THE AUSTRALIAN ENTOMOLOGIST

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The Australian Entomologist is a non-profit journal published in four parts annually

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COVER

Tellervo zoilus (Fabricius), mating in rainforest understory at Mission Beach, northern

Queensland. This species is one of a handful of true understory dwelling butterflies in

Australia. Males form leks, mating dances of several individuals, in sunny patches in

the morning. Females approach, then are led by a male to the underside of a nearby

leaf where copulation ensues. Sometimes the female leads and initiates genital contact.

As in many butterflies, the male, right, enters a catatonic state during ejaculation. Pen

and ink drawing by Caloundra ESQ member, Dr Albert Orr, whose illustrated books on

butterflies and dragonflies have won awards in Australia and overseas. His second book

on New Guinea Odonata has just appeared (see Australian Entomologist 43 (1): 38).

Australian Entomologist, 2016, 43 (4): 173-188 173

REVIEW OF THE AUSTRALIAN SPECIES OF ARCTORNIS

GERMAR, 1810 (LEPIDOPTERA: EREBIDAE: LYMANTRIINAE)

A.P. MACKEY

PO Box 404, Yandina, Qld 4561

Abstract

Four new species of Arctornis Germar, 1810 are described and illustrated from Australia:

Arctornis queenslandica sp. n., Arctornis lucens sp. n., Arctornis ravenshoeae sp. n., Arctornis

cairnsae sp. n. and Arctornis commoni sp. n. All are confined to northeastern Queensland.

Introduction

The genus Arctornis Germar, 1810 is widely distributed throughout the

Oriental tropics, Sundaland and New Guinea and also extends into the

Palaearctic region. It is extremely diverse and, as currently defined, probably

includes at least 200 species. Only in Borneo has a major part of the fauna

been revised (Holloway 1999). The genus has long been recognised as being

represented in Australia (as Redoa Walker, 1855), with the first recorded

species wrongly identified as the Oriental Arctornis submarginata Walker,

1855 (Turner 1921). The Australian checklist of Lepidoptera dealt with the

issue of A. submarginata and listed one undescribed species (Edwards 1996).

In his review of the Bornean species, Holloway (1999) provided the current

interpretation of Arctornis and broadly defined the genus on the basis of the

male genitalia: the harpe is well developed and arises from the sacculus and

the aedeagus is short and usually has a pair of finger-like processes

developed on the rim of its orifice. Tymbals are present in the abdomen of the

male. Adults are frequently very similar in general facies, being satiny white,

sometimes with a small black discal spot. Correct determination usually

requires examination of the genitalia. This similarity also makes it difficult to

associate males with females.

In the material examined for this present study, several specimens bear labels

indicating that they had been barcoded. Sexes were associated in three

species by use of this barcoding, which was based on similarity of COI

sequences. Details of analyses, sequence data, geographical and ecological

data are available through the Barcode of Life Data System (BOLD:

Ratnasingham and Hebert 2007) and BOLD process ID's are given for the

relevant specimens under material examined. In the descriptions below, both

sexes are similar unless otherwise indicated, while forewing length is the

distance from the base of the wing at its articulation to the tip of the apex.

The known Australian fauna comprises five species, which are described and

illustrated below.

Abbreviations: AM - Australian Museum, Sydney; ANIC - Australian

National Insect Collection, Canberra; APM - collection of A.P. Mackey,

Yandina; UQIC — University of Queensland Insect Collection (housed at the

Queensland Museum); QM - Queensland Museum, Brisbane.

174 Australian Entomologist, 2016, 43 (4)

Arctornis Germar, 1810

Cassidia Walker, 1862

Chatracharta Walker, 1862

Ciaca Walker, 1865

Cobanilla Moore, [1883] 1882-3

Kanchia Moore, [1883] 1882-3

Lymantralex Collenette, 1938

Redoa Walker, 1855

Scarpona Walker, 1862

Topomesa Walker, 1866

Type species: Arctornis l-nigrum (Müller, 1764).

Arctornis queenslandica sp. n.

(Figs 1-7)

Types. Holotype 6, QUEENSLAND: 14 km W by N of Hope Vale Mission, 8.x.1980,

E.D. Edwards, ANIC Genitalia Slide no. 19839, in ANIC. Paratypes: 1 9, 14 km W

by N of Hope Vale Mission, 8.x.1980, E.D. Edwards, ANIC Genitalia Slide no.

19838, in ANIC; 1 &, Kuranda, 1-15.ii.2008, D.C.F. Rentz, ANIC Genitalia Slide no.

19846, in ANIC; 1 9, Hervey's Range, 8.xi.2006, G. Cocks, G290, DNA Barcode

LOQT506-06, gvc6341, in APM.

Additional material examined. QUEENSLAND: 2 <3), Hervey's Range, June 2006 &

26.1v.2007, G. Cocks, DNA Barcodes: LOQT380-06, gvc5833-1L and LOQTB344-

07, gvc7087-1L respectively, in APM; 1 3, 1 9, Hervey's Range, l.v. 2009, G.

Cocks, in APM; 2 43), Cairns, 8.1.1987 & 1.1.1989, A.P. Mackey, in APM; 1 4, West

Claudie River, 4 km SW of road junction, 1.xii.1986, G. Daniels & M. Schneider, in

UQIC.

Figs 1-2. Arctornis queenslandica sp. n: (1) Holotype male, Hope Vale Mission, Qld;

(2) Paratype female, Hope Vale Mission, Qld.

Description (Figs 1-2). Head lightly scaled, white, with pair of purplish

brown dots between eyes just below antennal bases; pair of thin purplish

brown bars between antennal bases which extend almost to mid-line; labial

palp upturned, white, with purplish brown tip; antennal filament with fine

dusting of white scales that become worn away, pectinations pale reddish

Australian Entomologist, 2016, 43 (4) 175

yellow, long in male but very short in female. Thorax white; legs white, pro-

and mesothoracic legs with purplish brown spot towards proximal end of

tibia and on first tarsal segment. A faint spot may be present on joint between

femur and tibia. Abdomen lightly scaled, silvery white. Forewing length:

male 18-20 mm; female 28 mm. Both forewing and hindwing lightly scaled

and glistening white. Scales in central part of wings appear to be lost easily

leaving transparent white wing membrane and some sparse areas of

glistening scales along margins and discal vein; this effect more apparent in

female where most scales are lost. Underside similar except that scales less

glistening.

+ ^

Figs 3-4. Arctornis queenslandica male genitalia: (3) ventral view, Hope Vale

Mission, Qld, slide ANIC19839; (4) aedeagus, Kuranda, Qld, slide ANIC19846. Scale

bars ca 1 mm.

Male genitalia (Figs 3-4). Tegumen and vinculum lightly sclerotised and

loosely united; tegumen hood-like; vinculum a narrow U-shaped strap; saccus

weak; uncus well sclerotised, short, broad, deep and bifid, slightly hooked,

posterior face crenulate, punctulate and setose with long setae dorsally and

laterally; valva elongate, subrectangular and with bluntly rounded apex, costa

folded inwardly, sacculi large and well developed, adpressed but not fused

basally, outer surface of dorso-distal part of valva bears small, triangular lobe

covered with long, fine, setae, internal ventro-distal part of valva is in form of

stout, downturned beak, harpe well developed and partly embedded in pouch

formed between sacculus and costal fold, flat, slender, slowly tapering to fine

point, about two thirds the length of the valva; juxta moderate but lightly

sclerotised, slightly reflexed, strongly concave, dorsal edge emarginate in

176 Australian Entomologist, 2016, 43 (4)

ventral view; aedeagus short, broad and deep, weakly sclerotised except for

two slender, finger-like dorso-lateral projections from the carina penis,

caecum reflexed ventrally, vesica without ornamentation.

Female genitalia (Figs 5-7). Papillae anales shallowly and bluntly triangular,

covered with long fine setae; apophyses posteriores slender and slightly

curved and tapering, extending well into segment VIII; segment VIII broad

and collar like, apophyses anteriores very short and truncate; pseudopapillae

long, narrow, tapering slightly from posterior to anterior, setose; lamella

postvaginalis almost membranous, crescent-shaped, bearing a series of long

setae along its posterior edge; the deep channel-shaped sinus vaginalis

occupies most of the medial area of sternite VIII, the lamella antevaginalis

forms a small cup at its anterior part and at the bottom of the cup is a

complex of sclerotised folds; the opening of the ostium bursa is situated at the

end of the channel and broadens toward thin-walled sac of corpus bursae;

signum large and ovate, covered with rugged, short, stout spines.

Figs 5-7. Arctornis queenslandica female genitalia, Hervey's Range, Qld, slide G290:

(5) overview; (6) sterigma; (7) signum. Scale bars ca 1 mm (5-6) or 0.5 mm (7).

Australian Entomologist, 2016, 43 (4) 177

Diagnosis. This is the largest of the Australian Arctornis species and the only

one with the satiny, translucent sheen to the wings. The male and female

genitalia are distinctive and unlike any other Australian species.

Etymology. queenslandica (Latin adjective) — pertaining to Queensland.

Distribution. North Queensland: coastal ranges from Townsville north to Iron

Range in Cape York Peninsula.

Remarks. Sexes, originally associated by size, were confirmed to be

conspecific by barcoding. Specimens in collections often show pale cream

discolouration along the forewing costa, at the vein bases and on the thorax

and abdomen, but these marks are not found on freshly caught material. This

species is similar to Arctornis perfecta (Walker, 1862) from Sundaland and

Sulawesi, but in A. perfecta the harpe is much longer, extending well beyond

the end of the valve, and in the female the lamella postvaginalis has a large

medial lobe on the anterior margin (Holloway 1999: figs 354, 378).

Arctornis lucens sp. n.

(Figs 8-14)

Types. Holotype á, QUEENSLAND: Mission Beach near Tully, 30.x.1967. R.

Dobson, ANIC Genitalia Slide no. 19834, in ANIC. Paratypes: 1 9, Cairns, 1-

20.vii.2005, P. Hebert, G336, DNA Barcode: LOQC411-05, 05-QLD-00411; 1 d,

Kuranda, 1-15.x1i.2005, D.C.F. Rentz. ANIC Genitalia Slide no. 19833, ANIC DNA

no. 000971, in ANIC.

Additional material examined. QUEENSLAND: 1 4, Kuranda, 16-31.xii.2005,

D.C.F. Rentz. ANIC DNA no. 000970; 1 £, 1 km SE of Mt. Cook, Cooktown,

13.x.1980, E.D. Edwards, ANIC Genitalia Slide no. 19836; 1 (4, Mt. Webb National

Park, 50 km N. Cooktown, 11-14.vii.1976, G.B. & S.R. Monteith, ANIC Genitalia

Slide no. 19842, all in ANIC; 1 ĝ, Barrine, April 1948, K469601, in AM; 3 &d,

Kuranda, 1.iv.1928, 3.iv.1928 & 1.iv.1928, G143, G144 & G150, all in UQIC.

Figs 8-9. Arctornis lucens sp. n: (8) Holotype male, Mission Beach, Qld: (9) Paratype

female, Cairns, Qld.

178 Australian Entomologist, 2016, 43 (4)

Description (Figs 8-9). Head white, frons lightly speckled with brownish

orange, vertex brownish orange; labial palp upturned, brownish orange;

antennal scape brownish orange ventrally and laterally, white dorsally,

filament white, pectinations long in male, shorter in female, pale brownish

yellow. Thorax moderately scaled, white; legs white, fore and mid tibia with

large purplish brown spot sometimes with brownish orange edges proximally,

all legs with purplish brown spot on first tarsal segment and brownish orange

fifth tarsal segment. Abdomen lightly scaled, silvery white. Forewing length:

male 15-16 mm; female 19 mm. Both forewing and hindwing glistening

white, forewing with coruscations caused by irregular longitudinal ridges of

scales, hindwing less glistening particularly within discal region; forewing

with minute black discal spot at inflection of r-m; termen of fore- and

hindwing with faint greyish tinge especially towards the middle. Underside of

both wings milky white.

Figs 10-11. Arctornis lucens male genitalia, Mission Beach, Qld, slide ANIC19834:

(10) ventral view; (11) aedeagus. Scale bars ca 1 mm.

Male genitalia (Figs 10-11). Tegumen and vinculum united; tegumen hood-

like, narrowing considerably laterally to join very narrow vinculum, which

widens ventrally; saccus weak; uncus short, broad, slightly hooked, shoulders

broad, dorsally towards base a small patch of long fine setae either side of

mid-line, a ventral, membranous transverse ridge extending as lobe on each

side of uncus and bearing long fine setae may represent socii; valva lightly

sclerotised, broad, elongate, subrectangular, broadening slightly distally,

Australian Entomologist, 2016, 43 (4) 179

Figs 12-14. Arctornis lucens female genitalia, Cairns, Qld, slide G336: (12) overview;

(13) sterigma; (14) signum. Scale bars ca 1 mm (12, 14) or 0.5 mm (13).

sacculi well developed, harpe moderate, well sclerotised, tubular, tapering to

blunt point, strongly reflexed ventrally, distal half bearing rows of small

spines; annellus lightly sclerotised; juxta lightly sclerotised, r-shaped in

lateral view, subrectangular plate in ventral view, longer sides emarginate;

aedeagus short, broad and deep, lightly sclerotised except for two well

180 Australian Entomologist, 2016, 43 (4)

sclerotised, dorso-lateral finger-like lobes arising from the carina penis,

coecum relatively long, reflexed ventrally, vesica without ornamentation.

Female genitalia (Figs 12-14). Papillae anales short, broad, rounded and

rather narrow in terminal view, covered with long setae, apophyses

posteriores moderate, projecting about half way into segment VIII, ribbon-

like, truncate; segment VIII a broad collar, zone of long setae around the

posterior margin, mid-ventrally a short broad channel extending about half

Way across sternite marking opening of sinus vaginalis, lamella antevaginalis

a simple lip at anterior end of channel, lamella postvaginalis at posterior end

of the channel very lightly sclerotised, rectangular with poorly defined

transverse median ridge, apophyses anteriores ribbon-like, very short;

pseudopapillae well developed, elongate; an almost membranous, narrowly

crescentic sclerite bearing some long setae present intersegmentally between

segments VIII and the ovipositor lobes; ostium bursae opening directly

behind lip of lamella antevaginalis; ductus bursae very short, membranous,

leading into neck of pyriform corpus bursae; signum large, inverted leaf-

shaped covered with rugged, short, stout spines, smaller towards a

longitudinal, medial area.

Diagnosis. This is the only Australian species in which males have a robust

and strongly reflexed harpe. Females can be distinguished by the rugged

spines of the signum.

Etymology. lucens (Latin adjective) — shining.

Distribution. North Queensland: coastal areas from Mission Beach north to

Mt Webb.

Remarks. This is the most commonly collected Australian Arctornis species

and is likely to be more widely distributed than recorded here. Males and

females were associated by barcoding.

Arctornis ravenshoeae sp. n.

(Figs 15-21)

Types. Holotype 6, QUEENSLAND: 9 miles SSE of Ravenshoe, 21.iv.1969, LF.B.

Common & M. Upton, 2750 ft, ANIC Genitalia Slide no. 19835, in ANIC. Paratypes:

1 9, Kuranda, 1-15.iv.2005, D.C.F. Rentz, ANIC Genitalia Slide no. 19832, ANIC

DNA no. 000698; 1 9, Mission Beach nr. Tully, 24-28.1967, R. Dobson, ANIC

Genitalia Slide no. 19841; 1 4 Cooper's Creek, 26.viii.2003, P.D.N. Hebert, G541,

DNA Barcode: LOQB517-05, Moth 210.03CC, all in ANIC.

Additional material examined. QUEENSLAND: 1 4, 4 mls W of Babinda,

10.iii.1964, LF.B. Common & M. Upton; 1 ĝ, Kuranda, 15-30.vi.2004, D.C.F. Rentz,

ANIC Genitalia Slide no. 19843; 1 £, Kuranda, 1-15.vii.2005, D.C.F. Rentz, ANIC

Genitalia Slide no. 19845; 1 ĝ, Lizard Island, 1.v.2001, P.D.N. Hebert, G333, DNA

Barcode: LOQB302-05, Moth 302.01LZ, all in ANIC; 1 9, Lake Barrine, April 1939,

E.J. Dumigan, G 158, in UQIC.

Australian Entomologist, 2016, 43 (4) 181

Figs 15-16. Arctornis ravenshoeae sp. n: (15) Holotype male, Ravenshoe, Qld; (16)

Paratype female, Mission Beach, Qld.

Description (Figs 15-16). Head white, frons speckled with light orange,

vertex light orange bar between antennae; labial palp white, external surface

lightly speckled with orange, more densely orange towards tip; antennal

scape light to deep orange, filament white, pectinations long in male, shorter

in female, pale brownish yellow. Thorax, white, lightly scaled; legs white,

fifth tarsal segment orange, first two legs purplish brown spot on proximal

part of tibia and similar spot on first tarsal segment, leg 3 may have small

often insignificant pale purplish spot on first tarsal segment. Abdomen lightly

scaled, white. Forewing length: male 15-17 mm; female 18-19 mm. Forewing

and hindwing glistening white, forewing with slight coruscations caused by

fine, irregular longitudinal scale ridges, hindwing less glistening particularly

within discal region; in some males forewing with a minute and

inconspicuous black discal spot on r-m, discal spot minute but distinct in

female; fringes silvery white but in some specimens a hint of grey along

termen. Underside of both wings milky white.

Male genitalia (Figs 17-18). Tegumen and vinculum loosely united; tegumen

hood-like, a narrow strap laterally; vinculum narrow U-shaped band

widening only slightly in mid line, saccus not developed; uncus short, broad,

slightly hooked, shoulders broad, dorsally towards the base with small patch

of long fine setae either side of mid-line, a ventral, membranous transverse

ridge extending as a lobe on each side of uncus and bearing long fine setae

may represent socii; valva lightly sclerotised, subrectangular, widening

slightly distally, broadly and shallowly bilobed at apex, lobes bearing long

fine setae, sacculi well developed, harpe extending almost complete length of

valve, ribbon-like with longitudinal rows of small, robust spines, lightly

curved and tapering to a sharp point; juxta lightly sclerotised, r-shaped in

lateral view, a subrectangular plate in ventral view with two large lateral

depressions either side of a median ridge, both long edges emarginate;

aedeagus short, broad and deep, lightly sclerotised except for two well

sclerotised, stout, dorso-lateral finger-like lobes arising from carina penis,

coecum relatively long, reflexed ventrally, vesica without ornamentation.

182 Australian Entomologist, 2016, 43 (4)

Figs 17-18. Arctornis ravenshoeae male genitalia: (17) ventral view, Cooper's Creek,

Qld, slide G541; (18) aedeagus, Ravenshoe, Qld, slide ANIC19835. Scale bars ca 1

mm.

Female genitalia (Figs 19-21). Papillae anales short, broad, in terminal view

rounded and rather narrow, densely setose, apophyses posteriores moderate,

slightly tapering and extending about half way into segment VIII; segment

VIII a broad collar with zone of long setae around posterior margin, mid-

ventrally a short broad, partly sclerotised channel extends about half way

across sternite and marks opening of sinus vaginalis, lamella antevaginalis

forming slight lip at anterior end of channel, lamella postvaginalis present at

posterior end of channel forming outwardly projecting shelf that in some

views appears as crescentic plate, apophyses anteriores short, flat, barely

projecting into segment VII; pseudopapillae well developed, elongate,

widening slightly anteriorly; ventrally intersegmental membrane between

VIII and ovipositor lobes has an almost membranous sclerite which forms

narrow crescentic band; ductus bursae short, membranous; corpus bursae

pyriform; signum large, inverted leaf-shaped, covered with short, stout spines

except for longitudinally elongate central area.

Diagnosis. Of the Australian Arctornis species, A. ravenshoeae is only likely

to be confused with A. /ucens and males of A. commoni but the genitalia are

distinct. Males of A. ravenshoeae may be distinguished by the long, straight

harpe; females by the presence of the shelf-like lamella postvaginalis and a

signum with gracile spines absent in elongate central area.

Australian Entomologist, 2016, 43 (4) 183

—

Figs 19-21. Arctornis ravenshoeae female genitalia: (19) overview and (20) sterigma,

Kuranda, Qld, slide ANIC19832; (21) signum, Mission Beach, Qld, slide

ANIC19841. Scale bars ca 1 mm (19-20) or 0.5 mm (21).

184 Australian Entomologist, 2016, 43 (4)

Etymology. ravenshoeae (Latin adjective) — pertaining to Ravenshoe, in

recognition of the type locality of the species.

Distribution. North Queensland: east coast ranges and adjacent areas from

Lizard Island south to Mission Beach.

Remarks. The purplish brown spots on the legs may be obvious or very pale

and insignificant; they may be narrowly fringed with orange and the spot on

the first tarsal segment of leg 2 may be almost black. The sexes were

associated by barcoding.

Arctornis cairnsae sp.n.

(Figs 22-24)

Types. Holotype 3, QUEENSLAND: Whitfield Range near Cairns, 3.iv.1975, M.S. &

B.J. Moulds, ANIC Genitalia No. 19837, in ANIC. Paratype d, Kuranda, 1-

15.xii.2004, D.C.F. Rentz, 335 m, ANIC Genitalia Slde no. 19844, in ANIC.

Additional material examined. QUEENSLAND: 1 ĝ, Lake Barrine, 7.1.1939, EJ.

Dumigan in UQIC; 2 $4, Lake Barrine, April 1948, K469603 & K469605, in AM.

Fig. 22. Arctornis cairnsae sp. n. Holotype male, Whitfield Range, Cairns, Qld.

Description (Fig. 22). Male, head white, frons pale buff, vertex brownish

orange; labial palp white ventrally, external lateral surface brownish orange;

antennal scape brownish orange, filament pale golden yellow, pectinations

long, brownish yellow. Thorax pale cream, white ventrally, sparsely scaled;

legs, femur white, tarsus and tibia pale cream. Abdomen sparsely scaled, pale

cream, slightly glistening. Forewing length: male 15.5-16 mm. Wings lightly

scaled, ground colour of fore- and hindwings pale ivory with slight brown

irroration particularly along veins and termen, which at some angles appear

glistening pale gold; fringes of termen brown. Forewing with small but

distinct black, discal spot on r-m. Hindwing paler towards anal margin, which

has long white fringes. Underside pale ivory.

Male genitalia (Figs 23-24). Tegumen and vinculum loosely united; tegumen

hood-like, very thin and strap-like laterally; vinculum narrow V-shaped band

Australian Entomologist, 2016, 43 (4) 185

with slight downward bow in ventral midline, saccus not developed; uncus

short and broad with wide shoulders, slightly narrowing towards blunt,

slightly hooked end, shoulders bear a small group of long fine setae dorsally

either side of mid line; socii may be represented by ventral, membranous

transverse ridge extending as lobe each side of uncus and which bears long

fine setae; valva subrectangular, long, sharply elbowed dorsally close to base,

gently tapering distally to bluntly rounded valvula, sacculus well developed

and sharply angled at basal corner, another rather slight angle at about one

third to one half of ventral margin of the valve, harpe well developed, ribbon-

like, about half length of valve, strongly curved at one third and gradually

tapering to fine point; juxta r-shaped in lateral view, a subrectangular plate in

ventral view with ventral margin and antero-ventral angles gently rounded,

lip of dorsal margin supporting aedeagus slightly bilobed; aedeagus short,

broad and deep, lightly sclerotised except for two short, well sclerotised,

dorso-lateral finger-like lobes arising from carina penis, coecum relatively

long, reflexed ventrally and slightly inflated at its anterior end, vesica without

ornamentation.

Figs 23-24. Arctornis cairnsae male genitalia, Whitfield Range, Cairns, Qld, slide

ANIC19837: (23) ventral view; (24) aedeagus. Scale bars ca 1 mm.

Diagnosis. The colouration of the forewings of A. cairnsae is distinctive

among Australian Arctornis.

186 Australian Entomologist, 2016, 43 (4)

Etymology. cairnsae (Latin adjective) — pertaining to Cairns in recognition of

the type locality of the species.

Distribution. Only recorded from the Cairns region of northern Queensland.

Remarks. Only the male is known.

Arctornis commoni sp.n.

(Figs. 25-27)

Types. Holotype 3, QUEENSLAND: Iron Range, 7.iv.1964, IL.F.B. Common & M.S.

Upton, ANIC Genitalia Slide no. 19840, in ANIC. Paratypes: 2 64, Gordon Creek

area, Claudie Riv. district, 22-23.vi.1982, M.A. Schneider, UQIC Reg.# 812816 and

81202, in UQIC.

Fig. 25. Arctornis commoni sp. n. Holotype male, Iron Range, Qld.

Description (Fig. 25). Head white, frons heavily irrorated with orange, vertex

orange; labial palp white, external surface irrorated with orange, more

heavily towards the tip; antennal scape orange, filament white, pectinations

long, pale brownish yellow. Thorax white, lightly scaled; legs white, first and

second legs with small, pale orange spot on proximal part of tibia, first and

fifth tarsal segments light orange, third leg only the fifth tarsal segment

marked with pale orange. Abdomen sparsely scaled, white. Forewing length:

male 15-16 mm. Wings lightly scaled, glistening white, forewing

coruscations caused by irregular longitudinal ridges of scales and with a

minute black discal spot on r-m, hindwing matt white across basal half;

fringes grey along termen. Underside matt white.

Male genitalia (Figs 26-27). Tegumen and vinculum united; tegumen hood-

like, narrowing laterally to join vinculum; vinculum narrow, V-shaped band,

saccus not developed; uncus short and broad, shoulders wide, hardly

narrowing toward blunt, slightly hooked end, shoulders bearing small group

of long, fine setae dorsally either side of mid line; socii may be represented

by ventral, membranous transverse ridge extending as lobe each side of uncus

and which bears long, fine setae; valva subrectangular, long, sharply

Australian Entomologist, 2016, 43 (4) 187

convex dorsally close to base, gently tapering distally to bluntly rounded

valvula, sacculus well developed and rounded at basal corner, harpe well

developed, narrow, ribbon-like, extending about half length of valve, sharply

tapering to blunt point; juxta r-shaped in lateral view, subrectangular plate in

ventral view, the ventral margin medially notched, the antero-ventral angles

gently rounded, lip of dorsal margin supporting the aedeagus slightly bilobed;

aedeagus short, broad and deep, lightly sclerotised except for two well

sclerotised, dorso-lateral finger-like lobes arising from carina penis, coecum

relatively long, reflexed ventrally, vesica without ornamentation.

Figs 26-27. Arctornis commoni male genitalia, Iron Range, Qld, slide ANIC19840:

(26) ventral view; (27) aedeagus. Scale bars ca 1 mm.

Diagnosis. Arctornis commoni has a similar fascies to A. lucens and A.

ravenshoeae but the short, straight parallel-sided harpe easily distinguishes it

from the latter two species. It is possible that worn or slightly discoloured

specimens of A. commoni may be confused with A. cairnsae and these

species have similar male genitalia. They may be distinguished by the valve

in A. cairnsae which is narrower and more or less parallel-sided in its basal

half, the sacculus of A. cairnsae is angled along the valve margin, the harpe

of A. cairnsae is shorter after the elbow, the aedeagus of A. cairnsae is

broader, the finger-like processes on the carina penis therefore appear more

widely separated at their base, and they are more gracile than in A. commoni,

the coecum in A. cairnsae is slightly inflated but is not in A. commoni.

188 Australian Entomologist, 2016, 43 (4)

Etymology. commoni (Latin noun in genitive case) — in honour of Dr Ian

Common, who helped lay the foundations for the study of Australian moths

and who first collected this species.

Distribution. Only recorded from Iron Range in Cape York Peninsula,

northern Queensland.

Remarks. Only the male is known. The thorax and wing bases are often

discoloured.

Discussion

The Australian Arctornis species have fascies typical of the genus: broad,

triangular wings that are generally glistening or satiny white or, occasionally,

pale brownish or yellowish. It is possible that Australian species also occur in

New Guinea, which is known to have a diverse Arctornis fauna although only

three species are described. The Australian species were compared with the

described New Guinea species but proved quite distinct. The biology of the

Australian species is completely unknown but is likely to be similar to that of

other species in the genus. The little that is known for the genus was

summarised by Holloway (1999). Most of the known host plant families

occur in Australia, as do some known host plant genera such as Mangifera,

Terminalia, Cinnamomum, Litsea and Camellia. Although Mangifera and

Camellia are not native to Australia, they may be potential hosts for

Australian Arctornis given the polyphagous habit of many lymantriine

species.

Acknowledgements

Sincere thanks to the staff of ANIC, AM, UQIC and QM for access to

specimens in their care and to You Ning Su for making available the

photomicroscope and for instructions on its use. Dave Britton, Graeme

Cocks, Greg Daniels, You Ning Su and Susan Wright have been particularly

helpful in providing access to materials in their care. I thank Graeme Cocks,

David Mitchell and You Ning Su for providing access to barcode results. I

am grateful to Marianne Horak for her critical reading of the manuscript.

References

EDWARDS, E.D. 1996. Lymantriidae. Pp 275-277, in: Nielsen, E.S. Edwards, E.D. and Rangi,

T.V. (eds), Checklist of the Lepidoptera of Australia. Monographs of Australian Lepidoptera

Vol. 4. CSIRO Publishing, Collingwood.

HOLLOWAY, J.D. 1999. The moths of Borneo: family Lymantriidae. Malayan Nature Journal

53: 1-188.

RATNASINGHAM, S. and HEBERT, P.D.N. 2007. BOLD: the barcode of life data system

(www.barcodinglife.org). Molecular Ecology Notes 7: 355-364.

TURNER, A.J. 1921. Revision of the Australian Lepidoptera — Liparidae. Proceedings of the

Linnaean Society of New South Wales 45: 474-499.

Australian Entomologist, 2016, 43 (4): 189-210 189

A REVIEW OF THE SUBGENUS BULLADACUS DREW &

HANCOCK OF BACTROCERA MACQUART (DIPTERA:

TEPHRITIDAE: DACINAE), WITH DESCRIPTION OF TWO NEW

SPECIES FROM PAPUA NEW GUINEA

R.A.I. DREW! and D.L. HANCOCK?

! International Centre for the Management of Pest Fruit Flies, Griffith University, Qld 4111

78/3 McPherson Close, Edge Hill, Cairns, Qld 4870

Abstract

The Bactrocera Macquart subgenus Bulladacus Drew & Hancock is reviewed. Bactrocera

(Bulladacus) trilobata sp. n. and Bactrocera (Bulladacus) wanangiae sp. n. are described from

Papua New Guinea. Bactrocera obtrullata White & Evenhuis, 1999, is placed as a new synonym

of B. diaphana (Hering, 1953); both were described from Papua Province in eastern Indonesia.

The Solomon Islands species B. unipunctata (Malloch) is transferred from subgenus Bactrocera

to Bulladacus. Males of B. aceromata White & Evenhuis are newly described. Records of B.

mcgregori (Bezzi) from Singapore, West Malaysia and the Andaman and Nicobar Islands are

regarded as misidentifications of B. cinnabaria Drew & Romig. Bactrocera pacificae Drew &

Romig is newly recorded from Papua New Guinea. A key to the 20 known species is included.

Introduction

The Bactrocera Macquart subgenus Bulladacus Drew & Hancock is defined

as having the posterior lobe of the male surstylus short, abdominal sternite V

of male deeply concave on posterior margin [this combination of characters

placing it in the Bactrocera group of subgenera], plus a combination of all or

most of the following characters: antenna shorter than usual for the genus,

pecten of cilia present on male abdominal tergite III, distinct oval shining

spots (ceromata) on abdominal tergite V absent, one pair of scutellar setae,

prescutellar acrostichal seta present, supra-alar seta present or absent, wing

cell bcu with extension short, wing of male with bulla present near apex of

cell bcu extension (Drew and Hancock 1995, White and Evenhuis 1999,

Drew and Romig 2013). In addition, both the anatergite and katatergite are

largely yellow, the notopleural lobe is often partly or entirely red-brown,

fuscous or black, facial spots are often absent and most species have a broad

anepisternal yellow stripe that reaches the postpronotal lobe and an isolated

medial black vitta on tergite V that often crosses onto tergite IV but seldom

extends onto tergite III. The male bulla and pecten of cilia on abdominal

tergite III are absent in B. aceraglans White & Evenhuis but a patch of cilia

occurs on the wing where the bulla is normally situated. In males where a

distinct bulla is present, the cell bcu extension is often longer and narrower

than usual. No species in the subgenus shows a response to known male

lures.

Bulladacus appears to be most closely related to subgenus Calodacus

Hancock that, like it, has comparatively short antennae, a short cell bcu

extension and shows no response to known male lures (Hancock 2015); the

latter subgenus differs primarily in the presence of ceromata on abdominal

tergite V. Known hosts of Bulladacus species are primarily the fruit of

190 Australian Entomologist, 2016, 43 (4)

Gnetum gnemon (Gnetaceae) and Terminalia species (Combretaceae), with

only occasional records from other plant families.

The following abbreviations have been used: QDAF - Queensland

Department of Agriculture and Fisheries collection, Brisbane; QM —

Queensland Museum, Brisbane. Descriptive terminology follows White et al.

(1999), with bilateral structures (vittae, setae, etc.) listed in the singular.

Bactrocera (Bulladacus) Drew & Hancock

Bactrocera (Bulladacus) Drew & Hancock, 1995: 9. Type species: Bactrocera

gnetum Drew & Hancock, by original designation.

Twenty species are known, the subgenus ranging from the Philippines and

southern Thailand/Andaman Islands to the South Pacific as far east as Fiji

and Samoa. Two species occur in Australia, five in the Pacific Islands and at

least nine in Papua New Guinea.

Bactrocera (Bulladacus) aceraglans White & Evenhuis

Bactrocera (Bulladacus) aceraglans White & Evenhuis, 1999: 506 (). Type locality:

Minj, Western Highlands, Papua New Guinea.

Description. See White and Evenhuis (1999). This species differs from all

others in the presence of a patch of cilia, rather than a bulla, alongside wing

cell bcu in males. The notopleural lobe is dark fuscous to black.

Distribution. Papua New Guinea (Western Highlands Province).

Host plant. Unknown.

Bactrocera (Bulladacus) aceromata White & Evenhuis

(Fig. 1)

Bactrocera (Bulladacus) aceromata White & Evenhuis, 1999: 509 (9). Type locality:

Tapini, Papua New Guinea.

Material examined. PAPUA NEW GUINEA: Madang Province — 29 $4, 28 99,

Baitabag and Ohu, March, April and September 2001, reared from Gnetum gnemon

and Gnetum costatum; Morobe Province — 24 34, 28 99, Lae, February, March and

April 2000, reared from Gnetum gnemon and Sandoricum koetjape. Specimens in

QDAF.

Description. Male. Head: height 1.2 mm; frons length 1.3 times breadth, red-

brown without dark markings; setae black: 2 frontal, 1 orbital; lunule red-

brown; ocellar triangle black; vertex red-brown; face entirely fulvous without

dark markings, length 0.5 mm; gena fulvous without dark markings, black

seta present; occiput red-brown, fulvous along eye margins; occipital row

with 3-6 strong black setae. Antenna with all segments red-brown; length of

segments 0.12 mm, 0.24 mm, 0.45 mm; first flagellomere (segment 3)

strongly truncate (square) across apex (not rounded as in most Bactrocera

species).

Australian Entomologist, 2016, 43 (4) 191

Fig. 1. Bactrocera (Bulladacus) aceromata White & Evenhuis, male.

Thorax. Scutum entirely red-brown with a broad transverse black band across

posterior margin. Yellow markings as follows: postpronotal lobe;

anepisternal stripe reaching postpronotal lobe dorsally, continuing to

katepisternum as a transverse spot, anterior margin straight; anatergite

(posterior apex black); anterior 2/3 of katatergite (remainder black); broad

parallel-sided lateral postsutural vitta beginning with a large spot anterior to

notopleural suture and ending at intra-alar seta; medial postsutural yellow

vitta absent. Postnotum red-brown centrally, black laterally. Notopleural lobe

red-brown. Scutellum yellow except for a narrow black basal band. Setae: 2

scapular; 2 notopleural; 1 anepisternal; 1 supra-alar; 1 postalar; 1 intra-alar; 1

prescutellar acrostichal; 1 (apical) scutellar.

Legs. All leg segments entirely fulvous except apical three segments of all

tarsi dark fuscous; mid tibia with an apical black spur.

Wing. Length 4.2 mm; cells bc and c pale fuscous; dense microtrichia

covering cell c only; remainder of wing colourless except fuscous cell sc,

narrow fuscous costal band confluent with R,; and remaining narrow around

apex of wing, a distinct large bulla across extension of cell bcu and pale

192 Australian Entomologist, 2016, 43 (4)

fuscous in area normally covered by anal streak; aggregation of microtrichia

around A,+Cu, less dense than in most species of Bactrocera; supernumerary

lobe weak.

Abdomen. Oval; tergites free; strong black pecten present on tergite III. All

tergites red-brown except for a narrow transverse black band across anterior

margin of tergite III and a moderately broad medial black vitta over anterior

1/2 of tergite V. No shining spots (ceromata) on tergite V. All abdominal

sternites red-brown; sternite V with a deep concavity on posterior margin.

Surstylus with posterior lobe short.

Female. As for male except no bulla on wing; pecten of cilia absent from

abdominal segment III; oviscape red-brown, dorsoventrally compressed and

tapering slightly in dorsal view; ratio of length of oviscape to length of tergite

V, 0.3:1; apex of aculeus needle shaped.

Distribution. Papua New Guinea (Madang, Morobe and Central Provinces

north of the Owen Stanley Range).

Host plants. Gnetum gnemon and Gnetum costatum. Bactrocera (Bulladacus)

species utilise Gnetum species as primary hosts across their geographic range,

except in Australia where Gnetum is absent; the record from Sandoricum

koetjape (Meliaceae) listed under material examined is possibly an error.

Comments. Bactrocera aceromata is similar to B. (Bulladacus) cinnabaria

Drew & Romig in possessing a red-brown scutum, mostly red-brown

abdominal tergites, wings without infuscation except for the costal band and

anal streak and femora entirely pale (fulvous to red-brown). It differs from B.

cinnabaria in having the costal band very narrow around the wing apex and

in lacking black lateral margins on abdominal tergites IV and V.

Bactrocera (Bulladacus) aenigmatica (Malloch)

Dacus aenigmaticus Malloch, 1931: 261 (9). Type locality: Malololelei, Upolu,

Western Samoa.

Bactrocera (Bulladacus) aenigmatica (Malloch): Drew and Hancock 1995: 9 ($9).

Description. See Drew (1989) and Drew and Hancock (1995).

Distribution. Western Samoa (Upolu and Savaii).

Host plant. Aglaia samoensis (Meliaceae) (Drew and Hancock 1995).

Comments. The lengths of the antennal segments, previously unrecorded, are

0.1 mm, 0.2 mm and 0.46 mm and the first flagellomere is apically rounded.

Bactrocera (Bulladacus) bullata Drew

Bactrocera (Bactrocera) bullata Drew, 1989: 124 ($9). Type locality: Abelam, East

Sepik district, Papua New Guinea.

Bactrocera (Bulladacus) bullata: Drew and Hancock 1995: 9.

Australian Entomologist, 2016, 43 (4) 193

Material examined. PAPUA NEW GUINEA: Madang Province — a large series of

specimens collected at Ohu, October 2000 and February to October 2001, recorded as

reared from Phaleria macrocarpa; a large series of specimens collected at Baitabag,

May to October 2001, recorded as reared from Phaleria macrocarpa; Morobe

Province — a large series of specimens, May 2000, reared from an unknown host at the

Lae Forest Research Institute. Specimens in QDAF.

Description. See Drew (1989).

Distribution. Papua New Guinea (East Sepik, Madang and Morobe

Provinces).

Host plants. Phaleria macrocarpa (Thymeliaceae) (Novotny et al. 2005 and

above records) and Gnetum gnemon (Gnetaceae) (Leblanc et al. 2012). A

record from Garcinia sp. (Clusiaceae) (Drew 1989) is probably an error.

Bactrocera (Bulladacus) bullifera (Hardy)

Dacus (Strumeta) bulliferus Hardy, 1973: 32 (i$). Type locality: Songkhla, Thailand.

Bactrocera (Bulladacus) bullifera: Drew and Hancock 1995: 9; Drew and Romig

2013 ($9).

Description. See Drew and Romig (2013).

Distribution. Southern Thailand, Peninsular Malaysia, East Malaysia

(Sarawak), Indonesia (Java, Sumatra).

Host plant. Gnetum gnemon (Gnetaceae) (Drew and Romig 2013).

Bactrocera (Bulladacus) captiva Drew & Romig

Bactrocera (Bulladacus) captiva Drew & Romig, 2013: 196 ($9). Type locality:

‘Philippines’.

Description. See Drew and Romig (2013).

Distribution. Philippines (precise locality unknown).

Host plant. Eugenia javanica (Myrtaceae) — possibly incorrect, the type

series and host record stemming from quarantine interception at Narita

Airport, Japan (Drew and Romig 2013).

Bactrocera (Bulladacus) cinnabaria Drew & Romig

Dacus (Strumeta) mcgregori: Hardy and Adachi 1954: 176 ($9). Singapore.

Misidentification.

Bactrocera (Bulladacus) cinnabaria Drew & Romig, 2013: 197 ($9). Type locality:

Arong, Car Nicobar, India.

Description. See Drew and Romig (2013).

Distribution. India (Andaman and Nicobar Is), West Malaysia and Singapore.

Host plants. Gnetum gnemon (Gnetaceae) (Hardy and Adachi 1954, Yong

1994) and Gnetum sp. (Drew and Romig 2013).

194 Australian Entomologist, 2016, 43 (4)

Comments. The abdominal pattern is a little variable, with the black medial

vitta on tergites IV-V often interrupted or absent on tergite IV. Records of B.

mcgregori from Singapore (Hardy 1973, 1974), West Malaysia (Yong 1994)

and the Andaman and Nicobar Islands (David and Ramani 2011) are regarded

as misidentifications of B. cinnabaria, with B. mcgregori differing in the

presence of a presutural yellow vitta and longer, more parallel-sided lateral

postsutural yellow vitta that extends anterior to the suture as a distinct spot.

Bactrocera (Bulladacus) diaphana (Hering)

(Fig. 2)

Strumeta diaphana Hering, 1953: 508 (9). Type locality: Bernhard Camp, Papua

Province, Indonesia.

Bactrocera (Bactrocera) diaphana: Drew 1989: 130.

Bactrocera (Bulladacus) obtrullata White & Evenhuis, 1999: 510 (i$). Type locality:

Dojo, Papua Province, Indonesia. Syn. n.

Material examined. PAPUA NEW GUINEA: Madang Province — 1 3, 2 99,

Baitabag, 17.1.2001, reared from Pimelodendron amboinicum; Morobe Province — 2

£, 4 99, Lae Forest Research Institute Botanical Gardens, 29.iii.2000, reared from

Sandoricum koetjape. Specimens in QDAF.

Description. Male. Head: Height 1.2 mm; frons length 1.4 times breadth, red-

brown with pale fuscous to fuscous centrally; setae black: 2 frontal, 1 orbital;

lunule fuscous; ocellar triangle black; vertex red-brown; face entirely fulvous

without dark markings, length 0.5 mm; gena fulvous without dark markings,

black seta present; occiput pale fuscous, fulvous along eye margins; occipital

row with 2-3 weak black setae. Antenna with all segments red-brown; length

of segments: 0.1 mm, 0.2 mm, 0.4 mm; first flagellomere rounded at apex.

Thorax. Scutum red-brown with two submedial black bands that are broad

posteriorly and narrowing anteriorly and joined across posterior area of

scutum with fuscous colouration, black on lateral margins between

postpronotal lobe and notopleuron and inside notopleuron. Yellow markings

as follows: postpronotal lobe; anepisternal stripe reaching to anterior

notopleural seta dorsally, continuing to katepisternum as a large spot, anterior

margin straight; anatergite (posterior apex black); anterior 2/3 of katatergite

(remainder black); broad parallel-sided lateral postsutural vitta beginning as a

large spot anterior to notopleural suture and ending at intra-alar seta; a lateral

yellow band between posterior margin of postpronotal lobe and yellow spot

anterior to notopleural suture; medial postsutural yellow vitta absent.

Postnotum red-brown centrally, black laterally. Notopleural lobe red-brown.

Scutellum yellow with a narrow black basal band that widens laterally. Setae:

2 scapular; 2 notopleural; 1 anepisternal; 1 supra-alar; 1 postalar; 1 intra-alar;

1 prescutellar acrostichal; 1 (apical) scutellar.

Legs. All leg segments entirely fulvous except hind tibiae dark fuscous

basally to fuscous apically; mid tibia with an apical black spur.

Australian Entomologist, 2016, 43 (4) 195

Fig. 2. Bactrocera (Bulladacus) diaphana (Hering), male.

Wing. Length 4.4 mm; cells bc and c dark fuscous and covered with dense

microtrichia; remainder of wing colourless except dark fuscous cell sc,

narrow fuscous costal band confluent with R>,; and continuing around costal

margin to apex of vein M, a pale tint in centre of wing, distinct large dark

bulla across extension of cell bcu and fuscous in area normally covered by

anal streak; a dense aggregation of microtrichia around Aj4*Cu;;

supernumerary lobe weak.

Abdomen. Oval; tergites free; a strong black pecten present on tergite III. All

tergites red-brown except for a narrow black band submedially on tergite

IHI, a narrow black band across anterior margin of tergite III which expands

over lateral margins, a medium width medial black vitta over tergite V and

fuscous to dark fuscous anterolateral corners of tergites IV and V. No shining

spots (ceromata) on tergite V. All abdominal sternites red-brown; sternite V

with a deep concavity on posterior margin. Surstylus with posterior lob short.

196 Australian Entomologist, 2016, 43 (4)

Female. As for male except face with a pair of medium-sized oval black

spots, notopleuron with anterior 1/2 fuscous and posterior 1/2 yellow, wing

with cells bc and c colourless and lacking a bulla over extension of cell bcu;

apex of aculeus needle shaped, ratio of length of oviscape to length of tergite

V, 1:1.

Distribution. Indonesia (Papua Province); Papua New Guinea (Madang and

Morobe Provinces).

Host plants. Recorded hosts are Pimelodendron amboinicum (Euphorbiaceae)

(Novotny et al. 2005 and above records) and Sandoricum koetjape

(Meliaceae). If these are correct plant identifications then they are a

significant departure from those of most other B. (Bulladacus) species.

Comments. The female holotype of B. diaphana was examined, described and

illustrated by Drew (1989). The rearing of both sexes from the same host fruit

has now made possible the correct subgeneric placement for this species and

the description of the male.

Bactrocera (Bulladacus) diaphana exhibits a low level of sexual

dimorphism. It is similar to B. (Bulladacus) mcgregori (Bezzi) in general

body and wing colour patterns and in possessing a lateral yellow vitta

between the postpronotal lobe and notopleural suture. It differs from B.

mcgregori in having apex of antennal first flagellomere rounded apically, a

narrower anepisternal stripe and face fulvous with a pair of black spots in

females. The original description of B. diaphana was based on a female

specimen and that of Bactrocera (Bulladacus) obtrullata White & Evenhuis

on male specimens. Now that we have studied both sexes reared from the

same host plant sample, we can confirm that B. obtrullata is a new synonym

of B. diaphana, being identical in all significant morphological characters.

Bactrocera (Bulladacus) eximia Drew

Bactrocera (Bactrocera) eximia Drew, 1989: 132 ($9). Type locality: Baku, Madang

Province, Papua New Guinea.

Bactrocera (Bulladacus) eximia: Drew and Hancock 1995: 9.

Description. See Drew (1989).

Distribution. Papua New Guinea (Madang and Central Provinces).

Host plants. Terminalia brassii and T. catappa (Combretaceae) (Drew 1989,

Leblanc et al. 2012).

Bactrocera (Bulladacus) flavinotus (May)

Afrodacus flavinotus May, 1957: 293 (9). Type locality: Atherton, Queensland.

Bactrocera (Bulladacus) neotigrina Drew & Hancock, 1999: 7 ($9), in Drew et al.

1999. Type locality: Helenvale nr Cooktown, Queensland. Syn. Drew and

Hancock 2000: 27.

Bactrocera (Bulladacus) flavinotus: Drew and Hancock 2000: 27.

Australian Entomologist, 2016, 43 (4) 197

Description. See May (1957) and Drew et al. (1999).

Distribution. Northern Queensland (Cooktown to Atherton and Gordonvale).

Host plant. Terminalia sericocarpa (Combretaceae) (Drew et al. 1999),

Bactrocera (Bulladacus) gnetum Drew & Hancock

Bactrocera (Bulladacus) gnetum Drew & Hancock, 1995: 9 ($9). Type locality:

Saivou, Vanua Levu, Fiji.

Description. See Drew and Hancock (1995).

Distribution. Fiji.

Host plant. Gnetum gnemon (Gnetaceae) (Drew and Hancock 1995).

Bactrocera (Bulladacus) mcgregori (Bezzi)

Chaetodacus mcgregori Bezzi, 1919: 426 ($9). Type locality: Batbatan Island,

Philippines.

Bactrocera (Bulladacus) mcgregori: Drew and Hancock 1995: 9.

Description. See Drew and Romig (2013).

Distribution. Philippines (Panay and Batbatan Is), Records from Singapore

(Hardy and Adachi 1954, Hardy 1973, 1974), West Malaysia (Yong 1994)

and the Andaman and Nicobar Islands (David and Ramani 2011) are

transferred to B. cinnabaria.

Host plant. Unknown. Previous records of Gnetum gnemon (Hardy and

Adachi 1954, Hardy 1973, 1974, Yong 1994) refer to B. cinnabaria.

Bactrocera (Bulladacus) pacificae Drew & Romig

Bactrocera (Bulladacus) pacificae Drew & Romig, 2001: 138 ($9). Type locality:

Vatina, Guadalcanal, Solomon Islands.

Material examined. PAPUA NEW GUINEA: Central Province — 14 $4, 14 99,

Hiritano Highway, Doa Forest, 22.x.1999, reared from Gnetum gnemon; 1 9, Rouna

Forest, 2.11.1999, reared from Gnetum gnemon, coll. Drew et al.; Madang Province —

1 ĝ, 2 99 (10.12001), 1 3 (111.2001), Ohu, reared from Gnetum gnemon; 1 9,

Baitabag, 25.x.2000, reared from Gnetum gnemon. Specimens in QDAF.

Description. See Drew and Romig (2001).

Distribution. Solomon Islands (Guadalcanal and Temotu Province (Lom Lom

and Nendo Islands) and Papua New Guinea (Central and Madang

Provinces).

Host plant. Gnetum gnemon (Gnetaceae) (Drew and Romig 2001, Leblanc et

al. 2012).

Comments. Bactrocera pacificae is distinct in possessing a red-brown

scutellum with lateral yellow margins. The above new records from Papua

New Guinea indicate a wider distribution than previously considered.

198 Australian Entomologist, 2016, 43 (4)

Bactrocera (Bulladacus) penefurva Drew

Bactrocera (Bactrocera) penefurva Drew, 1989: 151 (8). Type locality: 20 km SE

Port Moresby, Papua New Guinea.

Bactrocera (Bulladacus) penefurva: Drew and Hancock 1995: 9; Drew and Romig

2001 (£9).

Material examined. PAPUA NEW GUINEA: Madang Province — a large series of

specimens collected at Baitabag, January to March 2001, reared from Terminalia sp.;

Morobe Province — a large series of specimens collected at Bundun Conference

Centre Lae, Lae Forest Research Institute Botanical Gardens and Oomsis Forest,

February to May 2000, reared from Terminalia sp. Specimens in QDAF.

Description. This species has been described and illustrated by Drew (1989)

and Drew and Romig (2001). One additional character is that, in both sexes,

the apex of the antennal first flagellomere is rounded, not truncate as in some

B. (Bulladacus) species.

Distribution. Papua New Guinea (Central, Morobe and Madang Provinces)

and Solomon Islands (Guadalcanal).

Host plants. Terminalia catappa and T. kaernbachii (Combretaceae) (Drew

and Romig 2001). A record from Gnetum gnemon (Gnetaceae) (Leblanc et al.

2012, Hollingsworth et al. 2003: Table 5) appears to be based on a lapsus for

B. pacificae.

Bactrocera (Bulladacus) peterseni (Hardy)

Dacus (Strumeta) peterseni Hardy, 1970: 75 (8). Type locality: Tarawakan, Tawi

Tawi, Philippines.

Bactrocera (Bulladacus) peterseni: Drew and Hancock 1995: 9.

Description. See Drew and Romig (2013).

Distribution. Philippines (Tawi Tawi I.).

Host plant. Unknown.

Bactrocera (Bulladacus) tigrina (May)

Afrodacus tigrinus May, 1952: 335 (9); 1957: 296 (3). Type locality: Kamerunga,

Cairns, Queensland.

Afrodacus furvus May, 1957: 294 ($9). Type locality: Atherton, Queensland. Syn.

Drew 1989: 24.

Bactrocera (Bulladacus) tigrina: Drew and Hancock 1995: 9.

Description. See May (1952, 1957), Drew (1989) and Drew et al. (1999).

Distribution. Northern Queensland (Iron Range to Murray Falls near Tully).

Host plants. Terminalia sericocarpa and T. muelleri (Combretaceae) (Drew

et al. 1999).

Australian Entomologist, 2016, 43 (4) 199

Bactrocera (Bulladacus) trilobata sp. n.

(Fig. 3)

Type material. Holotype 6, PAPUA NEW GUINEA: Madang Province, Mis Village,

5°11'S 145°47'E, 25.vii1.2008, coll. Ctvrtecka, Brus & Rimandai, reared from Phaleria

macrocarpa. Paratypes: | 4, 5 99, same data as holotype. Holotype and 1 paratype

in QM (Reg. Nos T234939 (HT) and T234940 (PT)); 5 paratypes in QDAF.

Description. Male. Head: Height 1.1 mm; frons length 1.6 times breadth, red-

brown without dark markings; orbital setae black: 2 frontal, 1 orbital; lunule

red-brown; ocellar triangle black; vertex red-brown.; face entirely fulvous

without dark markings, length 0.4 mm; gena fulvous without dark markings,

weak pale seta present; occiput red-brown, fulvous along eye margins,

occipital row with 1-3 dark setae. Antenna with all segments red-brown;

length of segments: 0.1 mm, 0.15 mm, 0.5 mm; first flagellomere rounded at

apex.

Thorax. Scutum red-brown without dark markings. Yellow markings as

follows: postpronotal lobe; anepisternal stripe reaching postpronotal lobe

dorsally, continuing to katepisternum as a transverse spot, anterior margin

straight; anatergite (posterior apex red-brown); anterior 1/2 of katatergite

(remainder red-brown); narrow lateral postsutural vitta narrowing slightly

posteriorly to end just before intra-alar seta; no distinct spot anterior to

notopleural suture; medial postsutural yellow vitta absent. Postnotum red-

brown centrally, black laterally. Notopleural lobe red-brown. Scutellum

yellow with a narrow black basal band. Setae: 2 scapular; 2 notopleural; 1

anepisternal; 1 supra-alar; 1 postalar; 1 intra-alar; 1 prescutellar acrostichal; 1

(apical) scutellar.

Legs. All leg segments entirely fulvous except apical three segments of all

tarsi red-brown; mid tibia with an apical black spur.

Wing. Length 4.4 mm; cells bc and c pale fuscous with dense microtrichia in

outer corner of cell c only; remainder of wing colourless except fuscous cell

sc., narrow fuscous costal band confluent with R>,; and remaining narrow

around wing margin to end at apex of vein M, narrow fuscous transverse

band enclosing DM-Cu crossvein, broad fuscous anal streak, small fuscous

bulla around apex of cell bcu extension. No dense aggregation of microtrichia

around A 4Cu». Supernumerary lobe weak.

Abdomen. Oval; tergites free; a strong pecten present on tergite III. All

tergites red-brown except for a short, medial, narrow black vitta on posterior

margin of tergite V. No shining spots (ceromata) on tergite V.

Female. As for male except bulla on wing and pecten of cilia on abdominal

tergite III both absent; oviscape red-brown, dorsoventrally compressed and

tapering slightly in dorsal view; ratio of length of oviscape to length of tergite

V, 1:1, apex of aculeus trilobed (Fig. 3).

200 Australian Entomologist, 2016, 43 (4)

Etymology. Named as an adjective after the trilobed aculeus.

Distribution. Papua New Guinea (Madang Province).

Host plant. Phaleria macrocarpa (Thymelaeaceae).

Comments. Bactrocera (Bulladacus) trilobata sp. n. is similar to B.

(Bulladacus) aenigmatica (Malloch), B. (Bulladacus) cinnabaria Drew &

Romig and B. (Bulladacus) pacificae Drew & Romig in possessing a red-

brown scutum, a narrow costal band confluent with R>,3, lateral postsutural

yellow vitta short and narrow and ending before the intra-alar seta and

additionally to B. cinnabaria in having a broad anepisternal stripe reaching to

the postpronotal lobe. It differs from all three species in having the DM-Cu

crossvein enclosed with fuscous colouration and the apex of the aculeus

trilobed.

wet

Fig. 3. Bactrocera (Bulladacus) trilobata sp. n., male and female aculeus.

Australian Entomologist, 2016, 43 (4) 201

Bactrocera (Bulladacus) unipunctata (Malloch)

Dacus unipunctatus Malloch, 1939: 245 (4). Type locality Tulagi, Florida I.,

Solomon Islands.

Bactrocera (Bactrocera) unipunctata: Drew 1989: 167.

Description. See Drew (1974, 1989) and Drew and Romig (2001).

Distribution. Solomon Islands (Florida I.).

Host plant. Unknown.

Comments. The short antennal segments (0.08, 0.18, 0.5 mm), short wing cell

bcu extension and lack of ceromata on abdominal tergite V place this species

in Bulladacus and it is therefore transferred from subgenus Bactrocera. It

also has an oval abdomen with a single black medial spot on tergite V and

microtrichia over much of cells bc and c, typical of many Bulladacus species.

The pecten of cilia and bulla appear to be absent but the only known

specimen is damaged and teneral and both structures are possibly present.

Bactrocera (Bulladacus) wanangiae sp. n.

(Fig. 4)

Bactrocera (Bulladacus) sp. near aceraglans White & Evenhuis, 1999: 507 (d).

Wanuma, Madang Province, Papua New Guinea.

Type material. Holotype 3, PAPUA NEW GUINEA: Morobe Province, Wanang,

4.xii.2008, Ctvrtecka et al., reared ex unidentified fruit. Paratype 3, same data as

holotype. Types in QM (Reg. Nos T234941 (HT) and T234942 (PT)).

Description. Male. Head: Height 1.2 mm; frons length 1.8 times breadth, red-

brown with dark fuscous on anteromedial hump; orbital setae black: 2 frontal,

1 orbital; lunule red-brown; ocellar triangle black; vertex red-brown; face

entirely fulvous, length 0.5 mm; gena red-brown; occiput red-brown, fulvous

along eye margins, occipital row with 4-5 strong pale setae. Antenna abraded.

Thorax. Scutum black with red-brown bordering postpronotal lobe, around

notopleural suture and below and behind lateral postsutural vitta. Yellow

markings as follows: postpronotal lobe; anepisternal stripe reaching

postpronotal lobe dorsally, continuing to katepisternum as a small spot,

anterior margin straight; anatergite (posterior apex black); anterior 2/3 of

katatergite (remainder black); broad parallel-sided lateral postsutural vitta

beginning as a small spot anterior to notopleural suture and ending just

behind intra-alar seta; medial postsutural yellow vitta absent. No lateral

yellow area between postpronotal and notopleural lobes. Pleural areas dark

fuscous to black. Postnotum red-brown centrally, black laterally. Notopleural

lobe yellow. Scutellum yellow with a narrow black basal band. Setae: 2

scapular; 2 notopleural; 1 anepisternal; 1 supra-alar; 1 postalar; 1 intra-alar; 1

prescutellar acrostichal; 1 (apical) scutellar.

Legs. All segments entirely fulvous.

202 Australian Entomologist, 2016, 43 (4)

Wing. Length 5.4 mm; cells bc and c pale fuscous with dense microtrichia

over all of cell c and outer half of cell bc; remainder of wing colourless

except fuscous cell sc., narrow fuscous costal band confluent with R>,; and

remaining narrow around wing margin to end between apices of R4,5 and M,

narrow infuscation enclosing R-M and DM-Cu crossveins that is also

connected with infuscation along M, broad pale fuscous anal streak, distinct

red-brown bulla around apex of cell bcu extension. No dense aggregation of

microtrichia around A,+CuA,. Supernumerary lobe of medium development.

Abdomen. Abdomen oval; tergites free; a weak pale pecten present on tergite

III. All tergites red-brown except for a narrow medial black vitta over tergite

V. No shining spots (ceromata) on tergite V.

Fig. 4. Bactrocera (Bulladacus) wanangiae sp. n., male.

Australian Entomologist, 2016, 43 (4) 203

Female. Unknown.

Etymology. Named after the type locality.

Distribution. Papua New Guinea (Madang and Morobe Provinces).

Host plant. Unknown.

Comments. Bactrocera (Bulladacus) wanangiae sp. n. is similar to B.

(Bulladacus) gnetum Drew & Hancock in possessing a black scutum without

a medial postsutural yellow vitta, wing with a narrow fuscous costal band and

fulvous leg segments. It differs from B. gnetum in having a broad parallel-

sided lateral postsutural yellow vitta and infuscation enclosing R-M and DM-

Cu crossveins that is connected by infuscation along vein M. The ‘B. sp nr

aceraglans’ of White and Evenhuis (1999) is associated based on its original

description and illustration.

Bactrocera (Bulladacus) warisensis White & Evenhuis

Bactrocera (Bulladacus) warisensis White & Evenhuis, 1999: 512 (3). Type locality:

Waris, S of Jayapura, Papua Province, Indonesia.

Description. See White and Evenhuis (1999).

Distribution. Indonesia (Papua Province).

Host plant. Unknown.

Key to species of subgenus Bulladacus

] Scutum with a medial postsutural yellow vitta ....................eeeesseues 2

— Scutum without a medial postsutural yellow vitta ...................eesssss 7

2 Scutum red-brown or with a pair of submedial black vittae; male with

bulla either small and distinct or weak and undeveloped ................... 3

— Scutum mostly black; male with bulla distinct and well developed ....... 4

3 Scutum with a pair of dark submedial vittae; lateral postsutural yellow

vitta extending anterior to suture as a distinct spot; medial postsutural

yellow vitta narrowly triangular and almost as long as lateral vitta; wing

with a dark band over DM-Cu crossvein; male with a small distinct bulla

over apex of cell bcu extension (Papua New Guinea) ............... B. bullata

— Scutum without a pair of dark submedial vittae; lateral postsutural yellow

vitta not extending anterior to suture; medial postsutural yellow vitta

broadly triangular and distinctly shorter than lateral vitta; wing without a

dark band over DM-Cu crossvein; male with bulla weak and undeveloped

CPSU EAD, T Seu uteri <n ee de E Rl X ay eas Pata Pe p B. flavinotus

4 Sexes similar, the costal band narrow throughout and not expanded into a

broad, isolated apical patch in males; abdomen orange-brown with narrow

black lateral margins on tergites III-IV and a black medial vitta on tergite

V (Papua New Guinea) ............. eeeeeeeeeeeeeeeeenmmm D. eximia

204 Australian Entomologist, 2016, 43 (4)

— Sexes dissimilar, the costal band narrow in females, expanded into a

broad, isolated apical patch in males; abdomen with black areas as

sublateral spots or patches on tergites III-V (males) or with black

transverse bands on tergite III (and often I+II) and black lateral margins

on teretes II-V (females) Lesen rre eee pb ones ner pee Pt erret 5

5 Medial postsutural vitta reaching line of suture anteriorly; abdomen

without a dark transverse band or sublateral patches on tergite I--II; male

wing with apical patch crossing vein M and reaching almost to DM-Cu

crossvein (Papua New Guinea and Solomon Islands) ......... B. penefurva

— Medial postsutural vitta not reaching line of suture anteriorly; abdomen

with a dark transverse band or sublateral patches on tergite I-II; male

wing with apical patch as above or smaller and not crossing vein M ...... 6

6 Medial postsutural vitta broadly triangular and distinctly shorter than

lateral vitta; male wing with apical patch not crossing vein M and

approaching it well distad of DM-Cu crossvein (Australia) ...... B. tigrina

— Medial postsutural vitta broadly oval and almost as long as lateral vitta;

male wing with apical patch crossing vein M and reaching almost to DM-

Cu crossvein (Philippines) .............. sess B. captiva

7 Costal band broadly expanded medially with a triangular dark band across

both R-M and DM-Cu crossveins; male bulla present over apex of cell

bcu extension (Papua Province, eastern Indonesia) ........... D. warisensis

— Costal band narrow or expanded apically but not expanded across R-M

and DM-Cu crossveins; male bulla variably placed or absent .............. 8

8 Scutum black except for lateral postsutural yellow vitta; costal band very

narrow and of uniform width beyond apex of vein R24 .................ssess 9

— Scutum largely fulvous to red-brown or with dark submedial and

sometimes medial vittae; costal band narrow or broadened apically ..... 11

9 Scutum with lateral postsutural yellow vitta short and sharply tapered

posteriorly; abdomen with a broad black lateral vitta on tergites III-V and

a broad medial vitta from posterior margin of tergite III to tergite V; male

with bulla large and ovate (Fiji) ............ceeesssseecceeeesesseneeeeeeseeeees B. gnetum

— Scutum with lateral postsutural yellow vitta elongate and parallel-sided,

enclosing intra-alar seta; abdomen orange-brown, at most with a black

medial vitta on tergite V; male bulla indistinct or absent ..................... 10

10 Wing with a dark posterior stripe over DM-Cu crossvein to wing margin;

fore and apical halves of mid and hind femora fuscous; male with a patch

of long cilia in place of bulla (Papua New Guinea) ............. B. aceraglans

— Wing with a narrow infuscation over R-M and DM-Cu crossveins and

intermediate portion of vein M; femora fulvous; male with bulla small

and indistinct (Papua New Guinea) ........................... B. wanangiae sp. n.

Australian Entomologist, 2016, 43 (4) 205

Apical third of mid femur black or black-spotted, mid tibia at least partly

and hind tibia entirely dark fuscous to black; costal band broadened or

expanded apically; male bulla blackened; scutum with lateral postsutural

yellow vitta extending anterior to suture as a distinct spot; abdomen with

medial black vitta across tergites III-V connected anteriorly with a broad

transverse band across tergite III; anepisternal yellow stripe narrow, not

reaching postpronotal lobe anteriorly ..........ssessssessssresssssreeresesssrereessssee 12

All femora and tibiae fulvous except hind tibia often fuscous; costal band

narrow, not or only weakly expanded apically; male bulla often not

blackened; scutum with lateral postsutural yellow vitta often not

extending anterior to suture as a distinct spot; abdomen with medial black

vitta generally confined to tergites IV-V or V, seldom entering or crossing

tergite III but not connected with a transverse band across it; anepisternal

yellow stripe often broad and reaching postpronotal lobe anteriorly ..... 13

12 Apical third of mid femur, most of mid and all of hind tibiae dark fuscous

to black; costal band broadened apically and reaching vein M; scutum

with the dark medial and submedial vittae not broadly connected

posteriorly; a narrow presutural yellow vitta present between postpronotal

lobe and sutural spot (southern Philippines: Tawi Tawi) ...... B. peterseni

Apical third of mid femur with a black spot, basal third of mid and all of

hind tibiae dark fuscous to black; costal band expanded into a distinct

apical patch; scutum with the dark medial and submedial vittae broadly

connected posteriorly; presutural yellow vitta absent (Southern Thailand,

Peninsular Malaysia, Borneo, Sumatra and Java) ............... B. bullifera

Scutum with lateral postsutural yellow vitta normally extending anterior

to suture as a distinct spot; presutural lateral yellow vitta present; lateral

postsutural yellow vitta parallel-sided and enclosing intra-alar seta ...... 14

Scutum with lateral postsutural yellow vitta not extending anterior to

suture as a distinct spot; presutural lateral yellow vitta absent, if present

then reduced to a small yellowish patch immediately behind postpronotal

lobe; lateral postsutural yellow vitta short and not enclosing intra-alar seta

Pesca OEA AE IE NE Pea eee ba een ean EORR IC die LER HONE OL Ks ny BERD AA Keres 16

14 Anepisternal yellow stripe narrow, only reaching anterior notopleural seta

anteriorly; scutum with a broad black submedial vitta; presutural lateral

yellow vitta broadly connecting presutural spot with postpronotal lobe;

costal band pale and indistinct; male wing with a pale fuscous patch

across vein M between R-M and DM-Cu crossveins (Papua New Guinea

and Papua Province, eastern Indonesia) ................... sss B. diaphana

Anepisternal yellow stripe broad, reaching postpronotal lobe anteriorly;

scutum with or without a narrow black submedial vitta; presutural lateral

yellow vitta not broadly connecting presutural spot with postpronotal

lobe; costal band distinct; wing without a pale fuscous patch across vein

206 Australian Entomologist, 2016, 43 (4)

15 Scutum with narrow dark submedial vitta and a narrow presutural lateral

yellow vitta connecting or almost connecting postpronotal lobe with

sutural spot; costal band reaching vein M (Philippines) ........ B. mcgregori

— Scutum without dark submedial vitta and with a broad presutural lateral

yellow or orange vitta between postpronotal lobe and notopleuron; costal

band not reaching vein M (Papua New Guinea) .................. B. aceromata

16 Anepisternal yellow stripe broad, reaching postpronotal lobe anteriorly;

scutum red-brown; scutellum yellow with a narrow dark basal band ... 17

— Anepisternal yellow stripe narrow, not reaching postpronotal lobe

anteriorly; scutum and scutellum largely orange-brown ....................... 19

17 Abdomen with black lateral markings; antennal first flagellomere apically

truncate; male bulla large and ovate (Andaman and Nicobar Is, West

Malaysia and Singapore) ........... 0. ccc ecceceeeeeeeeeeessetteeeeeeeeeess B. cinniabaria

— Abdomen without black lateral markings; antennal first flagellomere

apically rounded; male bulla small and rounded or possibly absent ..... 18

18 Scutum with lateral postsutural yellow vitta sharply tapered posteriorly;

notopleural lobe yellow; crossvein DM-Cu not enclosed by a narrow

transverse fuscous band; small facial spots present; male bulla indistinct

or absent; pecten of cilia on tergite III in males weak or absent; female

unknown (Solomon Islands: Florida I.) ............................... B. unipunctata

— Scutum with lateral postsutural yellow vitta slightly tapered posteriorly;

notopleural lobe red-brown; crossvein DM-Cu enclosed by a narrow

transverse fuscous band; facial spots absent; male bulla distinct; pecten of

cilia on tergite III in males well developed; female aculeus apically

trilobed (Papua New Guinea) ................. sss B. trilobata Sp. n.

19 Scutum with a short, medial black vitta across line of suture; lateral

postsutural yellow vitta sharply tapered posteriorly; scutellum with a

narrow black lateral vitta; costal band not reaching vein M; male bulla

small and round and placed on vein Aj4-Cu» beyond apex of very narrow

cell bcu extension; abdomen with a narrow, interrupted black medial vitta

on tergites I-V and broader sublateral markings on tergites III-IV (males)

or with a distinct black medial vitta and broad lateral margins on tergites

III-V (females) (Western Samoa) ..................... eee B. aenigmatica

— Scutum without a medial black vitta; lateral postsutural yellow vitta

narrow but not distinctly tapered; scutellum yellow basolaterally; costal

band reaching vein M; male bulla large and ovate and cell bcu extension

vestigial; abdomen without lateral black bands and medial vitta confined

to tergite V (Papua New Guinea and Solomon Islands, including Nendo)

CUARTA baa AAD nah AAR nl CHER ARA LOR scald UCET REINO TERRAE B. pacificae

Discussion

Although not all Bulladacus species show all the diagnostic characters, their

overall similarity suggests that the subgenus is monophyletic. The lack of

Australian Entomologist, 2016, 43 (4) 207

shining spots (ceromata) on abdominal tergite V and the presence of either a

bulla or patch of cilia at or near the apex of wing cell bcu extension appear to

be synapomorphies for the subgenus. Bactrocera aceraglans from Papua

New Guinea and possibly B. unipunctata from Solomon Islands lack both the

bulla and abdominal pecten in males but otherwise appear to belong here.

The geographical distribution of the 20 known Bulladacus species is shown

in Table 1, placed within the six zones recognised by Hancock and Drew

(2015). Two species (B. penefurva and B. pacificae) occur in both Papua

New Guinea (Zone D) and the Solomon Islands (Zone F); all other species

are endemic to their particular zones. The lack of Bulladacus species from

Wallacea (Zone C) likely reflects undercollecting rather than a genuine

absence.

Table 1. Distribution of species in genus Bactrocera and subgenus Bulladacus in each

biogeographic zone and percent endemism in Bulladacus. For a map of zones A-F see

Hancock and Drew (2015).

Biogeographic Zone No. species of No. species of % Endemic

Bactrocera Bulladacus Bulladacus

(A) Indian subcontinent 75 0 —

(B) South-East Asia 225 5 100

(D) New Guinea 170 10 80

(E) Australia 76 2 100

(F) South Pacific 59 5 60

The subgenus appears to be closely associated with Gnetum gnemon, a

tropical shrub/tree distributed from NE India throughout much of SE Asia

and Australasia (excluding mainland Australia) as far east as Fiji and Samoa.

Some species, particularly the Australian B. tigrina and B. flavinotus, have

become adapted to Terminalia species, while the Samoan B. aenigmatica

utilises Aglaia samoensis. Although Bulladacus species might be expected to

occur throughout the range of Gnetum species, so far they are only known as

far west as southern peninsular Thailand and the Andaman Islands. Greatest

diversity exists on the island of New Guinea, where ten species are known,

suggesting that the subgenus arose there before dispersing westwards into SE

Asia and eastwards into the South Pacific.

The three strongly sexually dimorphic species, B. tigrina, B. penefurva and B.

captiva, occur in NE Australia, Papua New Guinea-Solomon Islands and the

Philippines respectively, with the latter two species appearing to be the most

closely allied morphologically. A similar faunal relationship between New

Guinea and the Philippines exists in the closely related acanthonevrine genera

Copiolepis Enderlein and Piocolepis Hancock (Hancock 2014). Of the

208 Australian Entomologist, 2016, 43 (4)

remaining SE Asian species, B. bullifera (Sundaland) and B. peterseni (Tawi

Tawi) appear to be an allopatric species pair: both have dark areas on the mid

femur and mid and hind tibiae, plus three dark scutal vittae, a complete dark

medial vitta on abdominal tergites III-V, a narrow anepisternal yellow stripe

that does not reach the postpronotal lobe, a broad apex to the costal band and

an oval or rounded, distinctly blackened bulla. Likewise, B. cinnabaria

(Sundaland) and B. mcgregori (Philippines) also appear to be an allopatric

species pair: both are generally pale species with a broad anepisternal yellow

stripe, a paler, ovate bulla and a short, apically truncate antennal first

flagellomere especially in males. The presence or absence of a presutural

lateral yellow vitta thus appears to be homoplasious, particularly since its

alignment, when present, differs in the various species showing it.

In New Guinea, B. aceraglans (Western Highlands) stands alone, having

largely fuscous femora, the bulla replaced by a patch of cilia and the pecten

absent. Of the others, B. wanangiae (Madang and Morobe Provinces) shares

with B. aceraglans the black scutum, while B. diaphana (West Sepik and

Indonesian Papua) + B. aceromata (Central Province), both with a presutural

lateral yellow vitta, and B. bullata (East Sepik) + B. warisensis (Indonesian

Papua), both with the postsutural lateral yellow vitta extending anterior to the

suture as a distinct spot, a black submedial scutal vitta, a dark wing band over

at least DM-Cu crossvein, a small bulla at the apex of a relatively elongate

cell bcu in males and an almost unpatterned abdomen, appear to be related

pairs. The latter five species all have fulvous femora and all six species have

a parallel-sided lateral postsutural yellow vitta that encloses the intra-alar

seta. The medial postsutural yellow vitta in B. bullata is narrower than in

other species showing this character and is presumably homoplasious,

particularly since the other species lack the presutural spots.

The new species B. trilobata (Madang) has the lateral postsutural yellow vitta

ending before the intra-alar seta and not extending anterior to the notopleural

suture; it also has a red-brown scutum and fulvous femora and appears to be

closest to B. unipunctata from the Solomon Islands. The Terminalia host

plants, overall appearance, very short antennal first flagellomere (0.4 mm)

and well developed supernumerary lobe of the male wing in B. eximia

suggest a relationship with the figrina-penefurva-captiva series.

The three South Pacific species B. aenigmatica (Samoa), B. gnetum (Fiji) and

B. pacificae (Solomon Islands and Papua New Guinea) all have a short lateral

postsutural yellow vitta and likely form a related series that possibly includes

B. unipunctata and B. trilobata. In B. gnetun the scutum is black, the

scutellum yellow, the anepisternal yellow stripe reaches the postpronotal lobe

and the hind tibia is weakly fuscous. In B. aenigmatica and B. pacificae the

scutum and scutellum are largely orange-brown, the anepisternal yellow

stripe does not reach the postpronotal lobe and the hind tibia is dark fuscous,

suggesting another allopatric species pair despite the differences in the bulla.

Australian Entomologist, 2016, 43 (4) 209

A similar faunal relationship between Fiji, Solomon Islands (particularly

Nendo) and Western Samoa is seen in the adramine Coelotrypes punctilabris

(Bezzi) (Hancock and Drew 2005).

The host plant switch from Gnetum to Aglaia in B. aenigmatica suggests that

it reached Samoa prior to the arrival of Gnetum. The occurrence of an

extended abdominal black vitta in both western and eastern outlying species

suggests that this character, widespread in many Bactrocera species, is

homoplasious and that the short vitta on tergites IV-V or V only, seen in most

Bulladacus species, is the plesiomorphic state for the subgenus.

Relationships of the Australian B. flavinotus, with its red-brown scutum,

rudimentary bulla and relatively weak supernumerary lobe, remain uncertain

but its Terminalia host plant, very short antennal first flagellomere (0.4-0.44

mm) and broad medial postsutural vitta suggest a plesiomorphic relationship

with the eximia-captiva series. As with B. aenigmatica, the host plant switch

likely reflects the absence of Gnetum from Australia and suggests that the

flavinotus-captiva series originated there. A more detailed analysis of

phylogenertic relationships will be possible after all subgenera have been

reviewed and the polarity of character states more accurately determined.

Acknowledgements

We thank Meredith Romig for the curation of specimens and Michelle Baker

for the illustrations.

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Australian Entomologist, 2016, 43 (4): 211-234 211

THE STATUS OF YOMA ALGINA (BOISDUVAL, 1832) AND Y.

SABINA (CRAMER, 1780) (LEPIDOPTERA: NYMPHALIDAE:

NYMPHALINAE) IN AUSTRALIA

TREVOR A. LAMBKIN! and ROSS KENDALL?

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

?17 Eldon Street, Indooroopilly, Old 4068

(Email: trevor.lambkin @ uqconnect.edu.au)

Abstract

Yoma Doherty, 1886 (Nymphalinae: Junoniini) comprises two species, Y. algina (Boisduval,

1832) and Y. sabina (Cramer, 1780), which we confirm in this study by using larval morphology

and colour pattern, pupal morphology, adult morphology (i.e. wing pattern elements, wing

colour, fore and hindwing shapes, male genitalia) and host plant specialisation (Y. algina

restricted to Hemigraphis spp: Acanthaceae). In total, we determine at least 22 phenotypic

character state differences between the two species and, in addition, confirm that both species

occur in Queensland. In Australia, Y. sabina occurs in the Top End of the Northern Territory,

throughout Torres Strait and from eastern Queensland from Cape York to Townsville, while Y.

algina is now confirmed to naturally occur in Queensland, currently known only on the east

coast of Cape York Peninsula from Somerset at Cape York to Peach Creek, 25 km NNE of Coen.

The subspecific status of Australian populations of Y. algina, tentatively assigned to Y. a. netonia

Fruhstorfer, 1912, will remain in doubt until more Australian material is collected. Butterfly

house populations of Y. algina in eastern Australia have existed since the late 1980s but were

confused with Y. sabina for many years. These populations are thought to originate from a single

female collected at Iron Range, Cape York Peninsula. Thus, the existence of Y. algina in

Queensland has led to confusion in regards to the life history of Y. sabina. A review of Y. s.

sabina from Papua New Guinea and Y. s. parva (Butler, 1876) from mainland Australia,

including Torres Strait, indicates that the two taxa do not differ phenotypically. In addition, the

original description of Y. s. parva was found to be based on a diminutive 'dry season form',

which is atypical of normal Y. sabina from Australia. Thus, because of the phenotypic continuity

of Y. sabina through Papua New Guinea into Australia and confusion with the nomenclature of

the Australian population caused by Butler, we regard Y. s. parva syn. nov. as a junior synonym

of Y. s. sabina. In addition, the diminutive ‘dry season form’ of Y. s. sabina occurring in Papua

New Guinea and Australia is here proposed as form parva stat. rev.

Introduction

The genus Yoma Doherty, [1886] occurs widely in the Oriental and

Australian Regions (d'Abrera 1978, Parsons 1998). Yoma sabina (Cramer,

[1780]) is a lowland butterfly (Igarashi 1985), which has a broad distribution

from Myanmar to Hong Kong in southern China, Taiwan, the Philippines,

through Thailand to Indonesia, the Moluccas, mainland New Guinea and,

within Australia, from Torres Strait into tropical mainland Queensland and

the Northern Territory (Bingham 1905, Tsukada 1985, Parsons 1998,

Bascombe ef al. 1999, Braby 2000, Ek-Amnuay 2006). The type locality of

Y. sabina is Ambon Island in the Moluccas, Indonesia (Edwards et al. 2001,

Parsons 1998).

Yoma algina (Boisduval, 1832), which is considered to be a second species

(d'Abrera 1978, Parsons 1998), in contrast is much more restricted in its

distribution, occurring predominantly in the New Guinea region from Waigeo

212 Australian Entomologist, 2016, 43 (4)

Island through to the Solomon Archipelagos and Vanuatu (Parsons 1998,

Tennent 2002). The type locality of Y. algina is ‘New Guinea’ (Parsons

1998) but is thought to be more precisely Waigeo Island in West Papua

Province, Indonesia (Fruhstorfer 1912-15).

All of the recent major works on Australian butterflies (Common and

Waterhouse 1972, 1981, Braby 2000, Orr and Kitching 2010) have the genus

as monotypic and consisting of just one species, Y. sabina, with Y. algina a

subspecies of Y. sabina occurring in New Guinea. Fruhstorfer (1912-1915),

Barrett and Burns (1951), d'Abrera (1978) and Tennent (2002), however,

considered the two taxa to be specifically distinct. This distinction was

supported by Parsons (1998), who reported collecting both taxa in Papua

New Guinea flying together at several locations, including Port Moresby.

Thus he considered the two taxa to be separate species and sympatric in their

occurrence, at least in some areas.

Both taxa are highly variable in their underside wing markings. Thus, Parsons

(1998) suggested eight subspecies of Y. algina in Papua New Guinea, with Y.

a. netonia Fruhstorfer, 1912 occurring in southern Papua New Guinea and Y.

s. sabina occurring in New Guinea and the Moluccas (Tsukada 1985, Parsons

1998). In Australia, Y. s. parva (Butler, 1876) occurs in the Wet Tropics of

Queensland, on Cape York Peninsula, on several Torres Strait islands and in

localised areas in the Northern Territory (Braby 2000).

On mainland Queensland, Y. s. parva can be generally uncommon in the

southern end of its range in the Wet Tropics but, further north, it is observed

more frequently: at Cooktown (Valentine 1988) and particularly in Torres

Strait (Braby 2000, T.A. Lambkin unpublished data), especially on Thursday

Island (Valentine 1988). The species frequents lowland rainforest, open forest

bordering rainforest (G. Sankowsky pers. comm.) and monsoon forest (Braby

2000). In addition, it is found in swampy areas (Valentine 1988) and in

upland rainforests of the Atherton Tableland (Olive 1978). Overall, the

species is mostly observed during or just after the wet season. Adult males fly

along sunlit tracks defending territories and perch (Valentine 1988) with

wings outspread, normally within 2-5 m from the ground (Braby 2016).

Females of Y. sabina are often observed flying low over the forest floor in

search of oviposition sites on or near low-growing herbaceous host plants

(Saguru and Haruo 2000, Braby 2016). Adults are also found within the

forest settled on the underside of foliage and, in these situations, are often

difficult to detect due to their cryptic underside colouring. In addition, Y.

sabina 1s known to aestivate in large numbers throughout the dry season in

Cape York Peninsula and Townsville, clustering in dense vines, caves and

old buildings (Sankowsky 2015, Braby 2016, P.S. Valentine pers. comm.).

Parsons (1998) and Tennent (2002) noted that adults of Y. algina frequently

perch along sunlit tracks and clearings in rainforest habitats.

Australian Entomologist, 2016, 43 (4) 213

The life history of Y. s. sabina from Papua New Guinea was described by

Szent-Ivany and Carver (1967) and Parsons (1998). From the Moluccas,

Saguru and Haruo (2000) illustrated the egg and final instar of Y. s. sabina. In

Australia, the life history of what was believed at the time to be Y. s. parva

but found here to be that of Y. algina was described from material collected at

Iron Range, Cape York Peninsula in Queensland (Wood 1987a).

Saguru and Haruo (2000) reported observing a female of Y. s. sabina

Ovipositing on the ground (in the Moluccas) and the egg was successfully

reared to an adult on Gendarussa vulgaris Nees. (a synonym of Justicia

gendarussa L.) (Acanthaceae). Additionally, Saguru and Haruo (2000)

observed females of Y. s. podium Tsukada, 1985 in Taiwan, ovipositing on

Blechum pyramidatum (Laam.) Urb. (Acanthaceae). Sankowsky (2015)

added that larvae of Y. s. parva usually rest on the ground when not feeding.

There is little published on the early stages of Y. algina apart from a

photograph of a larva, indicated to be instar 3, in Parsons (1998).

In Australia, Wood (19872) successfully reared larvae of what he thought at

the time to be Y. s. parva on what he believed to be a Ruellia sp. (now syn. of

Dipteracanthus Nees) (Acanthaceae). Later, Sankowsky (1991) confirmed D.

bracteatus (R.Br) (Acanthaceae) as a larval host plant for Y. s. parva at Iron

Range.

In Papua New Guinea, Szent-Ivany and Carver (1967) reared Y. s. sabina on

Hemigraphis reptans (G.Forst.) T. Anderson ex Hemsl. (Acanthaceae), while

Parsons (1998) provided a detailed description of the early stages of Y. s.

sabina but did not mention a larval host plant.

Yoma s. parva, until recently, was rarely encountered in the Wet Tropics of

Queensland but of late it has been observed more frequently due to the

widespread increase of its host plant (Dipteracanthus spp) in the area (J.

Olive pers. comm.). In early 2004 it became established in Townsville

(Valentine 2004, 2005), where it still occurs in 2016 (P.S. Valentine pers.

comm.). The availability of Y. s. parva from near populated areas in recent

years has made possible a closer examination of its life history, thereby

enabling a better comparison with that of Y. algina reared in butterfly houses

in eastern Australia.

Since the late 1980s, what was considered to be Y. s. parva has been reared in

many butterfly houses along the east coast of Australia. The initial stimulus

for the current study was in 2008, when one of us (RK) attempted to feed

larvae of what was then thought to be a butterfly house population of Y. s.

parva on D. prostatus (Poir.) Nees. [syn R. prostata]. This attempt was

unsuccessful and puzzling since Dipteracanthus spp were the known larval

hosts for Y. s. parva (Sankowsky 1991), yet none of the larvae would feed

and all subsequently died. Consequently, this led to additional investigation

of the two taxa.

214 Australian Entomologist, 2016, 43 (4)

Based on the current study, it is now apparent that these butterfly house

populations of Yoma are not Y. sabina but are a separate species, Y. algina.

This raised questions in regards to where in Australia the original culture

material was derived or did it somehow originate from New Guinea stock.

The exact origin of the original culture material that was used to commence

these in-house butterfly populations is unknown but it is likely that it

originated from a single female from Cape York Peninsula (G. Sankowsky

pers. comm.). In addition, this current study of the life histories of both taxa

has now indicated that the life history observations of Wood (19872) were of

Y. algina, not Y. s. parva as he supposed. Moreover, his larval host plant

identification was also incorrect (Wood 19872), as larvae of Y. algina, as

reported here, are restricted to Hemigraphis spp and do not feed on

Dipteracanthus spp (syn Ruellia spp).

As a result of an investigation of both taxa, described in this paper, viz. wing

pattern elements, species' distributions, larval host plant preferences,

immature stage morphologies and structures of male genital armature, we

confirm that there are two species of Yoma and that both species naturally

occur on mainland Australia. We also remove the confusion surrounding the

immature stages of both species by describing and illustrating the life history

of both in full for the first time. Thus, we unravel the confusion that has

surrounded these two species in Australia since the 1980s and further revise

the taxonomic status of Y. sabina in Australia.

Materials and methods

Rearing of immature stages and comparison of larvae and pupae

The immature stages of Y. s. parva were reared from eggs supplied ex wild-

caught females collected from the Wet Tropics, Queensland. Those of Y.

algina were reared from eggs supplied by the Melbourne Zoo ex butterfly

house population. Eggs of both species were reared to instar 3 in clear plastic,

round food containers (280 ml; 50 mm high, bottom radius 42 mm, top radius

55 mm) on the following plant material: Y. s. parva on D. prostatus, H.

alternata (Burm.f.) T. Anderson and H. ciliata S. Moore, and Y. algina on the

two Hemigraphis spp.

Larger larvae (instars 4 and 5) were later transferred to potted host plants in

mesh cages (840 high x 480 x 400 cm). Larvae pupated beneath leaves or

stems of the host plants, on the overhangs of pot plants, or on the ceilings of

mesh cages. All life history studies were conducted in Brisbane during

February and March at ambient conditions (i.e. max/min temperatures of

approximately 32/22?C). Each instar of both species was described, measured

and photographed. To determine any differences between the larvae and

pupae of the two species, final instar larvae and pupae of Y. s. parva (n = 24)

and Y. algina (n = 18) were compared.

Australian Entomologist, 2016, 43 (4) 215

Comparison of wing pattern elements among adults

To determine the extent of phenotypic variation between the two species,

wing pattern, including colours and spotting, plus wing shape, were

subjectively compared among 78 specimens of Y. s. parva (428 8, 369 9)

from Queensland, 16 specimens of Y. s. parva (5463, 1199) from the

Northern Territory, 29 specimens of Y. s. sabina (21 d, 899) from Papua

New Guinea; 18 specimens of Y. algina (856, 1099) ex butterfly house

culture, 10 specimens of Y. algina (65 4, 499) wild caught in Queensland

and eight specimens of Y. a. netonia (656, 29 9) from Papua New Guinea.

Request for data from butterfly collections

Commencing in 2009, requests with attached figures of both taxa were

disseminated throughout the butterfly collecting fraternity (n 2 8) and public

museums (n = 4) for information concerning Yoma specimens in their

possession. In brief, the requests carried images of both species, male and

female, and the recipients were requested to use wing pattern elements and

wing shape to discern if Y. algina existed in any of their collections (the

specifics of this request are provided in Appendix I).

Comparison of male genitalia

The genital armature of three males of each taxon (i.e. 25$ from Australia

and 13 from Papua New Guinea) were dissected from the abdomens and

prepared for examination. Each abdomen was treated with 10% (w/v)

aqueous potassium hydroxide (KOH) for 24 hours at room temperature,

similar to the method used by Braby (2000). Clarified genitalia were then

stored in glycerol and examined unmounted using a stereomicroscope. From

these examinations the genital armature of a representative of each taxon was

chosen and photographed, then structural comparisons were made between

the two taxa. Male genitalia examined of Y. s. parva were from Malanda and

Thursday Island, Queensland; those of Y. s. sabina were from Kapa Kapa,

Central Province, Papua New Guinea; and those of Y. algina were from the

Melbourne Zoo Butterfly House (2). In addition, male genitalia were

examined of one Y. a. netonia from Rigo, Central Province, Papua New

Guinea. Nomenclature of genital structures follows Braby (2000) and

Monastyrskii (2011).

Abbreviations

Repositories of material examined: AM — Australian Museum, Sydney;

ANIC - Australian National Insect Collection, Canberra; CGMC - Collection

of C.G. Miller, Lennox Head; GRFC - Collection of G.R. Forbes, Brisbane;

HUC - Collection in Harvard University, Cambridge, MA, USA; MV —-

Museum of Victoria, Melbourne; QM — Queensland Museum, Brisbane;

TLIKC - Joint collection of T.A. Lambkin and A.I. Knight, Brisbane; WJC —

Collection of W. Jenkinson, Beaudesert.

216 Australian Entomologist, 2016, 43 (4)

Names on labels: AA — A. Atkins; AIK — A.I. Knight; CGM - C.G. Miller;

CWM - C.W. McCubbin; EDE - E.D. Edwards; EM - E. Mann; GAW —

G.A. Waterhouse; GBM - G.B. Monteith; GD — G. Daniels; GD&MAS - G.

Daniels and M.A. Schneider; GRF — G.R. Forbes; GW — G. Wurtz; HR - H.

Rauber; JAK - J.A. Kershaw; IFBC&MSU - LF.B. Common and M.S.

Upton; JCLS — J.C. Le Souef; JFD — J.F. Donaldson; JO — J. Olive; JWCD —

J.W.C. D'Apice; LR - L. Radunz; MDB - M. De Baar; MSM&BJM - M.S.

Moulds and B.J. Moulds; OBL - O.B. Lower; PZ - P. Zborowski; RGE -

R.G. Eastwood; TAL — T.A. Lambkin; TAW&IDN - T.A. Weir and I.D.

Naumann; THG - T.H. Guthrie; WBB - W.B. Barnard; WJ&DB — W.

Jenkinson and D. Bell; WJ&RKP — W. Jenkinson and R.K. Poulier.

Material examined

Yoma sabina parva (Butler, 1876)

QUEENSLAND: 19, Bamaga, Cape York, 28.iii.1964, IFBC&MSU (ANIC); 1d,

Batavia Downs, Cape York Peninsula, 13-19.1.1993, PZ (ANIC); 19, Cairns, captive

bred Indooroopilly, Brisbane, 30.xii.2008, TAL (TLIKC); 19, same data except

AA185; 14, Cape York, (QM); 1, same data except (ANIC); 19, Cape York,

19.x.1927, WBB (QM); 18, 19, Captain Billy Creek road junction, Cape York

Peninsula, 11°41'S 142?42'E, 15.1.1992, GD&MAS (QM); 146, Claudie River,

25.v.1974, JWCD (ANIC); 13, Chillagoe, OBL (QM); 19, Coen, Cape York

Peninsula, 9-16.vii.1971, GBM (QM); 26 4, Cooktown, 20.iv.1922 (ANIC); 26 4,

299, same data except 2.viii.1979 (24 4), 7.vii.1964 (9), 8.vii.1964 (9), JCLS

(ANIC); 28. d, 29 9, 35 km NW of Cooktown, 7.v.2003, WJ&DB (WJC); 18, Dauan

Island, Torres Strait, 18.1.2004, TAL (TLIKC); 16, same data except 11.ii.2015

(TLIKC); 15, 29 9, same data except 12.11.2015; 19, Gordon Creek area, Iron Range,

Cape York Peninsula, 1.vii.1982, GD (QM); 34d, 29 9, Green Hill, Thursday Island,

Torres Strait, 23-31.vii.1983, TAL (TLIKC); 14, 19, same data except 20.ii.1994;

334, 19, same data except 24.ii.1994; 19, 19, same data except 24.iii.1994; 19,

same data except 9.11.2001; 19, same data except 3.11.2004, AIK; 19, same data

except 27-29.iii.1987, MDB; 14, Heathlands, Cape York Peninsula, 16.iii.1992,

GD&MAS (QM); 1ĝ, Iron Range NP, Cape York Peninsula, 27.v.2010, WJ&RKP

(WJC); 14, same data except, 26.v.-8.vi.??, JWCD (ANIC); 19, same data except 18-

31.viii.1999; 19, same data except 4-12.vii.1995; 1$, same data except 13.iv.1971,

AA; 13, same data except -viii.1968, THG; 18, 19, Malanda, captive bred

Indooroopilly, Brisbane, 30.xii.2009, TAL (TLIKC); 19, same data except AA180;

12, same data except AA178, em. 1.v.2010; 19, Mer Island, Torres Strait, 1.vi.1985,

JFD (TLIKC); 19, same data except 30.11.1986, MDB (TLIKC); 1, same data

except 30.11.1986, MDB (TLIKC); 19, same data except 9.iii.1995, TAL; 1i, same

data except 8-14.v.1998, JWCD (ANIC); 19, same data except 25.iv-5.v.1999, AIK

(TLIKC); 14, same data except 25.1.2011, TAL&AIK; 14, same data except

28.2011, TAL&AIK; 16, same data except 29.122011, TAL&AIK; 19, same data

except 9°54'S 144°02'E, 9.1.2015, TAL&AIK; 344, Mt Cook NP, Cooktown,

12.x.1980, EDE (ANIC); 284, Mt Webb NP, -.ix.1980, EDE (ANIC); 19, Port

Stewart Rd, Cape York Peninsula, 3.v.2003, WJ&DB (WJC); 19, Red Cliff, Cairns,

28.v.1973, JWCD (ANIC), 26 d, Somerset, 10.iv.1906 (ANIC); 18, 19, Thursday

Island, 5.vi.1969, CWM (ANIC); 19, Trinity Park, N of Cairns, 1.xii.2009, ex captive

9, JO (TLIKC).

Australian Entomologist, 2016, 43 (4) 217

NORTHERN TERRITORY: 53, 1099, Rocky Bay, Gove, 3-7.v.1992, JWCD

(ANIC); 1 9, Nhulunbuy, NT, Aug-Sept 1984, GW (ANIC).

Yoma sabina sabina (Cramer, [1780])

PAPUA NEW GUINEA: 2464, Brown River, near Port Moresby, 9°15'S 147°05'E,

2.1.1969, 15.ix.1975, HR (QM); 1c, Kwikila, Central Province, 19.11.1976, GRF

(GRFC); 14, Rigo, PNG, 17.x.1968, GRF (GRFC); 4G 4, 599, Kapa Kapa, Central

Province, 9°48'S 147°31'E, 20.xi1967 (9), 14.xi.1968 (9), 21.xi.1968 (d,

26.xii.1968 (9), 2.ii.1969 (4), 6.1.1969 (d, 29 9), 23.xi.1973 (3), HR (QM); 64d,

same data except 4.viii.1968 (i$), 10.viii.1968 (4), 11.viii.1968 (288), 3.11.1969

(243), 6.11.1969 (333), GRF (GRFC); 446, same data except, 16.iv.1973 GRF

(GRFC); 19, Kapa Gese, Central Province, PNG, 19.11.1976, GRF (GRFC); 19,

Kwikila, Central Province, 19.1.1976, GRF (GRFC), 14,19, Paga Hill, Port

Moresby, 18.1.1966, ex pupa, EM (QM); 146, same data except 19.1.1966; 14, Port

Moresby, PNG, 3.11.1971, GRF (GRFC).

Yoma algina (Boisduval, 1832)

QUEENSLAND: 1d, 19, Claudie River, JAK (AM); 16, Heathlands, Cape York

Peninsula, 11?45'S 142°35'E 15-26.1.1992, TAW&IDN (ANIC); 19, Iron Range, Cape

York Peninsula, 1-9.vi.1971, GBM (QM); 14, 19, Iron Range, 25.v.1973 (8),

28.v.1973 (9), CGM (CGMC); 18, bred as Y. sabina, MDB, labelled ‘ex breeding

stock supposed to originate from Iron Range Qld, set Nov. 1990, original stock

probably PNG’ (TLIKC); 19, Lockerbie Area, Cape York Peninsula, 13-27.iv.1973,

GBM (QM); 19, captive bred, ex. Melbourne Zoo colony, em. 21.xi.2009, TAL, Host

Hemigraphis (TLIKC); 44d, 999, same data except, em. 1.xii.2009 (299),

21.xii.2009 (14, 19), 22.xii.2009 (256, 299), 23.xii.2009 (13), 8.vi.2010 (19),

10.vi.2010 (19), 10.xi.2010 (19), 23.xii.2010 (19); 14, same data except AA168;

14, same data except AA1176; 14, same data except 30.iv.2010, AA170; 1, same

data except 1.v.2010, AA171; 14, Peach Creek, 25 km NNE of Coen, 2.xi.1979,

MSM&BJM (AM); 16, Philip Hill, Iron Range, 10.111994 RGE (HUC); 12,

Somerset, Cape York Peninsula, Lep 11424, 10. iv. 1906, 'doubtfully Australian,

GAW' (MV).

Yoma algina netonia Fruhstorfer, 1912

PAPUA NEW GUINEA: 1c, Bulolo, Morobe Province, 20.1.1970, LR (QM); 13,

Gabensis, Morobe Province, 7.vii.1973, GRF (GRFC); 14, Kapa Kapa, Kokoda

Track, Central Province, 15.iv.1968, HR (QM); 299, Laloki, Central Province,

18.11.1976, GRF (GRFC); 16, Rigo, Central Province, 15.x.1968, GRF (GRFC), 1d,

same data except 16.x.1968; 14, Wau, Morobe Province, 26.iii.1978, GRF (GRFC).

Results

Larval host plants

Larvae of Y. s. parva would not feed on Hemigraphis spp when young but

when larger readily accepted this host. All larvae of Y. algina fed freely on

Hemigraphis spp but refused to feed on D. prostatus. Thus, the results

indicated larvae of Y. algina to be host specific to several species of just

Hemigraphis and furthermore these larvae would not accept Dipteracanthus

spp. In addition, larvae of Y. s. parva differed greatly from those of Y. algina

218 Australian Entomologist, 2016, 43 (4)

in that they readily accepted D. prostatus and only as large larvae accepted H.

alternata and H. ciliata.

Life histories

Yoma sabina parva (ex Malanda and Trinity Park)

Egg (Fig. 1): (n = ca 40); dome shaped; smooth with 14 vertical ribs.

First instar larva (Fig. 2): (n 2 ca 40); head shiny black, covered in pale

olive-green setae, with two short blunt black horns; body cylindrical, dorsal

surface black, ventral surface with last abdominal and anal segment yellow-

green and semi-translucent; mesothorax, metathorax and abdominal segments

each with six branched spines with bristles, with the lateral basal spine and

another single bristle below each spiracle; prolegs yellow-green, legs black.

Second instar larva (Fig. 3): (n 2 ca 40); head as in first instar except setae

black and horns longer; body as in first instar, shiny black with branched

spines longer, with basal lateral spine originating from a circular bright

orange patch; prolegs and legs black.

Third instar larva (Fig. 4): (n 2 ca 30); similar to second instar except spines

longer and more branched, body with ventral bristly setae, a faint white

lateral line joining orange patches and a pair of faint dorsal parallel white

stripes running the full length of the body.

Fourth instar larva (Fig. 5): (n = 24); similar to third instar except body

covered with many bristly setae, a definite white lateral line intermittently

running the length of the body just above the setae, a faint lateral orange line

joining orange patches and dorsally a defined pair of white parallel stripes

running the length of the body.

Fifth instar larva (Fig. 6): (n 2 24); similar to fourth instar except pair of

spiny horns on head longer than the basal width between the two horns (Fig.

7); body black, smooth with noticeable white blotches forming two indefinite

parallel lateral white lines and two indefinite parallel dorsal white lines, these

dorsal white lines creating a noticeable dorsal black stripe between them;

dorsal and lateral surfaces of body densely covered with fine white setae;

dorsal black spines with a blue lustre with bases of spines blue.

Pupa (Figs 8-9): (n 2 24); mottled brown and grey; anterior end with two

projections; mesothorax with a sharp 90? dorsal ridge, two sharp lateral

projections on edge of each wing case; with metathorax and abdominal

segments possessing pairs of blunt spines; those spines on metathorax and

abdominal segments 1 and 2 white; the largest spines being on abdominal

segments 3 and 4, with pairs of large black and very small white blunt spines

on abdominal segment 3, those on remaining abdominal segments blunt and

black.

Australian Entomologist, 2016, 43 (4) 219

5 i rh

M M M

Figs 1-9. Early stages of Yoma sabina, Malanda, Qld: (1) egg (height 1 mm); (2) Ist

instar larva (length 7 mm); (3) 2nd instar larva (10 mm); (4) 3rd instar larva (18 mm);

(5) 4th instar larva (40 mm); (6) 5th instar larva (45 mm); (7) 5th instar larval head

capsule (5 mm wide); (8-9) pupa, lateral views (height 25 mm).

220 Australian Entomologist, 2016, 43 (4)

Yoma algina (ex Melbourne Zoo Butterfly House)

Egg (Fig. 10): (n 2 circa 50); dome shaped; smooth with 14 vertical ribs.

First instar larva (Fig. 11): (n 2 18); head shiny black, covered in cream-

coloured setae, with two short blunt black horns; body cylindrical, dorsal and

ventral surfaces black, with ventral surface, last abdominal and anal segment

orange and semi-translucent; mesothorax, metathorax and abdominal

segments each with six branched spines with bristles, with the lateral basal

spine and another single bristle below each spiracle; prolegs orange, legs

black.

Second instar larva (Fig.12): (n = 18); head as in first instar except setae

black and horns longer; body shiny black with branched spines much longer

than instar 1, with basal lateral spine originating from a circular dark orange

patch; prolegs brown and legs black.

Third instar larva (Fig. 13): (n 2 18); similar to second instar except spines on

body longer and more branched; body glabrous, with bristly setae on ventral

surface and a pronounced white lateral line joining orange patches; prolegs

brown with bases black.

Fourth instar larva (Fig. 14): (n 2 18); similar to third instar with white lateral

line more pronounced.

Fifth instar larva (Fig. 15): (n 2 18); similar to fourth instar except pair of

spiny horns on head as long as the basal width between the two horns (Fig.

16), body non-glabrous with dorsal and lateral surfaces densely covered with

fine white setae and bases of dorsal spines with a faint blue spot; body with a

pronounced cream-coloured lateral line.

Pupa (Figs 17-18): (n = 18); mottled brown; anterior end with two

projections; mesothorax with a sharp 90° dorsal ridge, two sharp lateral

projections on edge of each wing case; metathorax and abdominal segments

possessing pairs of sharp spines; those spines on metathorax (being large) and

abdominal segments 1 and 2 white; the largest spines being on abdominal

segments 3 and 4, with pairs of large brown and very small brown sharp

spines on abdominal segment 3; those on remaining abdominal segments

brown, sharp and curved upwards.

Comparison of larvae and pupae

The most notable differences between final instar larvae of the two species

are the positions of white or cream lines on the thoracic and abdominal

segments (Figs 6, 15): final instar larvae of Y. s. parva bear two ill-defined

parallel dorsal white lines (Fig. 6), while those of Y. algina bear a

pronounced cream-coloured lateral line (Fig. 15). In addition, final instar

larvae of Y. s. parva are overall paler in colour, bear fewer setae and are more

glabrous in body appearance than those of Y. algina, which appear darker,

bear more setae and have a more hirsuite body texture.

Australian Entomologist, 2016, 43 (4) 221

[

Pn tres ud MDC RICE eats at. "5 =

fy M

. =al ar mate ean "mo A aeu dune i

Figs 10-18. Early stages of Yoma algina, Melbourne Zoo culture: (10) egg (height 1

mm); (11) Ist instar larva (length 6 mm); (12) 2nd instar larva (9 mm); (13) 3rd instar

larva (20 mm); (14) 4th instar larva (30 mm); (15) 5th instar larva (45 mm); (16) 5th

instar larval head capsule (6 mm wide); (17-18) pupa lateral views (height 24 mm).

222 Australian Entomologist, 2016, 43 (4)

The younger instars (2, 3 and 4) of both species were also found to have

distinctive morphological differences (Figs 3-5, 12-14).

The final instar head capsules of both species are covered in facial setae and

bear two black horns. The Y. algina head capsule (Fig. 16) has more facial

setae than Y. s. parva (Fig. 7) and the horns of Y. algina (Fig. 16) are thicker

than those of Y. s. parva (Fig. 7).

The pupae of the two species are notably different in the pattern of thoracic

and abdominal spines, with pupae of Y. algina bearing much heavier and

longer spines (Figs 17-18) than those of Y. s. parva (Figs 8- 9). The pupae of

both species vary in colour, ranging from brown to dark grey, each with

white blotches.

Overall, our descriptions and illustrations of the immature stages of Y. s.

parva matched the illustrations of Y. s. sabina provided by Saguru and Haruo

(2000).

Comparison of wing pattern elements

Superficially, adults of the two Yoma species (Figs 19-32) resemble each

other but individuals can be readily separated using several wing pattern

elements. Thus, the upperside orange colour of both sexes of Y. algina is

much brighter and more vibrant (Figs 27-30, 32) than that of Y. s. parva (Figs

19-26, 31). Specifically for females, there are two orange spots in the

subapical forewing area of Y. s. parva (Figs 20, 22, 24-26), while the same

spots in Y. algina are white (Figs 28, 30, 32). Additionally, in females the

hindwing uppersides of Y. algina always bear distinct ocelli in the orange

submarginal areas (Figs 28, 30, 32).

On the underside, the wing patterns of Y. s. parva are much more variable

than those of Y. algina, with Y. s. parva possessing submarginal black spots

(Fig 19-26) rather than distinct ocelli in the same wing areas of Y. algina

(Figs 27-30). Moreover, both sexes of Y. algina always possess a broad

underside white band (Figs 27-30).

The shape of the forewing apex of both species differs, being pronounced and

falcate in Y. s. parva (Figs 19-26, 31) and only slightly projected in Y. algina

(Figs 27-30, 32). This is also the case for the short and blunt terminal tail on

the hindwing, that of Y. s. parva being relatively long but almost absent in Y.

algina.

Finally, based on the review of 29 specimens (214 6, 89 9) of Y. s. sabina

from Papua New Guinea and 94 specimens (478.8, 4799) of Y. s. parva

from Australia, plus their variability in general, we found no consistent

upperside wing pattern elements that can be used to confidently separate the

two subspecies. In addition, the underside wing patterns of both populations

are so highly variable that no differentiation between the two populations

could be made using wing underside patterns.

Australian Entomologist, 2016, 43 (4) 223

Figs 19-24. Yoma sabina (all figures not to scale, upperside left, underside right). (19,

21, 23) ác: (19) Malanda, Qld, 30.xii.2009, TAL [forewing length 38 mm]; (21)

Dauar Island, Torres Strait, Qld, 12.11.2015, TAL [42 mm]; (23) Mer Island, Torres

Strait, 25.1.2011. TAL&AIK [41 mm]. (20, 22, 24) 99: (20) Malanda, Qld,

30.xi1.2009, TAL [42 mm]; (22) Dauar Island, 12.11.2015, TAL [39 mm]; (24) Mer

Island, 30.11.1986, MDB [38 mm].

224 Australian Entomologist, 2016, 43 (4)

Figs 25-30. Yoma spp (all figures not to scale, upperside left, underside right). (25-26)

Yoma sabina 9 9: (25) Cairns, Qld, 30.xii.2008, TAL [forewing length 42 mm]; (26)

Green Hill, Thursday Island, Torres Strait, 20.11.1994, TAL [36 mm]. (27-30) Yoma

algina: (27) 3, ex Melbourne Zoo colony, em. 22.xii.2009, TAL [35 mm]; (28) 9, ex

Melbourne Zoo colony, em. 10.xi.2010, TAL [39 mm]; (29) 4, Iron Range, Qld,

1990, MDB [35 mm]; (30) 9, ex Melbourne Zoo colony, em. 22.xii.2009, TAL [38

mm].

Australian Entomologist, 2016, 43 (4) 225

Figs 31-32. Live adults of Yoma spp in captivity in typical perched positions on forest

edges (figures not to scale): (31) Y. sabina £ [forewing length 42 mm]; (32) Y. algina

9 [40 mm]. (Photographs courtesy of G. Sankowsky, Tolga).

Request for data from butterfly collections

The request for collection data from institutional (4 out of 4 responded) and

private collections (7 out of 8 responded) produced 10 specimens (in three

institutional and two private collections) of wild caught Y. algina (6 3, 4

99), all from Cape York Peninsula, Queensland (Figs 33-44). The collection

data further indicated that the specimens all originated from the east coast of

Cape York Peninsula, i.e. extending north from Peach Creek Crossing near

Coen, through Claudie River, Iron Range and Heathlands, to Somerset and

Lockerbie at Cape York. Over half the specimens (6) originated from the

Claudie River basin (Figs 39-44) and the earliest known specimen was

collected in 1906 at Somerset (Figs 33-35, in MV). AII five locations contain

tracts of rainforest or semi-deciduous monsoonal vine thicket. Other wild

caught specimens not illustrated are: £, 9, Claudie R, 1914 (AM); ĝ, Peach

Ck, 25 km NNE of Coen, 1979 (AM); c, Heathlands, 1992 (ANIC); d,

Philip Hill, Iron Range, 1994 (HUC).

Comparison of male genital armature

The male genital armature (Figs 45-46) of the two species are markedly

different, with just about all structures distinctly dissimilar. Most notable are:

the relatively small claspers (valvae) of Y. algina; the relatively very thin

apical tip of the phallus (aedeagus) of Y. s. parva; the unique structure of the

gnathos of each species is very diagnostic, that of Y. algina being very large

with spines; although the unci in both species are hook-shaped, the uncus of

Y. algina is much more so. There are also noticeable differences in the shapes

of the tegumen, vinculum and saccus of each species. Aside from these

differences, Parsons (1998) reiterated the comment by Fruhstorfer (1912-

1915) on the general complexity of the structures of the male genitalia of

Yoma spp. This study certainly found the same, particularly the complicated

structure of the clasper, uncus and gnathos (Figs 45-46: c, d and e

respectively).

226 Australian Entomologist, 2016, 43 (4)

— "

LEP-11424

Lockerbie Area, |

G.B. Monteith

QM Reg. Na.

T185733

j1-9.vi*1971

36. B. Monteith

M Reg. Na.

| 7189734 |

Figs 33-44. Some of the wild caught Australian specimens of Yoma algina with their

origins (all Cape York Peninsula) and repositories: (33-35) 4, Somerset, 1906 (MV);

(36-38) 9, Lockerbie, 1973 (QM); (39-41) £, 9, Iron Range, 1973 (CGMC); (42-44)

Q, Iron Range, 1971 (OM); forewing lengths of specimens were not recorded.

Australian Entomologist, 2016, 43 (4) 227

Figs 45-46. Male genital armature, lateral left view, dorsal surface uppermost: (45)

Yoma algina, ex Melbourne Zoo culture, captive bred Indooroopilly, Brisbane, Qld,

10.vi.2010, TAL, host Hemigraphis (TLIKC); (46) Yoma sabina, Green Hill,

Thursday Island, Torres Strait, 3.11.2004, AIK (TLIKC). Approximate widths of

genitalia including saccus = 5 mm; names of structures labelled are: a — saccus, b —

tegumen, c — clasper, d — uncus, e — gnathos and f — phallus.

Discussion

In the early 1980s, a stock of Y. algina (believed at the time to be Y. s. parva)

was used to commence a commercial butterfly house culture of Yoma.

Despite the adults of both species being similar in their wing pattern

elements, the individuals of Y. algina flying in the butterfly houses appeared,

in general, to be much brighter in appearance than Y. s. parva, with a more

intense and vibrant orange colouration on the upperside of the wings,

particularly in the males. Because very few individuals of Y. s. parva had

been reared at that time, it was thought that the brightness of individuals

reared in butterfly houses was perhaps related to the fresh colour of newly

eclosed individuals.

The question remains, where did the butterfly house culture stock of Y. algina

originate? When first identified, this species was not known from Australia

and it was presumed that livestock originated from Papua New Guinea. Initial

discussions with several collectors and butterfly workers suggested that the

Yoma culture first established in the Melbourne Zoo Butterfly House came

from live material collected either from Iron Range by G. Wood (as per

Wood 19872), or from Weipa by the late Charles McCubbin who, just prior

to his death in 2010, told one of us (RK) that he collected live Yoma females

from Weipa for the Melbourne Zoo in the early 1990s, not long after the zoo

established its butterfly house, although Yoma is still unrecorded from Weipa

(Braby 2000, 2016). G. Sankowsky (pers. comm.) rejected the latter claim

and stated that Graham Wood, who was working at Iron Range in the 1980s,

sent him a gravid female and resulting progeny were supplied to the

Australian Butterfly Sanctuary at Kuranda, Queensland and, subsequently, to

the Melbourne Zoo in the late 1980s. Based on this latter information, the

most plausible scenario is that Melbourne Zoo's Y. algina most likely

originated from Iron Range, Cape York Peninsula, in the late 1980s.

228 Australian Entomologist, 2016, 43 (4)

The review of Yoma specimens in public and private collections indicated

that, in Australia, Y. algina occurs sympatrically with Y. s. parva exclusively

in eastern Cape York Peninsula, roughly from MclIlwrath Range (i.e. Peach

Creek Crossing) near Coen, north to the tip of Cape York Peninsula (i.e.

Somerset). The distributions of the two species are also known to overlap in

southern Papua New Guinea (Parsons 1998). The true taxonomic identity of

this isolated Australian population of Y. algina will remain uncertain until a

more thorough revision is made, particularly of material from New Guinea

and until more wild-caught Australian specimens become available. The

Australian population is tentatively assigned here to the southern Papua New

Guinean subspecies Y. a. netonia (Parsons 1998), based solely on the

geographical proximity of eastern Cape York Peninsula to southern Papua

New Guinea.

The introduction of Y. a. netonia from Iron Range into the butterfly houses in

eastern Australia added a degree of misunderstanding in the literature,

particularly with the life history and identification of Y. s. parva. Wood

(1987a) was confused with the host plants that he used to rear what he

thought at the time to be Y. s. parva. He illustrated, in monochrome, a final

instar larva that he identified as Y. s. parva, but which clearly matches the

mature larva of what we now know to be Y. a. netonia. A similar mistake,

this time in identifying the adults, was perpetuated by several authors who

illustrated Y. a. netonia instead of Y. s. parva in their publications on

Australian butterflies (viz. Valentine 1988, Braby 2000, 2004, 2016, Orr and

Kitching 2010).

Currently, the immature stages of Y. a. netonia have not been found in the

wild in Australia but, based on our host plant preference experiments,

Hemigraphis appears to be the preferred host plant. Based on records from G.

Sankowsky (pers. comm.) and Australia's Virtual Herbarium (2016), H.

ciliata appears to be the sole native Australian species of Hemigraphis

occurring in Queensland. Current records indicate that H. ciliata is restricted

to the eastern side of Cape York Peninsula, mostly from the Pascoe River to

the Rocky River in damp shady areas in very well developed rainforest,

although there are a few records of it further south around Cape Tribulation.

These distribution records, albeit in all probability incomplete, correlate

roughly with the known distribution of Y. a. netonia in Queensland. If H.

ciliata is the native host, then Y. a. netonia might also occur further south

around Cape Tribulation in the Wet Tropics.

Hemigraphis alternata is a naturalised species thought to originate in Java,

Indonesia. Australia's Virtual Herbarium (2016) records indicate that it 1s

currently known from tropical Queensland, between Port Douglas and

Cooktown and near Almaden. Other Hemigraphis spp in cultivation in

Queensland include H. reptans (Roth) J.R.J. Wood from the Pacific Islands

and H. urens from India. A cultivated Hemigraphis sp. is reported to be

Australian Entomologist, 2016, 43 (4) 229

common in gardens and council plantings in the wet tropical lowlands of

northern Queensland (G. Sankowsky pers. comm.).

Several native Dipteracanthus spp are recorded from tropical Australia

(Australia's Virtual Herbarium 2016), including D. australasicus F. Muell.

from much of tropical Australia and D. bracteatus from tropical Queensland

north of Cairns, the Top End of the Northern Territory and in Torres Strait.

The distribution of D. bracteatus roughly matches that of Y. s. parva. In

addition, G. Sankowsky (pers. comm.) has reared Y. s. parva on Brunoniella

australis (Cav.) Bremek (Acanthaceae), which occurs widely throughout

eastern Australia and the Northern Territory (Australia's Virtual Herbarium

2016) and therefore could be a natural host plant of Y. s. parva in the

Queensland tropics, including Iron Range. Braby (2016) recorded B.

spiciflora (F.Muell. ex Benth.) Bremek as a host of Hypolimnas alimena

lamina Fruhstorfer, 1903 (Nymphalinae: Junoniini) in Australia.

Based on his knowledge of Yoma in Cape York Peninsula, G. Sankowsky

(pers. comm.) further indicated that, in the wild, the two Yoma spp are

separated by disparate ecotones: Y. s. parva generally occurs in a variety of

habitats from more open forest to lowland rainforest (Valentine 1988) where

D. bracteatus and B. australasicus grow, while Y. a. netonia is restricted to

rainforest where H. ciliata grows.

Although Y. s. parva is well known from mainland Australia (Braby 2000,

2016), much less is known of its distribution throughout Torres Strait, where

the species appears to be confined to islands with stands of semi-deciduous

monsoon or vine forest. These islands include: in the south of the strait,

Thursday (Waiben) (Waterhouse and Lyell 1914, Lambkin and Knight 1983,

De Baar 1988, Valentine 1988), Horn (Ngurupai) (unpublished data, T.A.

Lambkin: December 1993, January 1994), Prince of Wales (Muralug)

(Waterhouse and Lyell 1914) and Hammond (Keriri) (unpublished data, T.A.

Lambkin: April 1989); in the central region, Moa (Waterhouse and Lyell

1914, Valentine and Johnson 1993); in the east, Darnley (Erub) (Waterhouse

and Lyell 1914, Johnson 1983), Murray (Mer) (Waterhouse and Lyell 1914,

Wood 1987b, Lambkin and Knight 1990) and its neighbouring island Dauar

(TLIKC); and Dauan Island (TLIKC) in the north. A useful map indicating

the positions of these islands and others in Torres Strait is illustrated in Braby

(2000: p. 17).

In this study, evidence from several data sources, i.e. larval colour pattern,

morphology and host plant requirements, pupal morphology, adult wing

shape, colour pattern elements and male genitalia, support a specific

distinction between the two taxa, based on the principles of the species

concept (de Queiroz 2007). Although the two species are similar in their

biology and biotype requirements and share some sympatry in their

geographical distributions, our investigations identified at least 22 phenotypic

character states that the two taxa do not share.

230 Australian Entomologist, 2016, 43 (4)

Six character states were found in the final instar larvae that differentiated the

two species: the pattern and position of cream-coloured lines on, and the

basal colour and texture of, the thoracic and abdominal segments; the degree

of thoracic and abdominal setae; and, on the head, differences in the degree

of facial setae and thickness of the horns. There was at least one character

state that differentiated the pupae, with the thoracic and abdominal spines of

Y. a. netonia being heavier and longer that those of Y. s. parva. Differences in

wing pattern elements (i.e. colour, pattern arrangement and shape) between

the two taxa were notable, with seven character states that differed between

the two species. The male genitalia of the two taxa are dissimilar in the shape

and size of the claspers, phallus, gnathos, uncus, tegumen, vinculum and

saccus (seven character state differences). Finally, Y. a. netonia appears to be

host specific to Hemigraphis spp (another character state difference).

This study also highlighted the similarity of the nominate subspecies of Y.

sabina to the Australian taxon Y. s. parva. Both taxa meet at a theoretical line

situated on the southern coast of Papua New Guinea: to the north of this line

is Y. s. sabina while to the south is Y. s. parva. We questioned the basis of

this separation during our study of Y. sabina from Australia and Papua New

Guinea. Thus, our review of the description of Y. s. parva by Butler (1876)

has revealed that his description (as Rhinopalpa parva) was based, in all

probability, on a single male specimen (Edwards et al. 2001) collected at

Cape York by Rev. J.S. MacFarlane. More interestingly, Butler's (1876)

description matches the wing pattern elements of a phenotypically distinct,

diminutive ‘dry season form’ referred to by Waterhouse and Lyell (1914)

(Figs 47-48). This form is typically observed just after the commencement of

the first rains of the monsoonal wet season.

Figs 47-48. Yoma sabina sabina form parva (figures to scale, upperside left,

underside right): (47) @, Dauar Island, Torres Strait, Qld, 12.ii.2015, TAL [forewing

length 30 mm]; (48) 9, Dauar Island, Torres Strait, Qld, 11.11.2015, TAL [forewing

length 34 mm].

Australian Entomologist, 2016, 43 (4) 231

It is obvious now that Butler (1876) had before him a specimen of this form,

even remarking at the end of his description that 'This is the smallest

Rhinopalpa that I have seen' and based his description of parva on this

specimen, not on what is the more typical *wet season' Y. sabina that occurs

in Australia and Papua New Guinea (Figs 19-26). Being smaller in size is one

of the key characters of the ‘dry season form’, having more orange in the sub-

marginal areas of the forewing upperside and, in the male, possessing a

distinctive tawny spot in the upperside forewing cell, all features that Butler

(1876) described. This diminutive form is now known from both Papua New

Guinea (Parsons 1998) and Australia (Braby 2000).

Thus, our study validates Fruhstorfer (1912-1915), Barrett and Burns (1951),

Parsons (1998) and Tennent (2002) in their treatment of the two taxa as

distinct species and, moreover, confirms the natural occurrence of Y. a.

netonia on mainland Queensland. Based on the known distributions of the

larval hosts in Cape York Peninsula and beyond into the Wet Tropics and

possibly Torres Strait, the distribution of Y. a. netonia may be more extensive

than current collection records indicate.

In addition, the staggered distribution of Y. s. parva across Torres Strait is

likely an artefact of butterfly collecting efforts on specific islands, with some

inhabited islands with suitable habitats still not surveyed well for butterflies.

Thus, Y. s. parva in Torres Strait is also probably more widespread than

presently known and this distribution is almost certainly related to the

distribution of its larval host plants (Dipteracanthis spp and possibly

Brunoniella spp).

Furthermore, based on our review of Y. sabina from Papua New Guinea and

Australia, it was found that the two populations differed little phenotypically

and, accordingly, we regard the subspecies Y. s. parva (Butler, 1876), syn.

nov. as a junior synonym of Y. s. sabina. Provisionally, until a proper

revision of the 'dry season form' is undertaken, we also propose this

diminutive 'dry season' form as Y. s. sabina form parva stat. rev.

Finally, further intensive collecting of Yoma spp may reveal the presence of

Y. a. netonia in areas such as the western side of Cape York Peninsula, in the

Wet Tropics and in the southern Torres Strait, in particular on the southern

islands of Prince of Wales and Hammond. These two islands have extensive

areas of suitable habitat and have been visited relatively infrequently by

butterfly workers.

Acknowledgements

We thank the local community councils and island Elders of several Torres

Strait islands for allowing entry into their communities and providing

assistance during the collection of material used in this study. Appreciation is

given to D. Britton (AM), the late M. De Baar (now TLIKC), R.G. Eastwood

(ex Harvard University, USA) (HUC), E.D. Edwards (ANIC), G.R. Forbes

232 Australian Entomologist, 2016, 43 (4)

(GFC), W. Jenkinson (WJC), CG Miller (CGMC), K. Walker (MV) and S.

Wright (QM), for allowing access to specimens in their care or for providing

identification of specimens or supplying images. G. Thompson (QM)

provided the high quality photographs of the two QM specimens. In addition,

appreciation is given to G. Sankowsky (Tolga, Atherton Tableland) for his

valuable personal communication and provision of unpublished data on host

plant taxonomy, distributions and images. Taxonomic clarification of host

plants was also provided by R. Barker (State Herbarium of South Australia).

Eggs of Y. a. netonia were kindly provided by the Melbourne Zoo Butterfly

House and those of Y. s. parva were generously provided by W. Graham

(Malanda, Atherton tableland) and J. Olive (previously of Trinity Park,

Queensland). A review of the first draft of the manuscript by M.P. Zalucki

(University of Queensland) was greatly appreciated. This paper partially

fulfils the requirements for a Master of Philosophy degree undertaken by the

senior author at the University of Queensland, Brisbane.

References

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Appendix I

Specifics of the request for data from butterfly collections to ascertain the

existence of wild caught specimens of Australian Y. algina:

‘A recent examination of several purported Australian specimens indicates

that two species of Yoma could occur in Australia, viz. Y. sabina and Y.

algina. Yoma algina is predominately a New Guinean species and

superficially resembles Y. sabina. In general, the underside wing pattern of Y.

algina is less variable than sabina; the orange colouring of the algina male is

brighter than sabina; the forewings of algina are less falcate and in the

underside submarginal areas algina has ocelli instead of the black dots of

sabina. Specifically, in the females, algina has submarginal ocelli on the

upperside hindwings, and the apical markings of the forewing upperside are

white instead of orange as in the female of sabina. To help elucidate this

issue and confirm if Y. algina naturally occurs in Australia, we would

appreciate receiving collection data of Yoma spp, together with an

identification based on the attached figures that show wing pattern

differences between the two species and as outlined above.’

Australian Entomologist, 2016, 43 (4): 235-237 235

EUREMA BLANDA SARAHA (FRUHSTORFER) (LEPIDOPTERA:

PIERIDAE: COLIADINAE) REDISCOVERED IN TORRES STRAIT,

QUEENSLAND, AUSTRALIA

P.R. WILSON! and I. JOHNSON?

'2 Billabong Drive, Bundaberg, Qld 4670

?260 Grassdale Road, Gumdale, Qld 4154

Abstract

A female Eurema blanda saraha (Fruhstorfer, 1912) was collected on Dauan Island on 9 March

2016 and represents the first record from Torres Strait, Queensland, since 1909.

Introduction

Braby (2010) stated that Eurema blanda (Boisduval, 1836) is known, within

Australian limits, only from Christmas Island (Indian Ocean) and Darnley

Island in eastern Torres Strait. The record for the latter locality was based on

a single male originally in the Australian Museum, reputedly collected by H.

Elgner on 20 December 1909. The reliability of Elgner's record was in doubt,

however, since the species had not been encountered for over 100 years,

despite extensive surveys throughout the islands.

The original record was provisionally referred to subspecies E. blanda

indecisa (Butler, 1898) by Waterhouse and Lyell (1914) and Waterhouse

(1932) but subsequently replaced by E blanda saraha (Fruhstorfer, 1912), the

mainland New Guinea subspecies, in the first and revised editions of

Common and Waterhouse (1972, 1981) and in Braby (2000). In Papua New

Guinea, E. blanda saraha is widespread throughout the mainland and may be

common to abundant (Parsons 1998).

During a visit to Dauan Island in early March 2016, a single female of E.

blanda was collected on 9 March, feeding on flowers in monsoonal vine

scrub. The significantly larger size initially distinguished it from the very

abundant E. hecabe (Linnaeus, 1758).

Discussion

Eurema blanda resembles E. hecabe and E. alitha (C. & R. Felder, 1862) but

is distinguished by the sex brand in the males, the shape of the black marginal

bands on the upperside of each wing, the presence of three brown dots in the

cell of the forewing underside and the larger size compared with other

Australian Eurema Hübner, 1819 species (Common and Waterhouse 1981,

Parsons 1998).

The Dauan Island female of E. blanda (Figs 1-2) has the broad black

marginal band at the apex with the inner margin less sharply indented at vein

M; when compared with that of E. hecabe and E. alitha. The marginal band

on the hindwing upperside of the Dauan Island specimen is narrow and not

distinctly scalloped. The wingspan of 46 mm is consistent with that of E.

blanda saraha from Papua New Guinea (female 50 mm) (Parsons 1998).

236 Australian Entomologist, 2016, 43 (4)

Figs 1-2. Eurema blanda saraha female from Dauan Island, Torres Strait,

Queensland: (1) upperside; (2) underside.

Braby (2000) described seasonal variation in many of the Australian Eurema

species, with the ‘summer form’ often having faint or occasionally absent

markings on the underside of both wings, while the *winter form' usually has

more prominent markings. The specimen from Dauan Island lacks brown

spos on the underside of the wings and is consistent with the ‘summer form’

of other Eurema species. It also lacks the three spots in the cell of the

forewing underside that are typical of E. blanda; however, these cell spots

may be reduced or absent in some ‘forms’ of E. blanda (http://www.

boldsystems.org/index.php/Taxbrowser | Taxonpage ?taxid- 104436).

Parsons (1998) listed Caesalpinia bonduc (L.) Roxb. and various Cassia

species (both Fabaceae: Caesalpiniodeae) as food plants. These plant species

occur on Dauan Island, with Caesalpinia being reasonably abundant on the

beach ridges along the northern and eastern sides of the island. Parsons

(1998) also stated that females are known to lay eggs in very large batches of

over 100 on the underside of the leaves of the food plants. However, no early

stages of the species were observed.

The Dauan Island female is identified as E. blanda saraha as it meets the

descriptions in Common and Waterhouse (1981) and Parsons (1998) and is

consistent with the female E. blanda saraha (Fig. 3) from Papua New Guinea

illustrated in Braby (2016). This specimen is the first recorded female from

Queensland and the first recorded from Torres Strait since 1909.

d 3

Fig. 3. Eurema blanda saraha from Papua New Guinea (reproduced from Braby

2016, with permission).

Australian Entomologist, 2016, 43 (4) 237

Further observations and specimens are required to determine whether the

species is vagrant or established on Dauan Island. The abundant E. hecabe

may have contributed to the lack of confirmed observations previously;

however, the distinctly larger size of E. blanda compared with other Eurema

species, particularly E. hecabe and E. alitha, is an important distinguishing

attribute.

Acknowledgements

The authors would like to thank the people of Dauan Island and the Council

for the opportunity to collect on the island, and special thanks to Liz &

Wayne Phillips, Cr Gaidan (Chair) and Mario Soki for their help. Thanks also

to Peter Valentine for technical comments, Steve Brown for assistance with

identification and Peter Samson for specimen photography. Photos from

Braby (2016) are reproduced with permission from CSIRO Publishing.

References

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238 Australian Entomologist, 2016, 43 (4)

MISCELLANEOUS NOTES

The following note is abstracted from the News Bulletin of the Entomological

Society of Queensland and appeared, with illustrations, in the Volume and

Part indicated.

Acraea terpsicore (Linnaeus) (Lepidoptera: Nymphalidae). — The Tawny Coster is

now well established in the Kowanyama area on the Gulf of Carpentaria, Queensland

and represents a significant range extension from the recorded range in the Northern

Territory. It was first recorded in the Darwin area of the Northern Territory in April

2012 and had spread to Kakadu National Park, east Arnhem Land and near

Kalumburu in the Kimberley [Western Australia] by mid 2013 (Braby et al. 2014).

A recent field trip from 8-19 August 2016 to the lower Mitchell River catchment,

including Dunbar Station, Rutland Plains Station, Koolatah Station and the

community lands of Kowanyama, revealed that the Tawny Coster was abundant

throughout the area. Records range from very common (20 individuals/hour) at the

Mitchell River crossing at Dunbar Station, very common (20/half hour) feeding on

Melaleuca flowers at a creek crossing 2.7 km NNE of Koolatah Station, uncommon (4

in 6 hrs) on Rutland Plains Station from the homestead to 30 km south on the Nassau

River, and common (20 in 5 hrs) on Kowanyama community land from the coast 22

km west of the town to 30 km NE of it.

The very rapid expansion of the range of A. terpsicore from the Northern Territory to

Queensland over a very short period indicates that further expansion is likely

throughout Cape York Peninsula and the Gulf of Carpentaria (if not already there) to

the east coast over the next couple of years.

Reference: Braby, M.F., Thistleton, B.M. and Neal, M.J. 2014. Host plants, biology

and distribution of Acraea terpsicore (Linnaeus, 1758) (Lepidoptera: Nymphalidae): a

new butterfly for northern Australia with potential invasive status. Austral

Entomology 53: 288-297.

WILSON, P. — Tawny Coster arrives in Queensland. — Volume 44 (6): 121-122

(September 2016).

Australian Entomologist, 2016, 43 (4) 239

RECENT LITERATURE

BAEHR, B.C. and WHYTE, R.

2016. The Peacock Spiders (Araneae: Salticidae: Maratus) of the Queensland Museum,

including six new species. Zootaxa 4154(5): 501-525. (http://dx.doi.org/10.11646/

zootaxa.4154.5.1)

2016. The first described male tube-web spider for mainland Australia: Ariadna kiwirrkurra

sp. nov. (Araneae: Segestriidae). Zootaxa 4189(3): 595-599. (http://dx.doi.org/

10.11646/zootaxa.4189.3.11)

BRABY, M.F.

2016. The complete field guide to butterflies of Australia. 2nd edition. CSIRO Publishing,

Clayton South; 382 pp.

BROADLEY, A., KAUSCHKE, E. and MOHRIG, W.

2016. Revision of the types of male Sciaridae (Diptera) described from Australia by

F.A.A. Skuse. Zootaxa 4193(3): 401-450. (http://dx.doi.org/10.11646/zootaxa.

4193.3.1)

BROCK, P.D., MARSHALL, J.A., BECCALONI, G.W. and HARMAN, A.J.E.

2016. The types of Phasmida in the Natural History Museum, London, UK. Zootaxa 4179(2):

151-208. (http://dx.doi.org/10.11646/zootaxa.4179.2.1)

COLEMAN, D.

2016. Insect itineraries - from Sierra Leone, West Africa to Sydney, New South Wales.

Humanities Australia 7: 46-54.

CRANSTON, P.S., MARTIN, J., MULDER, M. and SPIES, M.

2016. Clarification of Einfeldia Kieffer, 1922 (Diptera: Chironomidae) with E. australiensis

(Freeman, 1961), comb. n. based on immature stages. Zootaxa 4158(4): 491-506.

(http://dx.doi.org/10.1 1646/zootaxa.4158.4.3)

DEVICHE, P. and ENDERSBY, I.

2016. First confirmed record of Black-headed Skimmer, Crocothemis nigrifrons, in Victoria.

Victorian Entomologist 46(3): 58-59.

EDWARDS, E.

2016. A chronological bibliography and digest of information about Zelotypia stacyi Scott,

1869, the bent-wing ghost moth. Victorian Entomologist 46(5): 96-99, 108-110.

FARROW, R.

2016. Insects of south-eastern Australia: an ecological and behavioural guide. CSIRO

Publishing, Melbourne; 280 pp.

FRANKLIN, D.C. and WOOD, G.

2016. Two sight records of the Orange-banded Plane (Lexias aeropa) from the Iron Range

area, and a note on food plants. North Queensland Naturalist 46: 53-56. Available

online.

HARRIS, K.

2016. Victorian Neuroptera. Part 3. Hemerobiidae, brown lacewings, and Nymphidae split-

footed lacewings. Victorian Entomologist 46(3): 50-57.

2016. Victorian Neuroptera. Part 4. Osmylidae, Berothidae, beaded lacewings. Victorian

Entomologist 46(5): 100-107.

240 Australian Entomologist, 2016, 43 (4)

HENDRICH, L., BALKE, M. and WATTS, C.H.S.

2016. Description of a new Tiporus Watts, 1985 from northern Queensland, Australia

(Coleoptera: Dytiscidae, Hydroporinae). Zootaxa 4189(1): 174-182. (Attp://dx.doi.org/

10.11646/zootaxa.4 189. 1.10)

HILL, L.

2016. Meyrickella ruptellus (Noctuidae: Hypeninae), a rare vagrant in Tasmania. Victorian

Entomologist 46(3): 60-66.

2016. An extreme rain event brings two rare vagrant moths to Tasmania. Victorian

Entomologist 46(4): 88-89.

HORAK, M. and KOMAI, F.

2016. Cryptophlebia Walsingham, 1900, Thaumatotibia Zacher, 1915, and Archiphlebia

Komai & Horak, 2006, in Australia (Lepidoptera: Tortricidae: Olethreutinae:

Grapholitini). Zootaxa 4179(3): 441-477. (http://dx.doi.org/10.11646/zootaxa.4179.3.5)

KLOPFSTEIN, S.

2016. Revising Australian Pristomerus (Hymenoptera, Ichneumonidae, Cremastinae): species

with a tooth on the hind femur. Zootaxa 4168(2): 201-238. (http://dx.doi.org/10.11646/

zootaxa.4168.2.1)

LESCHEN, R.A.B., ESCALONA, H.E. and ELGUETA, M.

2016. Phylogeny of the Gondwanan beetle family Ulodidae (Tenebrionoidea). Zootaxa

4138(3): 441-473. (http-//dx.doi.org/10.11646/zootaxa.4138.3.2)

MATSUI, J. and VALENTINE, P.

2016. A new southern record of the Golden Jezebel (Delias aruna) on Cape York Peninsula.

North Queensland Naturalist 46: 69-70. Available online.

MOESENEDER, C.H. and HUTCHINSON, P.M.

2016. Navigator, a new endemic genus of Cetoniinae (Coleoptera: Scarabaeidae) from

Australia, with descriptions of two new species and behavioural studies. Zootaxa

4173(1): 530-556. (http://dx.doi.org/10.11646/zootaxa.4173.6.2)

MONTE, C., ZILIOLI, M. and BARTOLOZZI, L.

2016. Revision of the Australian species of Figulus MacLeay, 1819 (Coleoptera: Lucanidae).

Zootaxa 4189(3): 447-484. (http://dx.doi.org/10.11646/zootaxa.4169.3.2)

ORR, A.G. and KALKMAN, V.J.

2016. Two new species of Papuagrion Ris, 1913 (Odonata: Coenagrionidae) from New

Guinea with a survey of distribution records for the genus. Zootaxa 4173(1): 18-28.

(http://dx.doi.org/10. 1 1646/zootaxa.4173.1.2)

RENTZ, D.C.F.

2016. An adventive cockroach new to Queensland: implications for the future. North

Queensland Naturalist 46: 47-52. Available online.

THEISCHINGER, G.

2016. “Bird of Paradise Crane Fly", Dapanoptera richmondiana in suburban Sydney.

Victorian Entomologist 46(4): 88-89.

WANG, Y., DIETRICH, C.H. and ZHANG, Y.

2016. Australnirvana, a new leafhopper genus of Nirvanini (Hemiptera: Cicadellidae:

Evacanthinae) from Australia. Zootaxa 4168(1): 134-140. (Attp://dx.doi.org/10.11646/

zootaxa.4 1668. 1.6)

THE

AUSTRALIAN

ENTOMOLOGIST

VOLUME 43

2016

Published by:

THE ENTOMOLOGICAL SOCIETY OF QUEENSLAND

THE AUSTRALIAN ENTOMOLOGIST

The Australian Entomologist is a non-profit journal published in four

parts annually by the Entomological Society of Queensland. The

journal is devoted to entomology of the Australian region, including

New Zealand, Papua New Guinea and islands of the south-western

Pacific. Articles are accepted from amateur and professional

entomologists. The journal is produced independently and subscription

is not included with membership of the Society.

The Editorial Panel

Editor: D.L. Hancock

Assistant Editors: G. Daniels

G.B. Monteith

L.W. Popple

F. Turco

Business Manager: G.B. Monteith

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Email contact: geoff.monteith ? bigpond.com

ISSN 1320-6133

Printed for The Entomological Society of Queensland

by Bayfield Printing, Brisbane

THE AUSTRALIAN ENTOMOLOGIST

Contents

Volume 43, 2016

BAEHR, M.

A new genus and species of the tribe Zolini from southeastern Australia

(Coleoptera: Carabidae: Zolini)

A new species of the Tachys (s.1.) ectromoides-group from southeastern Queensland,

Australia (Coleoptera: Carabidae: Bembidini)

BATLEY, M., PAULY, A., GOLLAN, J.R., ASHCROFT, M.B. and SONET, G.

Re-identification of an exotic bee introduced to the Hunter Valley Region, New South

Wales — Seladonia hotoni (Vachal, 1903) (Hymenoptera: Halictidae)

BRABY, M.F.

Migration records of butterflies (Lepidoptera: Papilionidae, Hesperiidae, Pieridae,

Nymphalidae) in the ‘Top End’ of the Northern Territory

CLAYTON, J.

The genera Eoophyla Swinhoe and Nymphicula Snellen (Lepidoptera: Crambidae:

Acentropinae) in Fiji, with descriptions of four new species

DOWNES. M.F.

New southern range limit for Polyrhachis yorkana Forel (Hymenoptera: Formicidae),

with provisional data on castes

DOWNES, M.F. and EDWARDS, E.

An undescribed concealer moth, Stathmopoda sp. (Lepidoptera: Oecophoridae) in

nests of the weaver ant Polyrhachis australis Mayr (Hymenoptera: Formicidae)

DOWNES, M.F. and HARVEY, M.

The spider Oecobius concinnus Simon (Araneae: Oecobiidae) in nests of the ant

Polyrhachis australis Mayr (Hymenoptera: Formicidae) in Townsvillle, Queensland:

A new distribution record

DREW, R.A.I. and HANCOCK, D.L

A review of the subgenus Bulladacus Drew & Hancock of Bactrocera Macquart (Diptera:

Tephritidae: Dacinae), with description of two new species from Papua New Guinea

HANCOCK, D.L.

A review and keys to genera and some species of the fruit fly tribes Phytalmiini,

Phascini and Epacrocerini (Diptera: Tephritidae: Phytalmiinae)

HANCOCK, D.L. and DREW, R.A.I.

A review of the subgenus Austrodacus Perkins of Bactrocera Macquart (Diptera:

Tephritidae: Dacinae)

LAMBKIN, T.A.

First Australian records of Euploea wallacei melia Fruhstorfer, 1904 and Euploea

stephensii jamesi Butler, 1876 (Lepidoptera: Nymphalidae: Danainae) from Dauan

Island, Torres Strait, Queensland, with notes on the aggregation habits of Euploea

Fabricius species near flowering mangroves

LAMBKIN, T.A. and KENDALL, R.

The status of Yoma algina (Boisduval, 1832) and Y. sabina (Cramer, 1780)

(Lepidoptera: Nymphalidae: Nymphalinae) in Australia

MACKEY, A.P.

Review of the Australian species of Arctornis Germar, 1810 (Lepidoptera: Erebidae:

Lymantriinae)

iii

109

151

135

161

189

211

173

MATE-NANKERVIS, J.F.

Bambara testacea (Britten, 1926), an adventive featherwing beetle (Coleoptera:

Ptiliidae) new to the Australian fauna 5

MEYER, C.E., MILLER, C.G., BROWN, S.S. and WEIR, R.P.

First Australian record of Parthenos tigrina (M. Snellen van Vollenhoven, 1886)

(Lepidoptera: Nymphalidae: Nymphalinae) 113

MOLLET, B. and TARMANN, G.M.

A new organ in male zygaenid moths (Lepidoptera: Zygaenidae) 83

POPPLE, L.W.

A new species of Macrotristria Stål from the Shoalwater Bay region of central

Queensland (Cicadidae: Cicadinae) 101

SANDS, D.P.A., GRIMSHAW, P. and SANDS, M.C.

Acacia melvillei Pedley (Mimosaceae), a newly-recorded larval food plant for

Jalmenus eubulus Miskin (Lepidoptera: Lycaenidae) 165

TIKOCA, S., CLAYTON, J., HODGE, S, TUIWA WA, M., PENE, S. and BRODIE, G.

A record of Dactyloplusia impulsa (Walker, 1865) (Lepidoptera: Noctuidae:

Plusiinae) new to Fiji 35

WELLS, A. and KJER, K.

Norfolk Island's caddisfly is a New Zealander (Trichoptera: Hydroptilidae) 49

WILLIAMS, A.A.E. and WILLIAMS, M.R.

A new species of Synemon Doubleday (Lepidoptera: Castniidae) from Western Australia 119

WILSON, G.W. and RENTZ, D.C.F.

First observation in nature of effective predator defence behaviour in the spiny forest

katydid Phricta spinosa Redtenbacher (Orthoptera: Pseudophyllinae: Phrictini) 1

WILSON, P.R.

Notes on the pupa and host plant of Motasingha dirphia (Hewitson) (Lepidoptera:

Hesperiidae) 9

WILSON, P.R. and JOHNSON, I.

Eurema blanda saraha (Fruhstorfer) (Lepidoptera: Pieridae: Coliadinae) rediscovered

In Torres Strait, Queensland, Australia 235

MISCELLANEOUS NOTES 238

BOOK NOTICE 38

BOOK REVIEWS 46, 172

RECENT LITERATURE 88, 239

CORRIGENDA 108

Publication dates: Part 1 (pp 1-48) 22 March 2016

Part 2 (pp 49-108) 24 June 2016

Part 3 (pp 109-172) 3 September 2016

Part 4 (pp 173-240) 12 December 2016

ENTOMOLOGICAL NOTICES

Items for insertion should be sent to the Business Manager who reserves the

right to alter, reject or charge for notices.

NOTES FOR AUTHORS

Manuscripts submitted for publication can be submitted as either hardcopies

or electronically. Three copies (double spaced text and illustrations) of

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format should be sent in Microsoft Word. Digital illustrations should be sent

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for each figure. High resolution TIFFs or JPEGs (300 dpi at print size) must be

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and digital manuscripts on disc should be sent to:

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The Australian Entomologist

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Indooroopilly, Qld, 4068

Authors should refer to recent issues for layout and style. All papers will be

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considered unsuitable.

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mandatory provisions of the International Code of Zoological Nomenclature.

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Printed by Bayfield Printing, Unit 6/60 Kremzow Rd, Brendale, Q 4500, Ph: 1300 685 820

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THE AUSTRALIAN

Entomologist

Volume 43, Part 4, 12 December 2016

DREW, R.A.I. and HANCOCK, D.L.

A review of the subgenus Bulladacus Drew & Hancock of

Bactrocera Macquart (Diptera: Tephritidae: Dacinae), with

description of two new species from Papua New Guinea

LAMBKIN, T.A. and KENDALL, R.

The status of Yoma algina (Boisduval, 1832) and Y. sabina

(Cramer, 1780) (Lepidoptera: Nymphalidae: Nymphalinae) in

Australia

MACKEY, A.P.

Review of the Australian species of Arctornis Germar, 1810

(Lepidoptera: Erebidae: Lymantriinae)

WILSON, P.R. and JOHNSON, I.

Eurema blanda saraha (Fruhstorfer) (Lepidoptera: Pieridae:

Coliadinae) rediscovered in Torres Strait, Queensland, Australia

MISCELLANEOUS NOTES

RECENT LITERATURE

CONTENTS VOLUME 43, 2016

ISSN 1320 6133

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