The record of P. d.

sthenelinus occurring on Timor is uncertain as the butterfly was not listed for

Timor by Rothschild (1895), Tsukada and Nishiyama (1982) and Mendes and

Biva de Sousa (2010), although P. d. malayanus is now known from there

(Fig. 6). The record of P. d. sthenelus collected from Sumba in 1978 and

reported and illustrated by Tsukada and Nishiyama (1982) appears to be

misidentified, as their illustrated specimen is P. d. sthenelinus.

The eastern advance of P. d. malayanus has provoked discussion on how

subspecies in its path (i.e. sthenelinus, sthenelus and novoguineensis) could

maintain their viability after the arrival of P. d. malayanus (Smith and Vane-

Wright 2008). Smith and Vane-Wright (2008) noted that they were 'about to

witness’ a potential ‘natural experiment’ when P. d. malayanus spread into

the territories of P. d. sthenelinus and P. d. novoguineensis.

Australian Entomologist, 2017, 44 (2) 71

Smith and Vane-Wright (2008) were intrigued to know if the P. d.

sthenelinus and P. d. novoguineensis lineages could stay intact and coexist

with P. d. malayanus. They noted that if this happened, then it would be

reasonable to conclude that P. d. malayanus, P. d. sthenelinus and P. d.

novoguineensis all have reached separate species status. Now that P. d.

malayanus has been in Flores since 1997 (Morgun and Weimers 2012) and P.

d. sthenelinus still occurs in western Flores (2016), it suggests that these two

taxa have maintained separate lineages and thus might be separate species.

Figs 9-12. Final instar larvae of Papilio demoleus subspp: 9, 11 lateral views; 10, 12

dorsal and dorso-lateral views respectively: (9-10) P. d. demoleus on Citrus, Sri

Lanka; (11-12) P. d. sthenelus on Cullen tenax, Brisbane, Queensland. Figs 9-10

courtesy of G.M. and N.E. van der Poorten, Sri Lanka; Fig. 11 courtesy of Hongming

Kan, Brisbane, Qld; Fig. 12 courtesy of R. Kendall, Brisbane, Qld.

In addition, the molecular data of Smith and Vane-Wright (2008) indicated a

possible separation between the Australian taxon P. d. sthenelus (Figs 7-8)

and P. d. malayanus (Figs 1-2). This separation is supported by several life

history studies, which show that the final instar larvae of P. d. demoleus (Figs

9-10) and P. d. malayanus are variable but essentially the same, but are

notably different from those of P. d. sthenelus (Figs 11-12) (Rainbow 1907,

Tan and Khoon 2012, BOIC [Butterflies and Other Invertebrates Club] 2013,

Fernández Hernández and Minno 2015, van der Poorten and van der Porten

2016). The larva of P. d. sthenelus has two rows of subdorsal blunt spines,

whereas in P. d. malayanus these are absent. The larva of P. d. sthenelus

lacks the diagonal lateral banding on the abdominal segments or banding on

the thoracic segments that P. d. malayanus possesses, and P. d. sthenelus has

a row of lateral red spots just below the row of spiracles, which are absent in

P. d. malayanus. In addition, personal communication with R. Kendall

72 Australian Entomologist, 2017, 44 (2)

indicates that larvae of P. d. sthenelus are morphologically similar when fed

on either Cullen (Fig. 13) or Citrus.

Fig. 13. Cullen tenax, Brisbane, Queensland.

Thus, based on the molecular studies of Smith and Vane-Wright (2008), the

life history data presented above and the biogeographical data presented here,

specifically the coexistence of P. d. sthenelinus and P. d. malayanus, it is

possible that the three P. demoleus taxa (sthenelinus (Figs 3-5), sthenelus

(Figs 7-8) and malayanus (Figs 1-2)) are separate species. To help elucidate

this, a more thorough study and comparison of their biology would be

helpful, especially the morphology and colour pattern of final instar larvae of

P. d. sthenelinus, including its host preference.

Now that P. d. malayanus has reached Torres Strait and is poised to enter

mainland Australia, and based on the data presented above, it is possible that

it will coexist in Australia with P. d. sthenelus. If this happens, then it might

be further evidence for the valid separation of these two taxa and the

recognition of P. d. sthenelus as a distinct species.

In addition, if P. d. malayanus continues its range expansion into mainland

Australia, it presents a potential problem to commercial citrus producers,

since P. d. malayanus 1s known to defoliate young Citrus trees elsewhere

(Morgun and Weimers 2012).

Australian Entomologist, 2017, 44 (2) 73

Acknowledgements

I thank the local community councils and island Elders of Dauan and Mer

Islands, Torres Strait, for allowing entry into their communities and their

assistance during time spent on their islands. Appreciation is given to M.P.

Zalucki (University of Queensland) for his critical review of the manuscript,

and to S.S. Brown, I.R. Johnson, C.E. Meyer, C.G. Miller and P.R. Wilson

for allowing access to specimens in their care and for their personal

communications. I particularly wish to thank J. Nielsen for helpful discussion

and literature. I thank R. Kendall of Brisbane for his personal

communications on P. d. sthenelus utilising Citrus in Australia and for the

use of his image of a final instar larva of P. d. sthenelus from Brisbane. I also

thank Hongming Kan of Brisbane for the use of his image of another final

instar larva of P. d. sthenelus from Brisbane. I am also greatly appreciative of

George and Nancy van der Poorten of Sri Lanka for the use of their images of

final instar larvae of P. d. demoleus from Sri Lanka. In addition, my time

spent in Labuan Bajo, Flores has been aided greatly by the Catholic Brothers

of the Missionaries of the Poor, Labuan Bajo. This paper partially fulfils the

requirements for a Master of Philosophy degree undertaken by the author at

the University of Queensland, Brisbane.

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Australian Entomologist, 2017, 44 (2): 75- 84 75

ADDITIONAL CHARACTERS FOR SEPARATING ADULTS OF

PAPILIO DEMOLEUS STHENELUS W.S. MACLEAY, 1826

(LEPIDOPTERA: PAPILIONIDAE) FROM P. DEMOLEUS L.

SUBSPECIES OF BIOSECURITY CONCERN TO AUSTRALIA

JOHN E. NIELSEN

C /- Pathway Surveillance and Operational Science, Department of Agriculture and Water

Resources, GPO Box 858, Canberra, ACT 2602

Abstract

Papilio demoleus demoleus Linnaeus and P. d. malayanus Wallace are exotic pests of

biosecurity concern to Australia that have recently spread through the Indonesian Archipelago

and New Guinea, with P. d. malayanus recently recorded from Australia in the Torres Strait.

Although these exotic taxa are usually separable from the Australian subspecies (P. d. sthenelus

W.S. Macleay), some specimens of the latter subspecies were found to have characters that may

cause them to be misidentified as an exotic subspecies. Additional characters on the hindwing

underside are demonstrated to have diagnostic utility and are presented as a contribution to the

identification of P. demoleus in Australia for biosecurity purposes.

Introduction

Papilio demoleus Linneaus, 1758 is one of the most widespread members of

the Papilionidae, with five subspecies (Figs 1-12) distributed from the Middle

East through subtropical and tropical Asia south to Papua New Guinea and

Australia (Igarashi 1979, Smith and Vane Wright 2008, Tsukada and

Nishiyama 1982). Common names of this butterfly vary with region, but

include lime or citrus butterfly in Asia and chequered swallowtail in Australia

(Corbet and Pendlebury 1992, Orr and Kitching 2010).

In recent years, P. d. demoleus (Figs 1-2) and P. d. malayanus Wallace, 1865

(Figs 3-4) have expanded their ranges, with P. d. malayanus spreading

throughout the Indonesian Archipelago into Papua New Guinea (Tennent et

al. 2011, Morgun and Wiemers 2012). Papilio d. demoleus has also spread

through Indonesia via the Philippines and has now reached Ceram in the

Moluccas (Fig. 1). These range expansions are apparently due to the clearing

of rainforest for human development and associated plantings of Citrus in the

Philippines and Sumatra, creating suitable habitat for P. d. demoleus and P. d.

malayanus, respectively (Matsumoto 2002). Subsequent dispersal of both

subspecies has been relatively rapid, presumably aided by the strong flight of

adults and possibly by movement of nursery stock within islands. Larsen

(1984) inferred that the spread of P. d. demoleus to the Middle East was

probably also facilitated by plantings of Citrus. Papilio d. malayanus reached

the Bismarck Archipelago by 2005 (Tennent et al. 2011), is recorded from

Christmas Island (Braby 2004) and recently extended its distribution to

Torres Strait, Australia (Lambkin 2017). Elsewhere, butterflies released for

weddings have been implicated in the introduction of P. d. malayanus to the

Caribbean during the early 2000s (Eastwood et al. 2006), while unspecified

trade pathways were believed to have transported a specimen of P. d.

malayanus to Europe (Morgun and Wiemers 2012).

76 Australian Entomologist, 2017, 44 (2)

Figs 1-6. Papilio demoleus subspecies sensu Smith and Vane-Wright (2008) exotic to

Australia and Papua New Guinea: (1-2) P. d. demoleus: (1) $, Ambon Island,

Indonesia, June 2012, local collector vja H. Detani & D. Cassatt [JENC]; (2) 9,

Malalag, Mindanao Island, Philippines, ex pupa 10 May 2013, reared ex ova on Citrus

spp, L.R. & J.P. Ring [JENC]. (3-4): P. d. malayanus: (3) 3, (4) 9, Denpasar, Bali

Island, Indonesia, January 2010, H. Detani via D. Cassatt [JENC]. (5-6): P. d.

sthenelinus: (5) 3, (6) 9 [both ex G.A. Waterhouse collection, AMC].

Australian Entomologist, 2017, 44 (2) 77

Figs 7-12. Papilio demoleus subspecies endemic to Papua New Guinea and Australia

and an example of a mislabelled specimen: (7-8) P. d. novoguineensis: (T) 3, (8) 9,

*mrsby' [Port Moresby], ex W.W. Brandt collection [ANIC]. (9-10) P. d. sthenelus:

(9) 3, (10) 9, Buderim, Queensland, 26.682070°S, 153.085875°E, 10 September

2000, J.E. Nielsen, collected from north-south migratory flight. (11) example of P. d.

sthenelus 3 that superficially resembles an exotic subspecies, Prison farm, Glen

Innes, New South Wales, July 1969-December 1970 [ANIC]. (12) P. d. malayanus 9

misidentified as P. d. sthenelus, bearing data ‘Yeppoon, Queensland, 19 July 1962,

J.C. Le Souef and presumably mislabelled [ANIC].

78 Australian Entomologist, 2017, 44 (2)

Papilio demoleus demoleus and P. d. malayanus are pests of biosecurity

concern for Australia and are targeted through surveys performed by the

Northern Australian Quarantine Strategy (NAQS; Department of Agriculture

and Water Resources, unpublished). Both taxa primarily feed on Citrus and

can be pests of economic importance (CABI 2015, Corbet and Pendlebury

1992). In contrast, P. d. sthenelus W.S. Macleay, 1826 (Figs 9-10) feeds on

Fabaceae (Cullen Medik. and Psoralea L.), with few records from Citrus

(including pers. obs.) and one oviposition record on Melicope J.R.Forst &

G.Forst. (Braby 2000, Straatman 1962, Valentine et al. 1988). Similarly, the

poorly known P. d. novoguineensis Rothschild, 1908 (Figs 7-8) is a Fabaceae

specialist endemic to Papua New Guinea (Fenner and Lindgren 1974). The

life history and biology of a fifth subspecies, P. d. sthenelinus Rothschild,

1895 (Figs 5-6) from the Lesser Sunda Islands, Indonesia, remains unknown

(Matsumoto 2002, Lambkin, 2017).

Smith and Vane-Wright (2008) provided a key using wing pattern characters

to identify P. demoleus subspecies. In that key, P. d. sthenelus 1s separated

from P. d. demoleus and P. d. malayanus by the pale marking on the

forewing discocellular space. In subspecies of P. demoleus that are exotic to

the Australian region, this marking is divided into two spots by an area of

black scaling 70.5 mm wide (hereafter ‘divided discocellular forewing spot’).

In P. d. sthenelus this marking is considered to be not divided (Smith and

Vane-Wright 2008). While examining P. demoleus specimens in the

Australian National Insect Collection (ANIC), several specimens identified as

P. d. sthenelus were found with a divided discocellular forewing spot. These

specimens pre-dated the spread of P. d. demoleus and P. d. malayanus

through Indonesia and were not linked to any import pathway. In comparing

these specimens with P. d. demoleus and P. d. malayanus, it was noted that

several ventral hindwing markings that were not considered by Smith and

Vane-Wright (2008) might have diagnostic utility. This paper examines the

utility of these characters in clarifying the identity of P. demoleus specimens

collected in northern Australia.

Materials and method

The taxonomic arrangement used here follows Smith and Vane-Wright

(2008). Specimens of all subspecies of P. demoleus were examined in the

Australian National Insect Collection (ANIC: 1389 specimens), the

Australian Museum insect collection (AMC) and the author's private

collection (JENC). Additional specimens of P. d. demoleus and P. d.

malayanus were obtained from commercial sources, with the identity of these

specimens confirmed using Smith and Vane-Wright (2008). The hindwing

undersides of available specimens of P. d. sthenelus in the ANIC, and of P. d.

demoleus and P. d. malayanus in the JENC, were photographed using a

Nikon D90 DSLR with either a Nikkor Micro 105 mm handheld or a 40 mm

Nikkor Micro lens and with the camera mounted on a copy stand and

Australian Entomologist, 2017, 44 (2) 79

controlled using Digicam Control software. The digital editing software

GIMP was used to post-process all photos. Two subspecies, P. d.

novoguineensis and P. d. sthenelinus, were not considered for analysis due to

insufficient numbers of specimens being available.

Figs 13-16. Papilio demoleus subspecies, hind wings: (13) characters used and the

measurement for morphometric analysis: bdc: black discocellular crescent; odc: ochre

discocellular crescent; ydm: yellow discocellular marking; measurement of

(bdc--odc): ydm is (white double headed arrow): black double headed arrow. Note that

measurements are taken parallel to an axis formed by vein M2. (14-16) comparison of

ventral hindwing markings of diagnostic value in separating P. d. sthenelus from

subspecies of P. demoleus either endemic to Australia or of biosecurity concern: (14)

P. d. demoleus; (15) P. d. malayanus; (16) P. d. sthenelus. Specimens are 33 in

JENC: see Figs 1, 3 and 9 for collection data.

Measurements were taken of two characters for each specimen using the

‘measure tool” in GIMP (Figs 13-16). The first character was the combined

width of the black marking at the distal apex of the discocellular cell (black

discocellular crescent; bdc) and the ochre-coloured crescent adjacent to the

bdc (ochre discocellular crescent; odc). The width of these markings was

80 Australian Entomologist, 2017, 44 (2)

measured along an axis formed by vein M2, from where M2 intersects with

the discocellular cell. The second character, the length of the pale central

marking of the hindwing ventral discocellular cell (yellow discocelluar

marking; ydm), was also measured at its widest point along the same axis.

Specimens in which any marking being scored was obscured or missing due

to damage or aberration were excluded from analysis. Pairwise comparisons

of this ratio were made between P. demoleus subspecies using two-tailed

Mann-Whitney U tests with a confidence interval of 0.01. It is assumed that

no collector bias exists towards the characters considered here (i.e. the range

of markings exhibited by the specimens examined are representative of P.

demoleus populations generally). The ratio bdc:odc was also considered for

the same taxa using the above method.

Available specimens of P. d. sthenelus in the ANIC were also surveyed to

find specimens that violated the forewing character states used to separate

that subspecies from P. d. demoleus and P. d. malayanus according to

characters used in the key in Smith and Vane-Wright (2008).

Results

The median ratio between the markings (bdc+odc):ydc was found to differ

significantly (p « 0.00001) between all pairwise comparisons of the three

subspecies of P. demoleus examined (Table 1, Fig. 17). The differences were

so marked it was considered unnecessary to perform a protecting multivariate

analysis, especially given only three categories were compared. Across the

subspecies of P. demoleus examined, the width of the markings bdc-*odc

were widest relative to marking ydc in this order: P. d. demoleus > P. d.

malayanus > P. d. sthenelus; noting there was some overlap between P. d.

demoleus and P. d. malayanus. The median ratio of markings bdc:odc was

found to be significantly different between populations but had limited utility

due to overlap between all taxa sampled (data not shown).

A survey of specimens in the series of P. d. sthenelus in the ANIC (173

specimens) found 11 specimens (696 of all specimens examined) with a

divided discocellular forewing spot (Fig. 11) that could have keyed to an

exotic subspecies of P. demoleus using the key in Smith and Vane-Wright

(2008). However, one of these specimens, a female labelled as having been

collected at Yeppoon by J.C. Le Seouf (Fig. 12), was identified as a female P.

d. malayanus based on its (bdc+odc): ydm ratio.

Discussion

Biosecurity programs rely on reliable diagnostic tools being available (SPHD

2015). The variability of P. demoleus, including variation quantified here for

the character state used by Smith and Vane-Wright (2008) to separate P. d.

sthenelus from exotic taxa, makes reliance on a single character impractical

for biosecurity diagnostic purposes. For this reason, it is desirable that

diagnostic tools consider a number of characters for the sake of reliability.

Australian Entomologist, 2017, 44 (2) 81

Table 1. Summary of Mann-Whitney U-test statistics for pairwise comparisons of the

ratio (bdct+odc):ydm measured from sampled specimens of Papilio d. demoleus, P. d.

malayanus and P. d. sthenelus.

Sample Mean of

Taxon Mean Median U Z Significance

size Ranks

P. demoleus sthenelus 0.245

P. demoleus sthenelus

2 -10.8429 p « 0.0001

6.40586

2128 -8.7681

- P. demoleus malayanus

DE

Ratio idoine | yam

DE

p.a

Papa de mote denies Pape demoleus PaO ps Papia demoie us Sthe nøler

Fig. 17. Box and whiskers plot showing the spread of the ratio (bdct+odc):ydm

measured from sampled specimens of Papilio d. demoleus, P. d. malayanus and P. d.

sthenelus.

The morphometric analysis presented here demonstrates that the ratio formed

by the markings (bdc+odc):ydm is reliable for separating P. d. sthenelus from

both P. d. demoleus and P. d. malayanus, including for P. d. sthenelus

specimens that could be confused with exotic subspecies due to variation in

the forewing marking characters used by Smith and Vane-Wright (2008). It 1s

82 Australian Entomologist, 2017, 44 (2)

suggested that the (bdc+odc):ydm ratio presented here be used in addition to

the characters identified in the key by Smith and Vane-Wright (2008), also

illustrated by Lambkin (2017), when identifying specimens of P. demoleus

collected in northern Australia, especially if the specimen is being used to

support biosecurity decision making. In addition, molecular analysis used by

several phylogenetic studies (Eastwood et al. 2006, Zakharov et al. 2004) are

capable of separating the subspecies recognised by Smith and Vane-Wright

(2008). Of these, Eastwood et al. (2006) provided sufficient resolution to be

able to identify the origin of specimens introduced into the Americas.

Consideration should therefore be given to using molecular tests to provide

additional confidence in the identity of P. demoleus collected for biosecurity

purposes.

The P. d. malayanus specimen in ANIC collected by J.C. Le Seouf bearing

label data stating Yeppoon, Queensland as the collecting locality (Fig. 12) is

considered to have been accidentally mislabelled and is not taken to represent

early evidence of P. d. malayanus in Australia. Le Seouf had specimens of

exotic species in his collection, including some Malaysian taxa (M.F. Braby

pers. comm.) and there is evidence that other material he collected was also

incorrectly labelled (Dunn 1985).

Invasive species, including numerous arthropod taxa, pose a serious threat to

Australia. All entomologists, including amateur collectors, are encouraged to

be aware of the contribution they can make to Australian biosecurity by

collecting and reporting specimens of suspected exotic taxa. Plant Health

Australia (2016) provides guidance on how to report suspect exotic plant

pests if they are detected in Australian States or Territories.

Acknowledgements

Jacquie Recsei (Australian Museum, Sydney) and Ted Edwards and You-

Ning Su (Australian National Insect Collection, Canberra) are thanked for

granting access to specimens in their care. Les and Janice Ring generously

bred a series of P. d. demoleus for this work, while Fabian Douglas

contributed additional specimens of P. d. demoleus and, together with

Michael Braby, contributed helpful discussion regarding J.C. Le Seouf's

collection. Further material of P. demoleus was obtained from David Cassatt,

Ronald Hart and Pawel Lewenthal. Ross and Lilac Kendall provided

opportunities to observe larvae of P. d. sthenelus using Citrus spp in

Australia. Stacey Anderson and Luke Halling (NAQS) and Trevor Lambkin

assisted with helpful discussions, while Michael Braby, Julietta Brambilla,

Rod Eastwood, Ted Fenner, Albert Orr and John Tennent provided literature.

Caroline Martin and Gertraud Norton assisted with departmental clearance of

the manuscript, while Albert Orr generously reviewed drafts of the

manuscript and, with Ted Edwards, provided suggestions with statistical

analysis. I am especially grateful to my wife Haliz for supporting my work on

the Lepidoptera.

Australian Entomologist, 2017, 44 (2) 83

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

CSIRO publishing: Collingwood; xxvii + 976 pp.

BRABY, M.F. 2004. The complete field guide to butterflies of Australia. CSIRO: Collingwood;

339 pp.

CABI. 2015. Crop Protection Compendium. Wallingford, UK: CAB International. Available

online (accessed 12 Jan 2015): www.cabi.org/cpc

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

edition. (Edited by J.N. Eliot). Malaysian Nature Society; x + 595 pp.

DUNN, K.L. 1985. Specimens of interest in the J. C. Le Seouf collection of Australian

butterflies. Victorian Naturalist 101: 94-97.

EASTWOOD, R.G., BOYCE, S.L. and FARRELL, B.D. 2006. The provenance of Old World

lime swallowtail butterflies, Papilio demoleus (Lepidoptera: Papilionidae), recently discovered

in the New World. Annals of the Entomological Society of America 99: 164-168.

FENNER, T.L. and LINDGREN, E. 1974. The life history and larval foodplants of Papilio

demoleus L. (Lepidoptera: Papilionidae) in southern New Guinea. Papua New Guinea Science

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IGARASHI, S. 1979. Papilionidae and their early stages. Kondansha, Tokyo.

LAMBKIN, T.A. 2017. Papilio demoleus malayanus Wallace, 1865 (Lepidoptera: Papilionidae)

on Dauan Island, Torres Strait, Queensland and recent confirmation of P. d. sthenelinus

Rothschild, 1895 in the Lesser Sunda Islands. Australian Entomologist 44(2): 65-74.

LARSEN, T.B. 1984. Butterflies of Saudi Arabia and its neighbours. Stacey International,

London; 160 pp.

MATSUMOTO, K. 2002. Papilio demoleus on Borneo and Bali. Journal of the Lepidopterists’

Society 56(2): 108-111.

MORGUN, D.V. and WIEMERS, M. (2012). First record of the lime swallowtail Papilio

demoleus Linneaus, 1758 (Lepidoptera, Papilionidae) in Europe. Journal of Research on the

Lepidoptera 45: 85-89.

ORR, A.G. and KITCHING, R. 2010. The butterflies of Australia. Allen & Unwin: Sydney; 336

Pp.

PLANT HEALTH AUSTRALIA. 2016. Reporting suspect pests (1800 084 881). Available

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plant-pests/reporting-suspect-pests/

SMITH, C.R. and VANE-WRIGHT, R.I. 2008. Classification, nomenclature and identification

of lime swallowtail butterflies: a post-cladistic analysis (Lepidoptera: Papilionidae). Systematics

and Biodiversity 6: 175-203.

SPHD 2015. Operating guidelines for the subcommittee on plant health diagnostics and its

working groups. Available online (accessed 12 Aug 2016): http://plantbiosecuritydiagnostics.

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STRAATMAN, R. 1962. Notes on certain Lepidoptera ovipositing on plants which are toxic to

their larvae. Journal of the Lepidopterists’ Society 16: 99-103.

TENNENT, W.J., DEWHURST, C.F. and MULLER, C.J. 2011. On the recent spread of Papilio

demoleus Linnaeus, 1758, in eastern Papua New Guinea (Lepidoptera, Papilionidae). Butterflies

(Teinopalpus) 58: 30-33.

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1. Papilionidae. Plapac, Tokyo; 460 pp.

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Malanda; 71 pp.

ZAKHAROV, E.V., SMITH, C.R., LEES, D.C., CAMERON, A., VANE-WRIGHT, R.I. and

SPERLING, F.A.H. 2004. Independent gene phylogenies and morphology demonstrate a

Malagasy origin for a wide-ranging group of swallowtail butterflies. Evolution 58(12): 2763-

2782.

Australian Entomologist, 2017, 44 (2): 85-88 85

TRAIN ROBBERY: MENEMERUS BIVITTATUS (DUFOUR, 1831)

(ARANEAE: SALTICIDAE) STEALS LARVAE OF

TECHNOMYRMEX SOPHIAE FOREL, 1902 (HYMENOPTERA:

FORMICIDAE) IN TRANSIT

DANIEL C. HUSTON

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

(Email: Daniel. Huston(a)uqconnect.edu.au)

Abstract

The kleptoparasitic behaviour whereupon a spider steals material being transported by ants,

known as ‘snatching’, is reported from the jumping spider Menemerus bivittatus (Dufour, 1831)

in Queensland, Australia for the first time. Both male and female spiders were observed on

multiple occasions stealing larvae of the tropical pedicel ant Technomyrmex sophiae Forel, 1902

being transported by workers. The present report supports the view that this behaviour is

common across the pantropical range of the spider and likely represents an innate, rather than a

learned, foraging strategy.

Introduction

Salticidae, the jumping spiders, represent the largest family of spiders (WSC

2017), a group known for their well-developed visual system and complex

predatory behaviour (Richman and Jackson 1991, Jackson and Pollard 1996,

Bartos and Szczepko 2012, Bartos and Minias 2016). Many salticids

specialise in the exploitation of ants (e.g. Jackson and Nelson 2012), with

some species choosing to steal what the ants are currently transporting rather

than feeding upon them directly (Jackson er a/. 2008, Cushing 2012, Jackson

and Nelson 2012).

Species of the salticid genus Menemerus Simon, 1868 have been reported to

engage in kleptoparasitic behaviour, termed ‘snatching’, by Jackson et al.

(2008), who noted that this behaviour was originally mentioned by

Bhattacharya (1936) in reference to M. bivittatus (Dufour, 1831) from India.

This snatching behaviour was later well described from observations of three

species of Menemerus, including M. bivittatus, made by Jackson et al. (2008)

in Kenya. More recently, this behaviour was again reported in M. bivittatus in

Brazil (Halfeld 2015). These reports from Africa, India and South America

indicate that this snatching behaviour may be common in M. bivittatus across

its pantropical range. In Australia, M. bivittatus is commonly found in and

around human dwellings (Richardson et al. 2006).

Methods

On several occasions between 2015 and 2017, on the wooden exterior

balcony of the author's residence in Brisbane, Australia, individuals of

M. bivittatus (Fig. 1) were repeatedly observed snatching larvae of

Technomyrmex sophiae Forel, 1902 being transported by workers. Both male

and female M. bivittatus were observed engaging in this behaviour. Each of

these instances was carefully observed and two instances were recorded with

a digital video camera. A shortened video of these observations has been

86 Australian Entomologist, 2017, 44 (2)

made available on YouTube at the following URL: https://youtu.be/

KF9FJLHic20. Ants were initially identified as T. sophiae using the guide to

ants of Brisbane provided by Burwell (2007) and identification was

confirmed using the keys provided by Shattuck (2000) and Bolton (2007).

Spiders were identified using the resources provided by Prószyński (2016)

and ALA (2017).

Fig. 1. Male Menemerus bivittatus observed engaging in kleptoparasitic ‘snatching’

behaviour in Brisbane, Queensland.

Results

The behaviour exhibited by M. bivittatus was much as that described by

Jackson ef al. (2008). In the present study, spiders observed the moving

column of ants from a few centimetres distance until an individual ant

transporting a larva was selected (Figs 2A-B). The spider would then

intercept the moving ant by blocking its path and snatch the lava from the

mandibles of the ant (Figs 2C-F). Upon successful theft of the larva, the

spider would retreat from the ant column in order to feed (Figs 2G-I). Not all

snatching attempts were successful, with about 25% of observed attempts

failing. It is not known 1f these failures occurred due to a mistake on the part

of the spider, an individual ant possessing superior defensive technique, or

some other combination of factors. After feeding, the spider returned to

observing the ant column. When a T. sophiae worker was robbed of the larva

it was transporting, the ant ran in tight circles, interpreted as searching, for

approximately one second before continuing on its original path. On

occasion, the ants would pursue the spider for a short distance, although they

were quickly outmanoeuvred, with the ants returning to their original path.

Australian Entomologist, 2017, 44 (2) 87

Fig. 2. Female Menemerus bivittatus performing ‘snatching’ behaviour. Photograph

series taken from video: (A-C) orientation and identification of target ant; (D-F)

interception of target ant and theft of larva; (G-I) escape and feeding.

Discussion

Combined with the reports of snatching in India (Bhattacharya 1936), Africa

(Jackson et al. 2008) and South America (Hanfeld 2015), the present report

provides additional evidence demonstrating that snatching is likely used

88 Australian Entomologist, 2017, 44 (2)

across the entire pantropical distribution of the spider and is not a tactic

unique to specific populations. Thus, snatching in M. bivittatus appears most

likely an innate strategy, rather than a learned behaviour. To the author's best

knowledge, there has been no previous documentation of any interactions

occurring between M. bivittatus and T. sophiae, the latter an abundant species

of ant in and around Brisbane (Burwell 2000). The exploitation of this ant

species by M. bivittatus is thus likely to be a common occurrence.

References

ALA (Atlas of Living Australia). 2017. Atlas of Living Australia website (Accessed 15 Feb

2017) at: http://bie.ala.org.au/species/Menemerus+bivittatus

BARTOS, M. and MINIAS, P. 2016. Visual cues used in directing predatory strikes by the

jumping spider Yllenus arenarius (Aranea, Salticidae). Animal Behaviour 120: 51-59.

BARTOS, M. and SZCZEPKO, K. 2012. Development of prey-specific predatory behavior in a

jumping spider (Aranea: Salticidae). Journal of Arachnology 40: 228-233.

BHATTACHARYA, G.C. 1936. Observations of some peculiar habits of the spider (Marpissa

melanognathus). Journal of the Bombay Natural History Society 39: 142-144.

BOLTON, B. 2007. Taxonomy of the dolichoderine ant genus Technomyrmex Mayr

(Hymenoptera: Formicidae) based on the worker caste. Contributions of the American

Entomological Institute 35: 1-150.

BURWELL, C. Ants of Brisbane. Queensland Museum, Brisbane; 1+ 68 pp.

CUSHING, P.E. 2012. Spider-ant associations: an updated review of myrmecomorphy,

mymrecophily, and myrmecophagy in spiders. Psyche, 2012: Article ID 151989; 23 pp.

HALFELD, V.R. 2015. Compartamento predatório incomum de Menemerus bivittatus (Dufour)

(Araneae, Salticidae). EntomoBrasilis 8: 162-164.

JACKSON, R.R. and NELSON, XJ. 2012. Specialized exploitation of ants (Hymenoptera:

Formicidae) by spiders (Araneae). Myrmecological News 17: 33-49.

JACKSON, R.R. and POLLARD, S.D. 1996. Predatory behavior of jumping spiders. Annual

Review of Entomology 41: 287-308.

JACKSON, R.R., SALM, K. and POLLARD, S.D. 2008. Snatching prey from the mandibles of

ants, a feeding tactic adopted by East African jumping spiders. Journal of Arachnology 36: 609-

611.

PRÓSZYNSKI, J. 2016. Monograph of Salticidae (Araneae) of the World 1995-2015. Part II.

Global Species Database of Salticidae (Araneae). Online at: www.peckhamia.comv/salticidae/

salticidae.php

RICHARDSON, B.J., ZABKA, M., GRAY, MR. and MILLEDGE, G. 2006. Distributional

patterns of jumping spiders (Araneae: Salticidae) in Australia. Journal of Biogeography 33: 707-

719.

RICHMAN, D.B. and JACKSON, R.R. 1992. A review of the ethology of jumping spiders

(Araneae, Salticidae). Bulletin of the British Arachnological Society 9: 33-37.

SHATTUCK, S.O. 1999. Australian ants: their biology and identification. CSIRO Publishing,

Collingwood, Victoria; xi 226 pp.

WSC (World Spider Catalog). 2017. World spider catalog. Version 18.0. Natural History

Museum, Bern (Accessed 28 April 2017). Available online at: http://wsc.nmbe.ch

Australian Entomologist, 2017, 44 (2): 89-102 89

DESCRIPTION OF ANISYNTA CYNONE ANOMALA SUBSP.N.

(LEPIDOPTERA: HESPERIIDAE) FROM THE NORTHERN

TABLELANDS OF NEW SOUTH WALES, WITH A DISCUSSION OF

ITS VARIATION, SYMPATRY WITH AND SIMILARITY TO

ANISYNTA TILLYARDI WATERHOUSE & LYELL

D.P.A. SANDS! and M.C. SANDS?

!CSIRO Ecosciences Precinct, GPO Box 2583, Brisbane, Qld 4001

^67 Haven Road, Upper Brookfield, Old 4069

Abstract

A new subspecies of hesperiid, Anisynta cynone anomala subsp. n., from Torrington, Bluff Rock

Mountain and Bolivia Hill, Dutchman's Tableland, northern tablelands of New South Wales, is

described and distinguished from Anisynta c. cynone (Hewitson) from South Australia and

Victoria, A. c. gunneda L.E. Couchman from Gunnedah, NSW and the similar A. tillyardi

Waterhouse & Lyell. Both A. c. anomala subsp. n. and A. tillyardi are confirmed to occur at

Bolivia Hill, where some specimens of A. c. anomala subsp. n. resemble those of A. tillyardi,

suggesting possible hybridisation or convergence of the two species at this locality.

Introduction

Anisynta cynone (Hewitson, 1874) is a widely distributed skipper, occurring

patchily in South Australia (Fisher 1978), western Victoria (Dunn and Dunn

1991, Field 2013), southern New South Wales (Braby 2000) and on the

mountains and slopes of the Main Dividing Range in New South Wales

(Common and Waterhouse 1981, Braby 2000). Four subspecies were

recognised by Common and Waterhouse (1981): A. c. cynone and A. c.

gracilis (Tepper, 1882) from South Australia, A. c. grisea Waterhouse, 1932

from Victoria and southwestern New South Wales, and A. c. gunneda L.E.

Couchman, 1954 from Gunnedah and Mt Kaputar, New South Wales. Fisher

(1978), when referring to variation in A. cynone from South Australia,

indicated more material was needed to determine the subspecific status of A.

c. grisea, while Dunn and Dunn (1991) noted that subspecies gracilis, grisea

and cynone could not be separated without reference to the label data.

Subsequently, Braby (2000) did not accept A. c. grisea or A. c. gracilis as

valid subspecies but recognised A. c. gunneda, noting that specimens from

Mt Kaputar, New South Wales, were not typical of that subspecies from

Gunnedah, having a darker brown upperside and variable underside patterns.

Specimens of A. cynone from Bolivia Hill, New South Wales, were noted by

Dunn and Dunn (1991) to be larger than specimens of A. c. gunneda and the

uppersides superficially resembled A. tillyardi Waterhouse & Lyell, 1912,

while Braby (2000) noted that a female specimen of A. cynone from Bolivia

Hill could not be distinguished from that sex of A. tillyardi. Evans (1949)

provided images of male genitalia of Anisynta spp, showing slight differences

between A. cynone and A. tillyardi. However, Braby (2000) considered the

male genitalia of the two species to be indistinguishable. Sands (2009)

considered the wingspans of specimens of A. cynone from Torrington,

Dutchman’s Tableland, to be similar to those of A. tillyardi and suggested

90 Australian Entomologist, 2017, 44 (2)

further taxonomic studies would be required to determine the status of

specimens of A. cynone from Mt Kaputar, Torrington and Bolivia Hill.

Here we assign specimens of A. cynone from Torrington, Bluff Rock

Mountain and Bolivia Hill, on the northern New England Tablelands of New

South Wales, to a new subspecies, A. cynone anomala subsp. n. and

distinguish it from A. c. cynone and A. c. gunneda from Gunnedah, NSW.

We have not reviewed the status of A. cynone from Mt Kaputar, NSW but

show that the forewing lengths of these specimens (Fig. 27) are not

significantly different from those of 4. c. cynone and A. c. gunneda from

Gunnedah, but differ from those of Å. c anomala. We confirm that A. c.

anomala and A. tillyardi both occur at Bolivia Hill (G.R. Forbes pers.

comm.). As proposed by Braby (2000), Sands and New (2002) and Sands

(2009), we consider the following possibilities: (1) hybridisation between A.

c. anomala and A. tillyardi at Bolivia Hill; (2) an established tension zone (cf.

Barton and Hewitt 1985) involving the two species; or (3) convergence in A.

c. anomala that might explain the occurrence of some specimens of A. c.

anomala at Bolivia Hill that were previously considered to be hybrids.

Abbreviations: AMS — Australian Museum, Sydney; ANIC — Australian

National Insect Collection, CSIRO, Canberra; NMV — National Museum of

Victoria, Melbourne; QM — Queensland Museum, Brisbane; SAM — South

Australian Museum, Adelaide; CGM — Grant Miller, private collection; GF —

Graham Forbes, private collection, Brisbane; MCS — Michael Sands, private

collection, Brisbane; FWL — forewing length.

Anisynta cynone anomala subsp. n.

(Figs 1-6, 23-24, 27, 28a-e, h)

Types. Holotype 3, labelled ‘NEW SOUTH WALES, Silent Grove Road, 1128 m, 4.0

km N. Torrington, 29?16'54"S, 151?41'08"E, 14 March 2008, M.C. Sands’, in ANIC.

Paratypes: NEW SOUTH WALES: 12 33, Bolivia Hill, 36 km S Tenterfield,

8.11.1986, J.F.R Kerr; 2 33, 5 99, same data except 9.iii.1986; 9 99, same data

except 8.iii.1984, J. Kerr; 7 9 9, same data except 18.iii.1984; 1 ĝ, 1 9, Bolivia Hill,

36 km S Tenterfield, 16.11.1996, C.G. Miller; 1 9, Silent Grove Road, 1128 m, 4.0

km N Torrington, 29*16'54"S, 151?41'08"E, 12.iii.2008, M.C. Sands; 1 3, New

England Highway, Bolivia Hill, 1215 m, 29?20725"S, 151954"31”E, 1.1v.2006, M.C.

Sands; 1 3, Bolivia Hill, Tenterfield, 3.iii.1984, D.P.A. Sands, in ANIC; 1 3, 19,

29?15'59.]"S, 151?40'35.2"E, 6 km north of Torrington, 22.iii.2009, S.J. Johnson;

2 33, 2 99, Silent Grove Road, 1128 m, 4.0 km N Torrington, 29?16'54"S,

151?41'08"E, 14.iii.2008, M.C. Sands, in QM; 1 ĝ, 1 9, Silent Grove Road, 1128 m,

4.0 km N Torrington, 29?16'54"S, 151?41'08"E, 12.11.2008, M.C. Sands, in AMS;

1 3, Silent Grove Road, 1128 m, 4.0 km N Torrington, 29?16'54"S, 151?41'08"E,

14.iii.2008, M.C. Sands; 1 9, same data except 12.iii.2008, in NMV; I 3, 1 9, Silent

Grove Road, 1128 m, 4.0 km N Torrington, 29?16'54"S, 151°41’08”E, 14.ii1.2008,

M.C. Sands, in SAM; 2 33, 2 99,4 km Torrington, 17.iii.2013, C.G. Miller; 1 3,

Bolivia Hill, 36 km S. Tenterfield, 16.iii.1983, C.G. Miller; 1 3, 1 9, same data

except 23.11.1983; 1 9, same data except 21.11.1986; 1 9, Bolivia Hill, 37 km S

Australian Entomologist, 2017, 44 (2) 9]

Tenterfield, 21.iii.1986, C.G. Miller, in CGM; 2 33,4 99, Silent Grove Road, 1128

m, 4.0 km N Torrington, 29°16’54”S, 151?41'08"E, 12.iii.2008, M.C. Sands; 3 33,38

O9, same data except 14.iii.2008; 1 (4, same data except iv.2006; 24546,1989,

same data except 1.iv.2007; 1 3, Bolivia Hill, 3.iii.1984; 2 33, 2 99, Silent Grove

Road, 1080 m, 4.0 km N Torrington, 29?16'09"S, 151?41'08"E, 24.11.2008, D.P.A..

Sands, in MCS.

5 6

Figs 1-6. A. cynone anomala subsp. n.: (1-2) Holotype 3 (Torrington, NSW); (3-4)

aberrant 3 (Bolivia Hill); (5-6) Paratype 9 (Torrington, NSW). (1, 3, 5) uppersides,

(2, 4, 6) undersides.

Description. Male (Figs 1-4, 23-24). Antennal length (of holotype) 7.0 mm;

shaft dorsally grey-black, segments edged narrowly white, ventral surface

cream; club bowed beyond midpoint viewed dorsally, flattened viewed

laterally, dorsally grey-black, basal half cream viewed ventrally, apical half

92 Australian Entomologist, 2017, 44 (2)

orange; eyes black, edged with white setae dorsally; palpus dorsally black,

surrounded by long black and white setae, ventrally with long white setae;

thorax and abdomen dorsally black with brown setae at posterior, ventrally

light brown with long cream setae on thorax and abdomen terminal segments;

legs light brown with sparse long white setae, mid tibia with two long,

slender apical spurs, hind tibia with three spurs, two apical and one post

median. Forewing length (of holotype) 13.8 mm, costa almost straight,

weakly convex near base and R3 to apex; apex acute; termen weakly convex;

CuA, equidistant from M; and CuA», area between CuA> and 1A+2A, ca 1.5

x wider than CuA, and CuA», 1A+2A upwardly curved towards termen; cilia

light brownish cream, narrowly dark brown at vein ends of R5 to CuA,,

brown between CuA, and 1A+2A and tufted on inner margin. Upperside:

forewing grey-brown, becoming dark brown towards termen, base to cell and

area towards tornus, overlain with pale yellowish-brown scales, one

prominent pale yellow spot towards apex of cell; three short rectangular sub-

apical cream spots, crossed by dark brown veins at right angles to costa

between R4, Rs and Mi, two narrower sub-terminal spots closer to termen

between M, and M3, a row of four post median larger spots, sub-parallel to

termen, between veins M; to 1A+2A, with spots between CuA, and 1A+2A

elongate above 1A42A; hindwing base to 1/3 of costa convex, distal 2/3

almost straight; apex rounded, termen convex, inner margin almost straight,

slightly concave beyond midpoint; hindwing upperside medium grey-brown,

without spots or suffusion, base to median region overlain with pale

yellowish-brown scales; cilia weakly chequered, Sc+R; to Rs grey-brown, Rs

to CuA, cream, broadly brown at vein ends, narrowly cream 1A+2A to

tornus. Forewing underside, costa broadly pale orange-brown from base to

sub-apex, termen at apex broadly pale orange-brown, inner edge crenulated

Rs to M5; three short sub-rectangular sub-apical cream spots between R4 Rs

and M,, aligned at right angles to costa; ground colour posterior to cell, dark

brown from base to sub-apex and inner margin, reaching termen at tornus;

two cream cell spots and two post-median spots between M; and CuA,, CuA,

and CuA,; termen cream, broadly dark brown at vein ends R; to 1A+2A.

Hindwing underside, ground colour orange-brown, a band of post median

cream spots R; to I A+2A, interrupted at veins, inwardly edged with a band of

post median, elliptical reddish-brown spots ringed by brown, R; to 1A+2A;

an elongate greyish-cream patch from base to sub-termen between M, and

Ms, interrupted by two dark cell spots, two short cream patches between

Sc+R;, and Rs, separated by median dark brown spots, another between CuA;

and 1A+2A, both patches at cell sub-triangular; apical half of inner margin

fold 1A+2A to 3A, dark brown crossed by paler veins; marginal cilia Sc+R,

to 1A+2A, broadly cream between veins and at tornus, narrowly brown at

vein ends.

Male genitalia (Figs 28 a-e). Tegumen and vinculum ring wider basally than

dorsally, rounded at saccus; saccus produced; base of tegumen-uncus broad,

Australian Entomologist, 2017, 44 (2) 93

hooded; uncus slightly inwardly curved, expanded posteriorly with two semi-

circular lobes, separated by slightly concave margin, with dense fine setae on

dorsal and lateral surfaces; gnathos with pair of broad sub-triangular ridged

processes, broadest at base with sclerotized spinules; valva with ampulla and

fold separate from harpe, apex of dorsal lobe projecting well beyond ventral

lobe; rounded at apex and heavily developed sclerotized teeth, ventral lobe

with broad basal fold, with separate distal fold curved upwards posteriorly

with apex broad, rounded and edging teeth heavily sclerotized; apex of uncus,

dorsal and ventral lobes with long setae; juxta with heavily sclerotized ventral

plate, each lobe rounded, ventrally developed each side of orifice; aedeagus

sub-tubular, with sheath slightly expanded near orifice.

Female (Figs 5-6, 24). Similar to male, mostly with larger wingspans and

upperside with larger and darker cream forewing spots, hindwing cilia more

prominently chequered. Antennal length (Torrington paratype) 7.2 mm; shaft

dorsally grey-black, segments edged narrowly white, ventral surface cream;

viewed dorsally club bowed beyond midpoint, flattened laterally; dorsally

grey-black, viewed ventrally basal half cream, apical half orange; eyes black,

edged with white setae dorsally; palpus dorsally black, surrounded by long

black and white setae, ventrally with longer white setae; thorax and abdomen

dorsally black with brown setae at posterior, ventrally light brown with long

cream setae on thorax and abdomen terminal segments; legs light brown with

sparse long white setae, mid tibia with two long, slender apical spurs, hind

tibia with three spurs: two apical and one post median. Forewing length

(Torrington paratype) 13.5 mm, base of costa weakly convex, remainder

almost straight; apex obtuse; termen weakly convex. Forewing upperside

grey-brown, grading to dark brown towards termen, base to median area

overlain with pale orange-brown scales, a prominent pale yellow spot near

apex of cell, post-median and sub-terminal area with paler cream-yellow

spots: three short sub-rectangular cream spots aligned and at right angle with

costa between Ry Rs and Mi, a small post-median, pale yellow spot closer to

termen between M; and M»; four post median spots, sub-parallel to termen

M; to IA+2A, most prominent between M; and CuA>; with elongate spot

above 1A+2A smaller; cilia chequered cream apex to IA+2A, brown at vein

ends R; to I A+2A and at tornus; Forewing underside with costa broadly pale

orange-brown from base to apex, termen at apex broadly pale orange-brown,

inner edge crenulated from Rs to M3; 3 short sub-rectangular sub-apical

cream spots between Ry Rs and M,, in a line at right angles to costa; ground

colour posterior to cell dark brown, extending from base to sub-apex and

inner margin, reaching termen at tornus; one large cream cell spot and two

post-median spots between M; and CuA;, and CuA, and CuA»; termen cream,

broadly dark brown at vein ends R; to I A+2A. Hindwing costa (viewed from

beneath) with basal 1/3 convex, distal 2/3 almost straight towards apex, apex

strongly obtuse; termen convex, inner margin almost straight; underside,

ground colour orange brown; sub terminal cream patches between Sc+R;, M3

94 Australian Entomologist, 2017, 44 (2)

and CuA;, CuA, and CuA», and CuA> and 1A+2A; a row of irregular post

median cream-brown spots, edged reddish-brown between cream patches, M;

to 1A+2A; a sub terminal band of post median, oval or elliptical light brown

patches, each ringed by dark brown, edged distally by a sub terminal band of

irregular cream spots between veins Sc+R; and 1A+2A, and distally a sub-

terminal band of brown patches reaching termen, separated narrowly by

cream at veins, Sc+ R; to 1A+2A; apical half of inner marginal fold, 1A+2A

to 3A, dark brown; cilia Sc*R, to 1A+2A moderately chequered, narrowly

brown at vein ends, broadly cream between veins and at tornus.

Female genitalia. (Fig. 28 h). Slide mounted: papillae anales cupped, apically

tapered to blunt tip outwardly clothed in fine setae, apophyses posteriores

long, slender, extending length of segment eight, segment eight with a broad

ventral, bi-lobed sclerite, lobes flattened, slightly concave and similarly

shaped, apically rounded, ostium bursae with post vaginalis rectangular

sclerite with lateral edges rounded, strongly sclerotized, surrounded laterally

by two slender, tapered sclerites; corpus bursae simple, membranous, not

sclerotized.

Diagnosis. Anisynta cynone anomala subsp. n. (Figs 1-6) may be

distinguished from A. c. cynone (Figs 7-10) and A. c. gunneda (Figs 11-14)

by the greater wingspan of the former (for forewing lengths see Fig. 27) and

the colour and pattern of spots on the underside of the hind wings (compare

Figs 2, 4, 6, 23-24 with Figs 8, 10, 12, 14). On the upperside, the ground

colour of most A. cynone subspp, including A. c. anomala, is grey-brown, but

some specimens of A. c. cynone from South Australia are much darker (Fig.

7). The upperside forewing spots of A. c. anomala (Figs 1, 3, 5) are white to

pale cream in males and yellowish in females, whereas those of A. c. cynone

and Å. c. gunneda are almost white or pale cream (Figs 7, 9, 11, 13). On the

underside, the hindwing median spot and spots of the subterminal outer row

are white and well defined in A. c. cynone (Figs 8, 10) but much less well

defined and whitish cream in A. c. gunneda (Figs 12-13). In A. c. anomala the

spots of the postmedian inner row are distinctly elliptical in shape, longer

than wide, brown ringed by dark reddish brown (Figs 2, 6, 23-24); in A. c.

cynone these are orange-brown, tipped by very dark brown towards the

termen (Figs 8, 10); and in A. c. gunneda the spots are only slightly darker

than the ground colour (Figs 12, 14).

Anisynta c. anomala subsp. n. may be distinguished from A. tillyardi (Figs

15-22) by its smaller wingspan (see Fig. 27), differences in ground colour,

the shape of spots on the upperside (compare Figs 1, 3, 5 with Figs 15, 17,

19, 21) and the pattern of spots on the underside of the hind wings (Figs 23-

26). On the upperside, both sexes of A. c. anomala are grey-brown, whereas

the upperside of A. tillyardi is brown-black. A. c. anomala has the basal half

of both wings obscurely overlain with yellowish brown scales, whereas both

wings of A. tillyardi are overlain with orange-brown scales.

Australian Entomologist, 2017, 44 (2) 95

^

Figs 7-14. Anisynta cynone subpp: (7-10) A. cynone cynone (South Australia:

Alexandrina); (11-14) A. cynone gunneda (NSW: Gunnedah); (7-8, 11-12) 33, (9-10,

13-14), 9 9. (7, 9, 11, 13) uppersides, (8, 10, 12, 14) undersides.

96 Australian Entomologist, 2017, 44 (2)

21

Figs 15-22. Anisynta tillyardi: (15-16) 3, (19-20) 9, Mt McKenzie, northern NSW;

(17-18) 3, (21-22) 9, Bolivia Hill, northern NSW. (15, 17, 19, 21) uppersides, (16,

18, 20, 22) undersides.

Australian Entomologist, 2017, 44 Q) 97

The forewing spots of A. c. anomala are cream in males and slightly

yellowish in females. On the underside of the hind wings of both sexes of A.

c. anomala, the postmedian elliptical spots are longer than wide, whereas

those of similarly placed spots in A. tillyardi are neither elliptical nor longer

than wide.

The images of male genitalia figured by Evans (1949) show that the dorsal

lobes of the valvae of A. cynone differ from those of A. tillyardi. We noted

differences between valvae of A. c. cynone (Fig. 28f) and A. tillyardi (Fig.

28g), but the valvae of A. c. anomala (Fig. 28e) differed only slightly, with

the dorsal lobe of A. c. anomala being somewhat narrower and more

produced than the dorsal lobe of A. fillyardi. We could not distinguish

differences in the female genitalia of A. c. cynone and A. tillyardi.

Figs 23-26. Anisynta spp, hindwing undersides: (23-24) A. cynone anomala subsp. n.

(Torrington); (25-26) A. tillyardi (Mt McKenzie). (23, 25) 33, (24, 26) 99. Circle

indicates position of differentiating spots.

Variation. Range of forewing lengths: 33: 12.7-15.0 mm; 99, 13.3-15.8

mm. In addition to differences in forewing lengths, variation in both sexes of

A. c. anomala occurs in the size or presence of forewing spots and hindwing

spots and patches on the underside. The forewing spots on the upperside of

98 Australian Entomologist, 2017, 44 (2)

some males of A. c. anomala subsp. n. from Bolivia Hill and rarely

Torrington, may be small or obscure and closely resemble the pattern and

size of spots of male A. tillyardi (see Diagnosis). On the upperside of the hind

wings, females of 4. c. anomala may sometimes have a central cell spot

(C.G. Miller pers. comm.). The undersides of both sexes of A. c. anomala are

variable but the areas of cream between veins Mi, M» and M; are smaller

than those areas of A. tillyardi (see Diagnosis).

eo

€

O male

© female

MIT 38

>

A.c.cynone Ac.cynone Acgunneda A.c.gunneda A.c.anomala A lillyardi

Sth.Aust Vie. Gunnedah Mt. Kaputar Tor. & Boliv N.NSW

NSW NSW NSW

10 15

Fore wing lengths (mm)

5

Fig. 27. Anisynta cynone subspp and A. tillyardi, forewing lengths (mm): Å and 9

bars with number of specimens examined: A. c. cynone (South Australia: Alexandrina,

Malinong, Ashville); A. c. cynone (Victoria: Kerang); A. c. gunneda (inland NSW:

Gunnedah); A. c. gunneda (northern NSW: Mt Kaputar); A. c. anomala subsp. n.

(northern NSW: Torrington, Bolivia Hill); A. tillyardi (northern NSW: Ebor, Mt

McKenzie, Liston, Killarney).

Forewing lengths (mm) of a number of males and females of A. cynone

subspecies and A. tillyardi (Fig. 27) were analysed. Two-way analysis of

Australian Entomologist, 2017, 44 (2) 99

variance of forewing lengths, using factors sexes and taxa, found both were

significant (sex, F) 153 = 33.60, p< 0.001; taxa, Fsisg = 135.86, p«0.001;

without interaction, Fs 158 = 0.88 NS). Hence, there was a constant difference

between sexes for all taxa, with female forewings longer than males,

estimated at 0.82 + se 0.16. The LSD test identified three groups of forewing

lengths for: (1) A. c. cynone (South Australia, 12.15 mm + se 0.13; Victoria,

12.00 mm + se 0.23), A. c. gunneda (NSW: Gunnedah, 11.86 mm + 0.15 and

Mt Kaputar, 12.30 mm + se 0.31), with forewing lengths not significantly

different; (2) A. c. anomala (NSW: Torrington, Bolivia Hill, 13.90 mm + se

0.12), with forewings significantly longer than in A. c. cynone and A. c.

gunneda; and (3) A. tillyardi, with forewings significantly longer than in A. c.

anomala.

N på " 1

SD Ne. — DES

puse al à "wd DD b J

Urr. í l ^ x |

- AL MT p d

a - J re R? På

C

d

Y Ta

= TA:

== v

—À E oi "T LN / 1

€ EFN A

: P(X

— ( z. £ Pd ow

ib V ý

As wy)

f Ne aes / Re | \ \

» 2i i )

j ~ n^ |

på På /

FS "Tv i 7 å

y m. rd 4 SN i t / pe

— <= 1 l APT

| x | a h

g —

Fig. 28. Anisynta spp genitalia: (a-e, h) A. cynone anomala subsp. n. (Torrington); (f)

A. c. cynone (South Australia); (g) A. tillyardi. (a-g) 33: (a) undissected, lateral view;

(b-c) aedeagus retracted, juxta at apex, (b) lateral view, (c) dorsal view; (d) uncus and

tegumen, dorsal view; (e-g) valvae; (h) 9: sclerites with terminalia, ventral view.

Scale bar = 1 mm.

Distribution. New South Wales: Torrington, Dutchman’s Tableland (Sands

2009), Bluff Rock Mountain (one specimen, J. Moss) and Bolivia Hill, South

of Tenterfield.

100 Australian Entomologist, 2017, 44 (2)

Biology. At Mt Kaputar, inland NSW, larvae of A. c. gunneda are known to

feed on Austrostypa scabra (Lindl.) S.W.L. Jacobs & J. Everett and Poa aff.

sieberiana Spreng. (R. Mayo pers comm., Braby 2000). At Bolivia Hill,

females of A. c. anomala have been observed ovipositing on a Poa sp. (C.G.

Miller pers. com.), possibly P. sieberiana, but other possible food plants

include Austrostipa scabra, A. rudis subsp. rudis (Spreng.) S.W.L. Jacobs &

J. Everett, A. rudis subsp. nervosa (Vickery) S.W.L. Jacobs & J. Everett, A.

aristiglumis (F. Muell.), S. W.L. Jacobs & J. Everett, and A. ramosissima

(Trin.) S.W.L. Jacobs & J. Everett; (P. Grimshaw pers. comm. ).

At Torrington, Poa sieberiana 1s abundant (P. Grimshaw pers. comm.), and

likely to be a food plant for A. c. anomala. Other possible food plants for A.

c. anomala at Torrington include Austrostipa scabra, A. rudis subsp. rudis, A.

rudis subsp. nervosa and A. verticillata (Nees ex Spreng.) S.W.L. Jacobs & J.

Everett (P. Grimshaw pers. comm.). Poa labillardierei is known to be a food

plant for A. tillyardi (Atkins 1975) and is likely to be its food plant at Liston,

Queensland, where A. tillyardi is very abundant. However, at Mt McKenzie

where A. tillyardi is also abundant, P. sieberiana 1s the most common species

of Poa (P. Grimshaw pers. comm.) and at Bolivia Hill, where A. tillyardi is

not as abundant as A. c. anomala, a food plant of A. tillyardi might also be P.

sieberiana.

The favoured habitats at both localities are undisturbed areas in eucalypt

woodlands, where both sexes have been observed flying over patches of Poa

spp. At the roadside at Bolivia Hill, adults of 4. c. anomala have been

observed visiting yellow daisies (J.F.R. Kerr pers. comm.), including

Xerochrysum bracteatum (Vent.) Tzvelev, Chrysocephalum apetulatum

(Labill.) Steetz. and at least one introduced species.

Discussion

The extent of occurrence of A. c. anomala is not known but the subspecies is

thought likely to occur more extensively on the northwestern slopes near

Tenterfield, New South Wales. With the possibility of a cline between

populations of A. c. anomala and A. c. gunneda, the habitat, food plants and

current lack of life history data have influenced our decision to describe this

taxon as a subspecies of A. cynone and not recognise it as a distinct species.

We have also taken into consideration the views of Braby ef al. (2012)

relating to the usefulness of the subspecies concept. Whereas adults of A. c.

anomala prefer the dryer eucalypt woodlands on the western side of the Main

Range and tablelands, A. tillyardi prefers moist woodlands on the eastern

parts of the Main Range, except for the population occurring in moist

woodland at Mt McKenzie at the western edge of its range. Whereas A.

tillyardi occurs abundantly at Mt Mackenzie, it has not been found at

Torrington, a western locality for A. c. anomala. Few specimens of A.

tillyardi are known from Bolivia Hill but they include both sexes taken with

A. c. anomala on the same day at the same locality (G. Forbes pers. comm. ).

Australian Entomologist, 2017, 44 (2) 101

However, there may be some seasonal differences in times of appearance of

adults of the two species at nearby localities: for example, A. c. anomala

occurs mainly from mid March to early April at Bolivia Hill and at

Torrington, whereas A. tillyardi is most abundant from mid February to early

March, for example at Mount McKenzie (ca 25 km S of Bolivia Hill), north

of Tenterfield and at several locations in northern New South Wales,

including Liston.

Most specimens from Bolivia Hill previously thought to have been hybrids

are males and have the forewing spots reduced in number, obscured by dark

scales (e.g. Figs 3-4) or with some forewing spots absent. Very few males of

A. c. anomala have the elliptical underside spots reduced but some, with

undersides otherwise typical of A. c. anomala, have the forewing upperside

darker than usual and resemble A. tillyardi. Based on the recent records of A.

tillyardi from Bolivia Hill and specimens identical to those from Mt

McKenzie, we consider that the female specimen mentioned by Braby (2000)

might have been A. tillyardi.

The conservation significance of the population of A. c. anomala at Bolivia

Hill was noted by Sands and New (2002) but it is doubtful that threats can be

identified for either this area or the areas at Torrington, both now recognised

as National (State owned) Parks. The similarity between some specimens of

A. c. anomala and A. tillyardi at Bolivia Hill might be an example of

convergence rather than hybrdisation. However, hybridisation is known in

other closely related species of insects: for example, hybrids between

tenebrionid beetles from Namibia occurred in areas where their habitats

overlapped (Hamilton and Penrith 1977) and where hybridising continued

despite maintenance of their specific integrity in nearby habitats. If

hybridisation is occurring between A. c. anomala and A. tillyardi it might be

limited to an area of overlap near Bolivia Hill, where A. c. anomala and A.

tillyardi occur syntopically. We consider the population at Bolivia Hill might

represent a tension zone (cf. Barton and Hewitt 1985), where both species

occasionally hybridise but partially different seasonal emergence of adults

has maintained the integrity of both species, despite some gene flow. This

possibility might be resolved by future DNA studies in a way similar to those

of Rougerie et al. (2012), with their studies on hybridisation of hawk moths

in Tahiti. If hybridisation is shown to occur, the population of A. c. anomala

at Bolivia Hill might represent a Tension Zone involving overlap in the

distributions of A. c. anomala and A. tillyardi, as described by White et al.

(1969) for grasshoppers on Kangaroo Island in South Australia.

Acknowledgements

We thank Alex Stolarski, Graham Forbes, Russell Mayo and Grant Miller for

access to their specimens and Michael Braby, Ted Edwards, Ross Field, Tim

New, Ed Petrie and Tony Moore for their helpful discussions. Thanks also to

Max Whitten and Andreas Zwick for discussions on hybrid zones and the

102 Australian Entomologist, 2017, 44 (2)

potentials of future molecular studies and special thanks to John Kerr for his

constructive comments on an earlier draft of this paper. We also thank Paul

Grimshaw for plant identifications and Anne Bourne and Gunter Maywald

for statistical analyses and the composition of the figure on forewing lengths.

References

BARTON, N.H. and HEWITT, G.M.1985. Analysis of hybrid zones. Annual Review of Ecology

and Systematics 16: 113-148.

BRABY, M.F. 2000. Butterflies of Australia. Their identification, biology and distribution, 2.

vols. CSIRO Publishing, Melbourne; xxvii 976 pp.

BRABY, M.F. 2010. The merging of taxonomy and conservation biology: a synthesis of

Australian butterfly systematics (Lepidoptera: Hesperioidea and Papilionidae) for the 21*

century. Zootaxa 2707: 1-76.

BRABY, M.F, EASTWOOD, R. and MURRAY, N. 2012. The subspecies concept in

butterflies: has its application in taxonomy and conservation biology outlived its usefulness?

[Review article]. Biological Journal of the Linnean Society 106: 699-716.

COMMON, LF.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Revised edition.

Angus and Robertson, Sydney; xii + 498 pp; xiv + 612 pp.

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

and taxonomy. Part 2. Family Hesperiidae. Power Press, Bayswater, Victoria.

EVANS, W.H.1949. A catalogue of the Hesperiidae from Europe, Asia and Australia in the

British Museum (Natural History). British Museum (Natural History), London; xix + 502 pp.

FIELD, R.P. 2013. Butterflies. Identification and life history. Museum Victoria, Melbourne.

FISHER, R.H. 1978. Butterflies of South Australia. Woolman, South Australia.

HAMILTON, W.J. and PENRITH, M.L. 1977. Description of an individual possible hybrid

tenebrionid beetle and the habitat preference of the parental species. Canadian Entomologist:

109: 701-710.

ROUGERIE, R., HAXAIRE, J., KITCHING, I.J. and HEBERT, P.D.N. 2012. DNA barcodes

and morphology reveal a hybrid hawkmoth in Tahiti (Lepidoptera: Sphingidae). /nvertebrate

Systematics 26: 445-450.

SANDS, M.C. 2009. A new geographical record for Anisynta cynone (Hewitson) (Lepidoptera:

Hesperiidae: Trapezitinae). Australian Entomologist 36(1): 3-5.

SANDS, D.P.A. and NEW, T.R. 2002. The action plan for Australian butterflies. Environment

Australia, Canberra; 378 pp and CD.

WHITE, M J.D., KEY, K.H.L, ANDRE, L and CHENEY, J. 1969. Cytogenetics of the viatica

group of morabine grasshoppers II. Kangaroo Island populations. Australian Journal of Zoology

17(2): 313-328.

Australian Entomologist, 2017, 44 (2): 103-104 103

CONTRIBUTION TO THE GENUS RANOLUS BLAIR

(COLEOPTERA: DERMESTIDAE: RANOLINI), WITH NEW

SYNONYMY AND COMBINATIONS

JIRÍ HAVA! and JOHN LAWRENCE?

‘Department of Forest Protection and Entomology, Faculty of Forestry and Wood Sciences,

Czech University of Life Sciences, Kamycká 1176, CZ-165 21, Prague 6, Czech Republic

(Email: jh.dermestidae(a)volny.cz)

? CSIRO Australian National Insect Collection, GPO Box 1700, Canberra, ACT 2601 and

61 Glenbar Road, The Palms, Old 4570 (Corresponding author)

Abstract

The following new synonymy and combinations are proposed: Ranolus Blair, 1929 — Orphilodes

Lawrence & Slipinski, 2005, syn. n.; Ranolus australis (Lawrence & Slipinski, 2005), comb. n.;

Ranolus malleecola (Lawrence & Slipifiski, 2005), comb. n.; Ranolus minor (Lawrence &

Slipinski, 2005), comb. n. and Ranolus papuanus (Hava, 2015), comb. n.

Introduction

The genus Ranolus Blair, 1929 was originally described as a subgenus of

Attagenus Latreille, 1802, with the sole species A. (Ranolus) cavernicola

Blair, 1929 from Selangor, Malaysia. Háva and Kalík (2005) redescribed,

illustrated and compared it with all other known genera belonging to the tribe

Attagenini and considered it to be a separate genus. Háva, in Zahradník and

Hava (2014) transferred it, along with Orphilodes Lawrence & Slipinski,

2005, to the new tribe Ranolini of the subfamily Orphilinae. A second species

of Ranolus from Thailand was described by Háva (2014). Orphilodes

currently includes three Australian species (Lawrence and Slipinski 2005)

and one from Papua New Guinea (Háva 2015).

Subfamily Orphilinae LeConte, 1861

Tribe Ranolini Háva, 2014

Genus Ranolus Blair, 1929

Attagenus (Ranolus) Blair, 1929: 382. Type species: Attagenus (Ranolus) cavernicola

Blair, 1929: 382.

Orphilodes Lawrence & Slipinski, 2005: 236, syn. n. Type species: Orphilodes

australis Lawrence & Slipinski, 2005: 240.

Remarks. When Orphilodes was described by Lawrence and Slipinski in

2005, no comparison with Ranolus was made because Ranolus was then

considered to be a subgenus of the more distantly related genus Attagenus.

According to a recent study of both Ranolus and Orphilodes by the senior

author, including an examination of the type species of the former, it appears

that all included species of both nominal genera belong to one genus, the

oldest name for which is Ranolus. All morphological characters (pronotal

depressions, antennae, wings, male genitalia, legs, etc.) are identical. It may

be noted here that in the original paper describing Orphilodes on the basis of

adult species from Australia (Lawrence and Slipinski 2005), the distinctive

larva of the genus was also recorded from Sabah, close to the then known

104 Australian Entomologist, 2017, 44 (2)

range of Ranolus in Malaysia. The adult of Ranolus australis (Lawrence &

Slipinski) and a larva of an Australian Ranolus sp. are illustrated in Figs 1-2

and the following species are newly transferred:

Ranolus australis (Lawrence & Slipinski, 2005) (Orphilodes), comb. n.

Ranolus malleecola (Lawrence & Slipinski, 2005) (Orphilodes), comb. n.

Ranolus minor (Lawrence & Slipifiski, 2005) (Orphilodes), comb. n.

Ranolus papuanus (Håva, 2015) (Orphilodes), comb. n.

1

2

Figs 1-2. Ranolus spp: (1) R. australis adult, dorsal; (2) Ranolus sp. larva, lateral.

Acknowledgements

We are grateful to Sharon Shute (Natural History Museum, London) for the

loan of the holotype of Ranolus cavernicola. Dr. Adam Slipinski, Australian

National Insect Collection, provided the adult and larval figures.

References

BLAIR, K.G. 1929. Fauna of the Batu Caves, Selangor. XVII. Coleoptera. Journal of Federal

Malay Museum, Singapore 14: 381-387.

HÁVA, J. 2014. A new species of the genus Ranolus (Blair, 1929) from Thailand (Coleoptera:

Dermestidae: Orphilinae: Ranolini). Studies and Reports, Taxonomical Series 10(2): 403-408.

HÁVA, J. 2015. A new Orphilodes Lawrence & Slipinski, 2005 from Papua New Guinea

(Coleoptera: Dermestidae: Orphilinae). Arquivos Entomolóxicos 14: 222-226.

HÁVA, J. and KALÍK, V. 2005. A contribution to knowledge of the tribe Attagenini

(Coleoptera: Dermestidae: Megatominae) from the Malayan subregion. Acta Musei Moraviae,

Scientiae Biologicae, Brno 90: 209-214.

LAWRENCE, J.F. and SLIPINSKI, A. 2005. Three new genera of Indo-Australian Dermestidae

(Coleoptera) and their phylogenetic significance. Invertebrate Systematics 19: 231-261.

ZAHRADNIK, P. and HÁVA, J. 2014. Catalogue of the world genera and subgenera of the

superfamilies Derodontoidea and Bostrichoidea (Coleoptera: Derodontiformia, Bostrichiformia).

Zootaxa 3754: 301-352.

Australian Entomologist, 2017, 44 (2): 105-112 105

A REVIEW OF THE SUBGENUS JA VADACUS HARDY OF

BACTROCERA MACQUART (DIPTERA: TEPHRITIDAE: DACINAE)

D.L. HANCOCK! and R.A.I. DREW?

18/3 McPherson Close, Edge Hill, Cairns, Old 4870

"International Centre for the Management of Pest Fruit Flies, Griffith University, Old 4111

Abstract

The Bactrocera Macquart subgenus Javadacus Hardy is reviewed and five SE Asian species

recognised. Four additional species from SE Asia and Australia, previously included in

Javadacus, are transferred to subgenus Bactrocera, with B. unirufa Drew placed as a new

synonym of B. melanothoracica Drew. Based on its type species, subgenus Javadacus is

included in the Zeugodacus groip of subgenera. A key to species is included.

Introduction

The Bactrocera Macquart subgenus Javadacus Hardy was established for the

West Javanese species Bactrocera montana (Hardy) (Hardy 1983). Drew and

Romig (2013, 2016) included seven Southeast Asian species, two of which,

together with three Australian species (Drew 1989), are transferred here to

subgenus Bactrocera, based on their male sternite V and surstylus characters.

Based on the type species and others here included, the shallow posterior

emargination to sternite V and the elongate posterior surstylus lobes in males,

plus its overall appearance, place subgenus Javadacus in the Zeugodacus

group of subgenera as defined by Drew (1989). Virgilio ef al. (2015) and De

Meyer et al. (2015) referred it to the Bactrocera group of subgenera on the

basis of ‘B. unirufa Drew’, a species now known to actually belong in

subgenus Bactrocera (see Hancock and Drew 2015) and here placed in

synonymy with B. melanothoracica Drew. Accordingly, Javadacus species

were not transferred by De Meyer et al. (2015) to their ‘genus’ Zeugodacus

Hendel and they are also retained here within genus Bactrocera, following

Drew and Romig (2013) and Hancock and Drew (2015) in this regard.

Genus Bactrocera Macquart

Subgenus Javadacus Hardy

Dacus (Javadacus) Hardy, 1983: 26. Type species Dacus (Javadacus) montanus

Hardy, 1983, by original designation.

Definition. Abdominal sternite V of male with a shallow posterior

emargination; posterior lobe of male surstylus elongate and narrow; pecten of

cilia present on abdominal tergite III of male; postpronotal setae absent;

supra-alar setae absent; prescutellar acrostichal setae present; one pair of

scutellar setae; scutum with medial postsutural yellow vittae present and

lateral postsutural yellow vittae extending anterior to suture as distinct spots.

Response to male lures. Cue lure (4 species) or none known (1 species).

106 Australian Entomologist, 2017, 44 (2)

Included species. Bactrocera (J.) javanensis (Perkins) (= transtillum Hering),

B. (J.) montana (Hardy), B. (J.) scutellaria (Bezzi), B. (J.) semisurstyli Drew

& Romig, B. (J.) trilineata (Hardy).

Host plants. One species recorded from Coccinia grandis (Cucurbitaceae);

others unknown (Drew and Romig 2013).

Comments. This subgenus appears to be closely related to the large subgenus

Zeugodacus, differing in having both supra-alar and basal scutellar setae

absent.

B. (Javadacus) javanensis (Perkins)

Afrodacus javanensis Perkins, 1938: 132. Type locality Mt Ardjoeno, E. Java.

Strumeta transtillum Hering, 1952: 265. Type locality Idjen, Ongop-ongop, E. Java.

Syn. Drew and Romig, 2013: 208.

Dacus (Afrodacus) javanensis (Perkins): Hardy 1955: 9.

Bactrocera (Javadacus) javanensis (Perkins): Norrbom et al. 1999: 98; Drew and

Romig 2013: 208.

Distribution. East Java, Indonesia. Recorded from Mts Arjuno and Ijen.

Host plant. None known.

Male lure. None known.

Comments. Illustrated by Drew and Romig (2013).

B. (Javadacus) montana (Hardy)

Dacus (Javadacus) montanus Hardy, 1983: 27. Type locality Cibodas, W. Java.

Bactrocera (Javadacus) montana (Hardy): Norrbom et al. 1999: 98; Drew and Romig

2013: 208.

Distribution. West Java, Indonesia. Recorded from Mt Gede.

Host plant. None known.

Male lure. Cue lure; the record from methyl eugenol (Hardy 1983) appears to

be either an error or the result of contaminated lures.

Comments. Illustrated by Drew and Romig (2013).

B. (Javadacus) scutellaria (Bezzi)

Chaetodacus scutellarius Bezzi, 1916: 110. Type locality Goorghalli Estate, S.

Mysore, India.

Dacus (Bactrocera) scutellarius (Bezzi): Hardy 1977: 52.

Bactrocera (Bactrocera) scutellaria (Bezzi): Norrbom et al. 1999: 95.

Bactrocera (Javadacus) scutellaria (Bezzi): Drew and Raghu 2002: 328; Drew and

Romig 2013: 211.

Distribution. Southern India.

Australian Entomologist, 2017, 44 Q) 107

Host plant. None known.

Male lure. Cue lure.

Comments. Illustrated by Drew and Romig (2013).

B. (Javadacus) semisurstyli Drew & Romig

Bactrocera (Javadacus) semisurstyli Drew and Romig 2013: 212. Type locality Suli,

Ambon, Indonesia.

Distribution. Ambon I., Maluku, Indonesia.

Host plant. None known.

Male lure. Cue lure.

Comments. This species has a fulvous face with a pair of oval fuscous spots

and entirely red-brown femora. Illustrated by Drew and Romig (2013).

B. (Javadacus) trilineata (Hardy)

Dacus (Afrodacus) trilineatus Hardy, 1955: 12. Type locality Sarakki Village,

Bangalore, India.

Bactrocera (Javadacus) trilineata (Hardy): Norrbom et al. 1999: 98; Drew and Raghu

2002: 328; Drew and Romig 2013: 213.

Distribution. India, Sri Lanka, Thailand and Vietnam.

Host plant. Coccinia grandis (Cucurbitaceae) (Drew and Romig 2013).

Male lure. Cue lure.

Comments. Illustrated by Drew and Romig (2013). Specimens from Thailand

and Vietnam have a smaller black area on the notopleural lobe, broader

postsutural lateral yellow vittae and entirely fulvous hind femora; these are

currently regarded as conspecific (Drew and Romig 2013).

Excluded species

The following two Asian and two Australian species currently included in

subgenus Javadacus differ in lacking a medial postsutural yellow vitta and

the presutural extensions of the lateral postsutural vittae on the scutum. They

also have a deeply emarginate posterior margin to male sternite V and

produced but still relatively short posterior surstylus lobes (Figs 1-2). This,

plus their non-cucurbitaceous host plants, better places them in subgenus

Bactrocera, to which they are referred. The slightly produced posterior

surstylus lobe seen in these species also occurs in B. (Queenslandacus)

exigua (May) [HT examined, in Queensland Museum, Brisbane] but the short

cell bcu extension seen in the latter species suggests that it 1s not closely

related. In all known cases, males respond to methyl eugenol or isoeugenol,

chemically similar attractants that are rarely found outside the Bactrocera

group of subgenera.

108 Australian Entomologist, 2017, 44 (2)

B. (Bactrocera) aberrans (Hardy)

Dacus (Afrodacus) aberrans Hardy, 1951: 118. Type locality Lake Barrine,

Queensland, Australia.

Afrodacus mesoniger May, 1952: 8. Type locality Toowoomba, Queensland,

Australia. Syn Drew 1989: 189.

Bactrocera (Javadacus) aberrans (Hardy): Drew 1989: 189.

Distribution. Known with certainty only from SE Queensland (Toowoomba,

Mt Tamborine and Ashgrove) and the Atherton Tableland in NE Queensland.

Host plants. Cinnamomum oliveri, C. virens, Litsea leefeana, L. reticulata

and Neolitsea dealbata (Lauraceae) (Hancock et al. 2000).

Male lure. Weakly attracted to isoeugenol (Royer 2015). Reports from cue

lure (Huxham and Hancock 2002) are misidentifications.

Comments. Identification of this species is difficult and some previous

records (e.g. Huxham and Hancock 2002) refer to aberrant individuals (with

supra-alar setae lacking) of other Bactrocera species. Two males from

Cooktown with a relatively narrow anepisternal stripe, recorded by Royer and

Hancock (2012) as possibly B. unirufa, are now believed to belong to B.

aberrans. Illustrated by Drew (1989).

B. (Bactrocera) melanothoracica Drew

Bactrocera (Javadacus) melanothoracica Drew, 1989: 190. Type locality Yam I.,

Torres Strait, Australia.

Bactrocera (Javadacus) unirufa Drew, 1989: 191. Type locality Bellenden Ker

Range, Queensland, Australia; syn. n.

Bactrocera (Bactrocera) melanothoracica Drew: Hancock and Drew 2015: 101.

Bactrocera (Bactrocera) unirufa Drew: Hancock and Drew 2015: 101.

Distribution. Torres Strait islands and Cape York to Kennedy (near Cardwell)

in NE Queensland (Royer and Hancock 2012).

Host plant. Not recorded, but possibly Tabernaemontana pandacaqui

(Apocynaceae) (Royer and Hancock 2012).

Male lure. Methyl eugenol.

Comments. This species is very variable in the extent of the black areas on

the scutum but the area lateral of the postsutural lateral yellow vitta is always

red-brown. Prescutellar acrostichal setae are present and supra-alar setae are

normally absent. Male sternite V and surstylus lobes are as in B. (B.)

pallescentis (Hardy) (cf. Figs 1-2). Examination of holotypes of both B.

melanothoracica and B. unirufa |in Queensland Museum, Brisbane], plus a

large series of specimens from throughout the species’ range, indicates that

only one taxon is involved, with B. unirufa here placed in synonymy. Both

taxa were illustrated by Drew (1989).

Australian Entomologist, 2017, 44 (2) 109

B. (Bactrocera) nigrita (Hardy)

Dacus (Afrodacus) aberrans nigritus Hardy, 1955: 5. Type locality Singapore.

Bactrocera (Javadacus) nigrita (Hardy): Norrbom et al. 1999: 98; Drew and Romig

2013: 209.

Distribution. Recorded from Singapore and Vietnam.

Host plant. Cinnamomum iners (Lauraceae) (Hardy 1955).

Male lure. A single specimen has been collected at methyl eugenol (Drew

and Romig 2013).

Comments. Illustrated by Drew and Romig (2013).

B. (Bactrocera) pallescentis (Hardy)

Dacus (Afrodacus) aberrans pallescentis Hardy, 1955: 5. Type locality Ranikhet,

Uttar Pradesh, India.

Bactrocera (Javadacus) pallescentis (Hardy): Norrbom et al. 1999: 98; Drew and

Romig 2013: 210.

Distribution. Northern India (Uttar Pradesh).

Host plant. Mimusops sp. (Sapotaceae) (Hardy 1955).

Male lure. None known.

Comments. The male sternite V and surstylus lobes are shown in Figs 1-2;

they are from one of five paratype males now in the Natural History

Museum, London. This species was illustrated by Drew and Romig (2013).

1 2

Figs 1-2. Bactrocera (Bactrocera) pallescentis (Hardy), paratype male: (1) sternite V;

(2) surstylus lobes.

Key to species of subgenus Javadacus

I Scutum red-brown with a pair of broad, black submedial bands and a

yellow lateral band between postpronotal and mnotopleural lobes;

postsutural lateral yellow vittae short and triangular, ending well before

postalar setae; wing with costal band crossing vein R»+3 and broad across

apex of cell r4+5, ending at apex of vein M; abdominal tergite III orange-

110 Australian Entomologist, 2017, 44 (2)

brown with black lateral margins [eastern Indonesia (Ambon)] ...............

EA UPHRO AR B. (J.) semisurstyli Drew & Romig, 2013

— Scutum mostly black with no lateral yellow band between postpronotal

and notopleural lobes; lateral postsutural vittae long and parallel sided,

almost reaching or enclosing postalar setae; wing with costal band not

crossing vein R, except at apex and ending no more than halfway

between apices of veins R4+5 and M; abdominal tergite III black or with a

black medial vitta sc deret Lp ELEM etre te 2

2 Abdomen orange-brown with broad black lateral markings on tergites IV

and V and a black T-shaped pattern along anterior margin of tergite III

and medially on tergites III-V; face fuscous to black or with a black band

Jons Gr Mare 46 Potosi Ma LA Pese AR a 3

— Abdomen largely black, at most with submedial pale areas on tergite V

and posterior margins of tergites III and IV; face fulvous or with a pair of

latge DICE SPOTS inops eset rrt EE Lo ere rrt e etta d Ee Dama eda 4

3 Wing with a narow infuscation enclosing DM-Cu crossvein; face fulvous

with a narrow black band along oral margin [Indonesia (East Java)] .........

MBA diretti diii c a EEEN too die M ve tutte B. (J.) javanensis (Perkins, 1938)

— Wing without a narrow infuscation enclosing crossvein DM-Cu; face dark

fuscous to black and glossy [Indonesia (West Java)] ................sssss

ibo PURO: NUN Hee CN B. (J.) montana (Hardy, 1983)

4 Face with a pair of large, transverse, oval black spots; notopleural lobe

entirely black; scutellum black at apex; all femora extensively black on

apical half to two-thirds [southern India] ....................sssssssessssse

— Face fulvous without black spots; notopleural lobe at most with anterior

half black; scutellum entirely yellow at apex; fore, mid and often hind

femora entirely fulvous [India, Sri Lanka, Thailand and Vietnam] ...........

Neo cR TRE B. (J.) trilineata (Hardy, 1955)

Discussion

Subgenus Javadacus has no synapomorphies that clearly define it, being

separated from subgenus Zeugodacus solely on the basis of setal reduction

(supra-alar setae absent plus only one pair of scutellar setae). These are

unreliable characters and the subgenus is likely to be polyphyletic. The

Ambonese B. (J.) semisurstyli differs significantly from the other included

species and, apart from the setal characters, closely resembles B. (Z.)

buruensis White from Buru and Sulawesi and B. (Z.) flavipilosa (Hardy) from

Sulawesi in most morphological details, including those of the face, scutum,

wings, legs and abdomen (see Drew and Romig 2013). These three species,

all from Wallacea (Zone C of Hancock and Drew 2015), likely form a related

group within subgenus Zeugodacus.

Australian Entomologist, 2017, 44 (2) 111

The remaining species form two species-pairs: B. (J.) javanensis plus B. (J.)

montana from East and West Java respectively; and B. (J.) scutellaria from

southern India plus B. (J.) trilineata from much of South and Southeast Asia.

The former pair have largely pale abdomens with black T-shaped and lateral

markings, while the latter pair have largely black abdomens. These pairs

might also prove to be unrelated and derived independently from subgenus

Zeugodacus by character reduction. However, all five species are retained

here in Javadacus pending a detailed review of the large and complex

subgenus Zeugodacus sensu stricto.

Of the species here included in Javadacus, two occur in the Indian

subcontinent (Zone A), one being shared with SE Asia (Zone B). The two

Indonesian species (Zone B) are endemic to Java, while the Wallacean

species (Zone C) 1s endemic to Ambon.

Although recent molecular studies (e.g. Virgilio et al. 2015) suggest that

Zeugodacus represents a separate genus closer to Dacus Fabricius than to

other groups of Bactrocera subgenera, a recent study by Jiang et al. (2016),

while supporting a sister-group relationship between Zeugodacus and Dacus

on molecular grounds [shared plesiomorphies?], noted that further evidence

was required before taxonomic raising of Zeugodacus to genus could be

validated. Jiang et al. (2016) noted that such action should be based on more

complete taxon sampling, more comprehensive molecular data combining

mitochondrial genomes and nuclear genes and on more taxonomic, biological

and biogeographic evidence. Morphological and biological data contradicting

a direct Dacus-Zeugodacus relationship were discussed by Hancock and

Drew (2015) and no true synapomorphies linking them have been identified.

We therefore continue to regard Javadacus as belonging in the Zeugodacus

group of subgenera within genus Bactrocera.

Acknowledgements

We thank Daniel Whitmore (Natural History Museum, London) and Susan

Wright (Queensland Museum, Brisbane) for access to specimens.

References

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DE MEYER, M., DELATTE, H., MWATAWALA, M., QUILICI, S., VAYASSIERES, J.-F.

and VIRGILIO, M. 2015. A review of the current knowledge on Zeugodacus cucurbitae

(Coquillett) (Diptera, Tephritidae) in Africa, with a list of species included in Zeugodacus.

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and Oceanian Regions. Memoirs of the Queensland Museum 26: 1-521.

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rainforest habitat of the Western Ghats, India. Raffles Bulletin of Zoology 50: 327-352.

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East Asia. CAB International, Wallingford; 653 pp.

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Entomologist 42(2): 91-104.

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distribution and host plants of fruit flies (Diptera: Tephritidae) in Australia. Information Series

Q199067, Queensland Department of Primary Industries, Brisbane; iii + 75 pp.

HARDY, D.E. 1951. The Krauss collection of Australian fruit flies (Tephritidae—Diptera).

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of the Kansas Entomological Society 28: 3-15.

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and Hardy, D.E. (eds), A catalog of the Diptera of the Oriental region. Vol. III, Suborder

Cyclorrhapha (excluding Division Aschiza). University of Hawaii Press, Honolulu.

HARDY, D.E. 1983. The fruit flies of the genus Dacus Fabricius of Java, Sumatra and Lombok,

Indonesia (Diptera: Tephritidae). Treubia 29: 1-45.

HERING, E.M. 1952. Fruchtfliegen (Trypetidae) von Indonesien (Dipt.). Treubia 21: 263-290.

HUXHAM, K.A. and HANCOCK, D.L. 2002. New records of Dacinae (Diptera: Tephritidae)

from northern Queensland and Torres Strait, Australia. Australian Entomologist 29(4): 123-126.

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first complete mitochondrial genome of Dacus longicornis (Diptera: Tephritidae) using next-

generation sequencing and mitochondrial genome phylogeny of Dacini tribe. Scientific Reports

6: 36426; doi: 10.1038/srep36426. 22 pp.

MAY, A.W.S. 1952. New genera and species of Dacinae (Trypetidae, Diptera) from Queensland.

Queensland Journal of Agricultural Science (1951) 8: 5-13.

NORRBOM, A.L., CARROLL, L.E., THOMPSON, F.C., WHITE, I.M. and FREIDBERG, A.

1999. Systematic database of names. Pp 65-251, in: Thompson, F.C. (ed.), Fruit fly expert

identification system and systematic information database. Myia 9: ix + 524 pp.

PERKINS, F.A. 1938. Studies in Oriental and Australian Trypaneidae. Part II. Adraminae and

Dacinae from India, Ceylon, Malaya, Sumatra, Java, Borneo, Philippine Islands, and Formosa.

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north Queensland, Australia, and improved lures for some pest species. Austral Entomology 54:

411-426.

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Australian Entomologist, 2017, 44 (2): 113-120 113

A REVIEW OF THE PACIFIC ISLANDS SUBGENUS NOTODACUS

PERKINS OF BACTROCERA MACQUART (DIPTERA:

TEPHRITIDAE: DACINAE)

D.L. HANCOCK" and R.A.I. DREW?

18/3 McPherson Close, Edge Hill, Cairns, Old 4870

?International Centre for the Management of Pest Fruit Flies, Griffith University, Old 4111

Abstract

The Bactrocera Macquart subgenus Notodacus Perkins is reviewed and four species recognised,

including one undescribed. Based on both morphological and molecular evidence, the subgenus

is transferred from the Bactrocera group of subgenera to the Melanodacus group of subgenera. A

key to species is included.

Introduction

Subgenus Notodacus Perkins contains a small group of Pacific Island species,

one of economic importance, that belong in the widespread genus Bactrocera

Macquart. Traditionally included in the Bactrocera group of subgenera,

based on morphology of the male terminalia in its type species B. xanthodes

(Broun) (e.g. Drew 1972, 1989), subsequent information from later described

species, together with molecular evidence, indicate an actual placement in the

Melanodacus group of subgenera.

Genus Bactrocera Macquart

Subgenus Notodacus Perkins

Notodacus Perkins, 1937: 56. Type species Dacus xanthodes Broun, 1905 [=

Tephrites (Dacus) xanthodes Broun, 1904], by original designation.

Definition. Abdominal sternite V of male broad with a shallow to deep

posterior emargination (Figs 1-2); posterior lobe of male surstylus short;

pecten of cilia present on abdominal tergite III of male; postpronotal setae

present, placed posterolaterally; supra-alar and prescutellar acrostichal setae

present; one pair of scutellar setae; scutum with a long and narrow medial

postsutural yellow vitta and lateral presutural and postsutural yellow vittae;

scutellum large and bilobed; pair of large [non-shiny] spots (ceromata) on

abdominal tergite V present; head, thorax and abdomen fulvous to red-brown

and with a glossy, transparent appearance; aculeus apically acute.

Response to male lures. Methyl eugenol (2 spp) or none known (2 spp).

Included species. Bactrocera (N.) neoxanthodes Drew & Romig, B. (N.)

paraxanthodes Drew & Hancock, B. (N.) xanthodes (Broun), plus an

undescribed sp. (Drew et al. 1997).

Host plants. Recorded from various families, with one species polyphagous.

Comments. The posterolateral postpronotal seta, glossy, transparent

appearance and large, bilobed scutellum appear to be synapomorphies for this

subgenus. Other defining characters are largely plesiomorphic. The

114 Australian Entomologist, 2017, 44 (2)

placement of the postpronotal seta suggests it is not homologous with the

centrally placed seta seen in B. (Heminotodacus) dissidens Drew, or with the

small, posterocentrally placed seta seen in B. (Zeugodacus) hatyaiensis Drew

& Romig, with the seta evidently derived independently in each of these three

cases.

1 2

Figs 1-2. Bactrocera (Notodacus) spp, male sternite V: (1) B. (N.) neoxanthodes; (2)

B. (N.) xanthodes.

B. (Notodacus) neoxanthodes Drew & Romig (Fig. 3)

Bactrocera (Notodacus) sp. n. No. 2: Drew et al. 1997: 132.

Bactrocera (Notodacus) neoxanthodes Drew & Romig, 2001: 142. Type locality

Kwero, Loh I., Vanuatu.

Fig. 3. Bactrocera (Notodacus) neoxanthodes female from Vanuatu. Photo by Steve

K. Wilson O Pacific Community 2017.

Australian Entomologist, 2017, 44 Q) 115

Distribution. Vanuatu (Loh, Santo and Efaté islands).

Male lure. None known.

Host plants. Barringtonia edulis (Lecythidaceae) and Passiflora suberosa

(Passifloraceae) (Drew and Romig 2001).

Comments. Male sternite V (Fig. 1) broad and weakly emarginate posteriorly.

Adult illustrated in Fig. 3 and by Drew and Romig (2001) and Leblanc et al.

(2012).

B. (Notodacus) paraxanthodes Drew & Hancock

Bactrocera (Notodacus) paraxanthodes Drew & Hancock, 1995: 10. Type locality

Maré, New Caledonia [the handwritten locality ‘Maré’ was interpreted as ‘Mave’

in the original description].

Distribution. New Caledonia: main island and Maré Island (Loyalty Islands).

Records from Vanuatu and Samoa belong elsewhere (Drew et al. 1997).

Male lure. A possible weak attraction to methyl eugenol (Amice and Sales

1997, Drew et al. 1997).

Host plants. Schefflera gabriellae and Meryta sp. (Araliaceae) (Leblanc et al.

2012); a record from Tylophora biglandulosa (Apocynaceae) is doubtful, this

being an asclepiad and host of Dacus (Neodacus) aneuvittatus Drew.

Comments. Shape of male sternite V not discernible on available specimens

due to curvature of the lateral abdominal margins. This species was

illustrated by Drew and Hancock (1995).

B. (Notodacus) xanthodes (Broun) (Figs 4-5)

Tephrites (Dacus) xanthodes Broun, 1904: 306. Type localities Rarotonga, Cook Is;

Suva, Fiji; and Tonga [ex fruit imported into New Zealand].

Tephrites xanthodes Broun, 1905a [February]: 3. Type localities Rarotonga, Cook Is;

Suva, Fiji; and Tonga [ex fruit imported into New Zealand]. Preoccupied: Broun

1904.

Dacus xanthodes Broun, 1905b [June]: 327. Type localities Rarotonga, Cook Is; Suva,

Fiji; and Tonga [ex fruit imported into New Zealand]. Preoccupied: Broun 1904.

Chaetodacus xanthodes (Broun): Bezzi 1928: 105.

Dacus xanthodes (Broun): Malloch 1931: 260.

Notodacus xanthodes (Broun): Perkins 1937: 57.

Dacus (Notodacus) xanthodes (Broun): Hardy 1955: 434.

Bactrocera (Notodacus) xanthodes (Broun): Drew 1989: 170.

Distribution. American Samoa, Cook Islands (southern group), Fiji, French

Polynesia (Austral group), Niue, Rotuma, Samoa, Tonga, Wallis and Futuna;

eradicated from Nauru in 1999 (Leblanc et al. 2012). Records from Vanuatu

are of B. neoxanthodes Drew & Romig (see above).

Male lure. Methyl eugenol.

116 Australian Entomologist, 2017, 44 (2)

Host plants. This species is highly polyphagous, being recorded from fruit of

34 plant species in 20 families, including many of economic importance

(Leblanc et al. 2012). Breadfruit (Artocarpus altilis: Moraceae) is an

important host on many islands (Tora Vueti et al. 1997). Records from

Cucurbitaceae (Citrullus lanatus: Watermelon) appear to refer only to

damaged fruit (Leblanc et al. 2012).

Figs 4-5. Bactrocera (Notodacus) xanthodes from Samoa: (4) male; (5) female.

Photos by Steve K. Wilson O Pacific Community 2017.

Australian Entomologist, 2017, 44 (2) 117

Comments. This species was described three times from the same type series

bred from larvae found in fruit imported into New Zealand from Rarotonga,

Suva and Tonga (Broun 1904, 1905a, 1905b), with their chronological

sequence established by Norrbom ef al. (1999). However, no types were

designated and no specimens attributable to the original series are known to

exist (Drew 1989). Male sternite V (Fig. 2) broad and relatively deeply

emarginate posteriorly. Adult illustrated in Figs 4-5 and by Drew (1974,

1989). For a full description see Drew (1974).

B. (Notodacus) undescribed species 1

Bactrocera (Notodacus) paraxanthodes Drew & Hancock, 1995: 10. Partim: Western

Samoa records only; misidentification.

Bactrocera (Notodacus) sp. n. No. 1: Drew et al. 1997: 132; Leblanc et al. 2012: 35.

Distribution. Samoa.

Male lure. None known.

Host plants. Ficus sp. (Moraceae) (Drew and Hancock 1995), plus Meryta sp.

(Araliaceae) and Mammea glauca (Calophyllaceae) (Leblanc et al. 2012).

Comments. Examined specimens are teneral with curled abdomens, the shape

of male sternite V thus not discernible. This species has not been illustrated

but distinguishing characters were noted by Drew et al. (1997).

Key to species of subgenus Notodacus

* — presumed apomorphic characters.

I Wing with costal band narrow and uniformly fuscous from end of vein

R243 to apex; scutum red-brown, contrasting with fulvous abdomen and

with a narrow whitish to pale yellow medial vitta extending as a fulvous

area to hind margin and fulvous anterior to suture; scutellum red-brown

laterally, fulvous medially and with distinct whitish or yellow lateral

margins over at least basal half*; male sternite V deeply emarginate

posteriorly*; female oviscape extensively black posteriorly* [numerous

South Pacific islands; Figs 4-5] ............... B. (N.) xanthodes (Broun, 1904)

— Wing with costal band either broad and uniformly pale or narrow and pale

except fuscous at apex; scutum orange-brown with the yellow medial vitta

continuing anterior to suture; scutellum without distinct yellow margins

but an indistinct yellow area sometimes present; male sternite V either not

described or with a shallow emargination posteriorly; female oviscape

entirely or almost entirely fulvous ...............ssssssssseRR venn 2

2 Wing with costal band narrow and confluent with vein R;44, dark fuscous

at apex of cell r4+5 but paler elsewhere*; scutellum fulvous centrally and

broadly dark fulvous to pale fuscous laterally, sometimes with a small

yellow basolateral area [Vanuatu; Fig. 3] .....orrrrrvrrrrrvrrrrrrrvvrnrrrrrrvrnrrrrrrrrnnnn

DN. ACER B. (N.) neoxanthodes Drew & Romig, 2001

118 Australian Entomologist, 2017, 44 (2)

— Wing with costal band broad and confluent with vein R4+5, entirely pale,

not dark fuscous apically; scutellum not as above ... i.n 3

3 Wing without a pale, transverse infuscation enclosing crossveins;

scutellum fulvous with black lateral vittae reaching apical setae

posteriorly but ending narrowly before base anteriorly* [New Caledonia]

EE EN ER A B. (N.) paraxanthodes Drew & Hancock, 1995

— Wing with a narrow, pale transverse infuscation enclosing both crossveins

R-M and DM-Cu*; scutellum fulvous without black lateral vittae [Samoa]

eese D. (N.) undescribed sp. (No. 1 of Drew et al. 1997)

Discussion

Subgenus Notodacus species are known only from islands of the South

Pacific (Zone F of Hancock and Drew 2015). They have individual

apomorphies (see Key) and do not appear to form related pairs or triplets.

Rather, they appear to represent vicariant species derived from a single

ancestral entity, with the most polyphagous species, P. xanthodes,

subsequently dispersing (most likely by human introduction) throughout

much of the South Pacific. The remaining three species appear to be endemic

(and restricted) to single island groups (New Caledonia, Vanuatu and Samoa

respectively), suggesting that B. xanthodes originated in Fiji.

The male sternite V (Figs 1-2) is broader than in the Bactrocera group of

subgenera, resembling more that of the Zeugodacus and Melanodacus groups

(cf. Drew 1972, figs 1-2); hence, despite the relatively deep emargination in

the type species, Notodacus is referred here to the Melanodacus group of

subgenera. This is supported by molecular studies (Krosch et al. 2012,

Virgilio et al. 2015), which both placed B. (Notodacus) xanthodes in a clade

with other Melanodacus group subgenera such as Daculus Speiser,

Gymnodacus Munro and (Krosch et al. 2012) Paratridacus Shiraki. Leblanc

et al. (2015) placed B. (N.) xanthodes as basal to both Daculus and the

Bactrocera group.

Jiang et al. (2016) also showed that the Bactrocera and Melanodacus groups

of subgenera formed separate monophyletic clades and followed Krosch et al.

(2012) in placing Tetradacus Miyake as a separate lineage basal to both

groups, all separated from the Zeugodacus group of subgenera (and other

examined genera of Dacinae and Tephritinae) in possessing a TA

(apomorphy) instead of TAA stop codon (plesiomorphy) for the COI gene.

The presence of presutural and postsutural lateral yellow vittae and a medial

yellow vitta on the scutum, plus a single pair of scutellar setae, suggest a

relationship with subgenus Tetradacus (sensu Hancock and Drew 2015),

which has similarly short posterior surstylus lobes and a shallow

emargination to sternite V in males. Tetradacus, however, differs in having a

normally shaped scutellum and in lacking postpronotal, prescutellar

acrostichal and, usually, supra-alar setae.

Australian Entomologist, 2017, 44 (2) 119

Notodacus also resembles the B. (H.) aglaiae (Hardy) group of subgenus

Hemizeugodacus Hardy (sensu Hancock and Drew 2015) in possessing a

medial yellow vitta on the scutum and retaining both supra-alar and

prescutellar acrostichal setae, but the latter has two pairs of scutellar setae.

Hemizeugodacus, however, is likely to be the most closely related subgenus,

the lack of basal scutellar setae in Notodacus being possibly a consequence of

its distinctly modified scutellum.

Acknowledgements

We thank Susan Wright (Queensland Museum, Brisbane) for images of the

handwritten type labels of B. paraxanthodes and access to specimens. We

also thank Steve Wilson (Queensland Museum, Brisbane) and the Secretariat

of the Pacific Community, Nouméa (per Stuart Roberts) for provision of Figs

3-5 and permission to use them.

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EMERY, D.L. and NJ.

Popplepsalta aeroides Owen & Moulds (Hemiptera: Cicadidae):

description of the female

HANCOCK, D.L. and DREW, R.A.I.

A review of the subgenus Jaradacus Hardy of Bactrocera Macquart

(Diptera: Tephritidae: Dacinae)

HANCOCK, D.L. and DREW, R.A.I.

A review of the Pacific Islands subgenus Notodacus Perkins of Bactrocera

Macquart (Diptera: Tephritidae: Dacinae)

HÁVA, J. and LAWRENCE, J.

Contribution to the genus Rano/us Blair (Coleoptera: Dermestidae:

Ranolini), with new synonymy and combinations

HUSTON, D.C.

Train robbery: Menemerus bivitlatus (Dufour, 1831) (Araneae: Salticidae)

steals larvae of Technomyrmex sophiae Forel, 1902 (Hymenoptera:

Formicidae) in transit

LAMBKIN, K. J.

Mesonirvana Evans, 1956, replaces Loscarloides Evans, 1956

(Hemiptera: Cicadomorpha: Prosboloidea: Dysmorphoptilidae),

a homonym of Eoscartoides Matsumura, 1940 (Hemiptera: Cercopidae)

LAMBKIN, T. L

Papilio demoleus malayanus Wallace, 1865 (Lepidoptera: Papilionidae)

on Dauan Island, Torres Strait, Queensland and recent confirmation of

P. d. stheneZinus Rothschild, 1895 in the Lesser Sunda Islands

NIELSEN, J. E.

A tentative record of Papilio demoleus malayanus Wallace, 1865

(Lepidoptera: Papilionidae) from the Solomon Islands

NIELSEN, J. E.

Additional characters for separating adults of Papilio demoleus sthenelus

W.S. Macleay, 1826 (Lepidoptera: Papilionidae) from P. demoleus L.

subspecies of biosecurity concern to Australia

SANDS, D.P.A. and M.C.

Description of Anisynta cynone anomala subsp. n. (Lepidoptera: Hesperiidae)

from the northern tablelands of New South Wales, with a discussion of its

variation, sympatry with and similarity to lv/synta fillyardi Waterhouse & Lyell

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