Australian Entomologist, 2015, 42 (3): 105-106 105

TWO NEW RECORDS OF AUSTRALIAN TRIATOMA LAPORTE

(HEMIPTERA: REDUVHUDAE: TRIATOMINAE)

G.B. MONTEITH

Queensland Museum, PO Box 3300, South Brisbane. Old 4101

Abstract

Two additional specimens of Triatoma leopoldi (Schouteden, 1933) are recorded from Australia

together with photographs of a living specimen.

Introduction

Monteith (1974) recorded three males of Triatoma novaeguineae Miller,

1958 taken at mercury vapour light on the banks of the East Claudie River

(12.714°S, 143.287°E), Iron Range area (now within Kutini-Payamu NP) in

Cape York Peninsula, that being the first record of this medically important

group of blood-feeding insects from Australia.

Me T e

K Je "è iòs z ait te P

Figs 1-2. Living male of Triatoma leopoldi (Schouteden) collected at East Claudie

River in December 2010: (1) dorsal view; (2) lateral view showing rostrum — R.

(Photographs by Eric Vanderduys).

106 Australian Entomologist, 2015, 42 (3)

Monteith (1974) also suggested that the species may be a junior synonym of

Triatoma leopoldi (Schouteden, 1933) and this was implemented in the world

revision of the subfamily by Lent and Wygodzinsky (1979). Outside

Australia, the species is known from the islands of New Guinea, Biak and

Sulawesi (Lent and Wygodzinsky 1979, Maldonado Capriles 1990). All

Triatominae are vertebrate blood feeders (Lent and Wygodzinsky 1979, Justi

et al. 2014) and the listing of the species as ‘arthropod-feeder, predator’ by

Cassis and Gross (1995) is a lapsus.

A recent review of the potential for transmission of Chagas’ disease (caused

by the parasitic protozoan Trypanosoma cruzi Chagas) in Australia (Jackson

et al. 2014) estimated 1,928 infected residents in 2011 and noted the presence

of T. leopoldi as a potential Australian vector.

New records

Two more male specimens have been collected at MV light in the Iron Range

area, both lodged in the Queensland Museum, as follows: East Claudie River,

12.714°S, 143.287°E, 8-16.x11.2010, Monteith & Escalona; West Claudie

River, 12.739°S, 143.256°E, 28.x1.1986, G. Daniels & M. Schneider. The

West Claudie River locality is 4 km SW of the East Claudie locality. Both are

in lowland rainforest. Females have yet to be collected in Australia. The 2010

specimen was photographed alive by Eric Vanderduys and these are

reproduced with thanks to enable collectors to recognise this important

species (Figs 1-2). The distinctive colour and straight rostrum held close to

the underside of the head (Fig. 2) allows separation from other Australian

assassin bugs which have the rostrum curved away from the head underside.

Males are 19-20 mm in length. Overseas, females reach 23 mm.

References

CASSIS, G. and GROSS, G.F. 1995. Hemiptera: Heteroptera (Coleorrhyncha to Cimicomorpha).

In: Houston, W.W.K. and Maynard, G.V. (eds), Zoological Catalogue of Australia. Volume

27.3A. CSIRO, Melbourne; 501 pp.

JACKSON, Y., PINTO, A. and PETT, S. 2014. Chagas’ disease in Australia and New Zealand:

risks and need for public health interventions. Tropical Medicine and International Health 19(2):

212-218.

JUSTI, S.A., RUSSO, C.A.M., MALLET, J.R. dos S., OBARA, M.T. and GALVÃO, C. 2014.

Molecular phylogeny of Triatomini (Hemiptera: Reduviidae: Triatominae). Parasites & Vectors

7(149): 12 pp. Published online — doi:10.1186/1756-3305-7-149.

LENT, H. and WYGODZINSKY, P. 1979. Revision of the Triatominae (Hemiptera, Reduviidae)

and their significance as vectors of Chagas’ disease. Bulletin of the American Museum of Natural

History 163: 125-520.

MALDONADO CAPRILES, J. 1990. Systematic catalogue of the Reduviidae of the world

(Insecta: Heteroptera). University of Puerto Rico, Puerto Rico; 694 pp.

MONTEITH, G.B. 1974. Confirmation of the presence of Triatominae (Hemiptera: Reduviidae)

in Australia, with notes on Indo-Pacific species. Journal of the Australian Entomological Society

13: 89-94.

Australian Entomologist, 2015, 42 (3): 107-126 107

A REVIEW OF THE TREE, FIG AND FRUIT-INFESTING FLIES

OF THE AETHIOTHEMARA, DIARRHEGMA, DIRIOXA AND

THEMAROIDES GROUPS OF GENERA (DIPTERA: TEPHRITIDAE:

ACANTHONEVRINI)

DAVID L. HANCOCK

8/3 McPherson Close, Edge Hill, Cairns, Old 4870

Abstract

Indo-Australian, East Asian and African fruit flies placed in the Aethiothemara, Diarrhegma,

Dirioxa and Themaroides groups of genera and known or believed to breed in figs, overripe and

damaged fruit or beneath the bark of standing trees are reviewed. The 134 recognised species are

referred to 39 genera: Aethiothemara Hendel, Afrocneros Bezzi, Labeschatia Munro, Ocnerioxa

Speiser, Ptiloniola Hendel and Themarictera Hendel in the Aethiothemara group; Diarrhegma

Bezzi in the Diarrhegma group; Anchiacanthonevra Hardy, Dirioxa Hendel, Griphomyia Hardy,

Lumirioxa Permkam & Hancock, Micronevrina Permkam & Hancock, Mimoeuphranta Hardy

and Parachlaena Hering in the Dirioxa group; and Acanthonevroides Permkam & Hancock,

Alloeomyia Hardy, Aridonevra Permkam & Hancock, Buloloa Hardy, Cheesmanomyia Malloch,

Clusiosoma Malloch, Clusiosomina Malloch, Enoplopteron de Meijere, Hemiclusiosoma Hardy,

Hexaresta Hering, Neothemara Malloch, Nothoclusiosoma Hardy, Paedohexacinia Hardy,

Pseudacanthoneura Malloch, Pseudoneothemara Hardy, Quasirhabdochaeta Hardy, Rabaulia

Malloch, Rabauliomorpha Hardy, Taeniorioxa Permkam & Hancock, Termitorioxa Hendel,

Themarohystrix Hendel, Themaroides Hendel, Themaroidopsis Hering, Trypanocentra Hendel

and Walkeraitia Hardy in the Themaroides group. Lyronotum Hering, 1941 is placed as a new

synonym of Hexaresta Hering, 1941 and its type species, Acanthoneura seriata de Meijere,

1915, is placed as a new synonym of Hexaresta multistriga (Walker, 1859). Clusiosoma

semifuscum Malloch, 1926 is returned to synonymy with C. minutum (de Meijere, 1913). Keys

are included to Asian and Australasian genera plus species of Aethiothemara, Diarrhegma,

Dirioxa, Termitorioxa and Themaroides and males of the Clusiosoma pullatum group.

Introduction

This is the fifth in a series of papers reviewing and keying Indo-Australian,

Oceanian and East Asian fruit flies placed in the tribe Acanthonevrini sensu

Korneyev (1999) of subfamily Phytalmitnae and deals with species referred

to the Diarrhegma, Dirioxa and Themaroides groups of genera, together with

the Afrotropical Aethiothemara group. Diarrhegma occurs from India and

China to Indonesia, while the Dirioxa and Themaroides groups are largely

restricted to the Australian and Oceanian Regions. Previous papers dealt with

the Acanthonevra, Sophira, Rioxa and Dacopsis complexes of the

Acanthonevra group (Hancock 2011, 2012, 2014a, 2014b).

The Afrotropical Aethiothemara group was reviewed and keyed by Hancock

(1986) and contains six genera: Aethiothemara Hendel, Afrocneros Bezzi,

Labeschatia Munro, Ocnerioxa Speiser, Ptiloniola Hendel and Themarictera

Hendel. Keys to the species of Afrocneros (3 spp), Ocnerioxa (11 spp) and

Ptiloniola (3 spp) were provided by Munro (1967), while Labeschatia and

Themarictera are monotypic (Hancock 1986). A provisional key to the six

species of Aethiothemara, based on that of Hendel (1928), is provided below.

Larvae of Afrocneros excellens (Loew), A. mundus (Loew) and Ocnerioxa

sinuata (Loew) feed on the parenchymatous tissue of standing Cussonia

108 Australian Entomologist, 2015, 42 (3)

(Araliaceae) trunks (Munro 1967), while Themarictera flaveolata (Fabricius)

has been bred from fruit of Maerua cafra and ‘Capparis sp.’ (Capparaceae)

(Munro 1925, Hancock 2003).

For the Indo-Australian fauna, the Dirioxa group differs from the

Diarrhegma and Themaroides groups in having only a single apical spine on

the mid tibia rather than two. The Diarrhegma group contains only the genus

Diarrhegma Bezzi, with two species. Diarrhegma modestum (Fabricius) has

been reported from decaying wood (Bezzi 1913), including Moringa

pterygosperma (Moringaceae) (Hancock and Drew 1994b), and at least some

genera in the Dirioxa and Themaroides groups (e.g. Lumirioxa Permkam &

Hancock and Termitorioxa Hendel) are known to breed beneath the decaying

bark of standing trees (Permkam and Hancock 1995, Hancock 2002). Adults

of Acanthonevroides Permkam & Hancock, Lumirioxa and Micronevrina

Permkam & Hancock have been collected in Malaise traps baited with sawn

saplings (G.B. Monteith in Hancock 2013).

The Themaroides group is currently divided into three — the Clusiosoma

subgroup with nine genera, Neothemara subgroup with six genera and

Themaroides subgroup with ten genera (Hancock and Drew 2003). In the

Clusiosoma subgroup, larvae of Cheesmanomyia nigra (de Meijere),

Clusiosoma minutum (de Meijere) [as C. semifuscum Malloch], C. pleurale

Malloch, C. subpullatum Hardy, Clusiosomina puncticeps Malloch, Rabaulia

fascifacies Malloch, R. nigrotibia Hering and Trypanocentra_ nigrithorax

Malloch all feed in the fruit of figs (Ficus spp: Moraceae) (Hardy 1986,

Permkam and Hancock 1995, Hancock et al. 1998, Hancock et al. 2000,

Hancock and Drew 2003). In the Themaroides subgroup, larvae of

Termitorioxa termitoxena (Bezzi) feed beneath damaged bark of trees such as

Terminalia sp. (Combretaceae) and Delonix regia (Fabaceae); an initial

record from galleries of Mastotermes darwiniensis Froggatt (Bezzi 1919) is

believed to indicate feeding on the tree trunk rather than feeding on the

termites (Hancock 2002). Acanthonevroides nigriventris (Malloch) has been

collected on trunks of Acacia and Eucalyptus and attracted to sawn saplings

and all species in the genus are likely to be tree trunk feeders (Permkam and

Hancock 1995, Hancock 2013). Larval hosts have not been recorded for any

species in the Neothemara subgroup.

The New Guinea-Pacific genera Anchiacanthonevra Hardy, Mimoeuphranta

Hardy and Parachlaena Hering (Fig. 1) were transferred from the

Acanthonevra group to the Dirioxa group by Hancock (2012), which also

includes Dirioxa Hendel, Lumirioxa and Micronevrina. The New Guinea

genus Griphomyia Hardy was initially placed in tribe Trypetini by Hardy

(1987) but the presence of secondary scutellar and intrapostalar setae,

together with the setose aculeus, suggest it is better placed in the

Acanthonevrini. Its precise relationships remain obscure but the distinct

intrapostalar setae, single midtibial spine and shape of the spermathecae

Australian Entomologist, 2015, 42 (3) 109

suggest it is best placed in the Dirioxa group, at least provisionally. It differs

from other acanthonevrine genera in the bare or micropubescent, rather than

plumose, arista. Larvae of Lumirioxa araucariae (Tryon) feed within a wet

rot beneath the bark of Araucaria cunninghamii trees (Araucariaceae)

(Brimblecombe 1945). Larvae of Dirioxa pornia (Walker) attack a wide

range of overripe or damaged fruit and even fallen Araucaria cones

(Permkam and Hancock 1995, Hancock et al. 2000), while those of D.

fuscipennis Hancock & Drew have been bred from the fruit of Barringtonia

edulis (Lecythidaceae) and Poutaria guayana (Sapotaceae) (Hancock and

Drew 2003). The hosts of other genera in the group remain unknown,

although the attraction of Micronevrina hyalina Permkam & Hancock and M.

montana Permkam & Hancock to sawn saplings (Hancock 2013) suggests

they also breed in trees.

Fig. 1. Parachlaena greenwoodi (Bezzi): male from Fiji.

110

Australian Entomologist, 2015, 42 (3)

Two further East or Southeast Asian genera sometimes included in the

Acanthonevrini, Hexaptilona Hering and Paracanthonevra Hardy, belong in

subfamilies Blepharoneurinae or Trypetinae respectively (Korneyev 1999,

Han 1999) and are not included here.

Key to Asian and Australasian genera

Scutum largely ivory-white posteriorly and with black patches antero- and

posterolaterally; scutellum ivory-white and unspotted, wing with

pterostigma about half length of cell c; head with 2 pairs each of frontal

and orbital setae; mid tibiae with 2 long apical spines [S and SE Asia; 2

species, keyed below] .... Diarrhegma group ... Diarrhegma Bezzi, 1913

Not as above; scutum and scutellum fulvous to black without tvory-white

markings; other characters variable [Australian and Pacific Regions] .... 2

Mid tibia with only 1 long apical and no distinct, shorter spines; wing

vein Cu, and Cu base both bare; cell m with 1 or (rarely) 2 hyaline

jes LEOLE 1016) RAAS AE PENAN en a L, ETE E Dirioxa group ... 3

Mid tibia with 2 long and subequal or 1 long and 1 or more shorter apical

spines, wing vein Cu, and/or Cu base often setulose; cell m with 0-3

hyaline indentations ....................sassasassaaaa Themaroides group ... 9

Arista bare or micropubescent; upper orbital seta weak or absent; 1 pair of

frontal setae; R-M crossvein placed near basal third of cell dm and below

or before middle of pterostigma; cell bcu extension vestigial [Papua New

Guinea (mainland and Bougainville); 5 spp, keyed and illustrated by

Arey IBT] 28 en! Al le ðr cence cis ates Ma aa Griphomyia Hardy, 1987

Arista plumose; upper orbital seta often well developed; 1 or 2 pairs of

frontal setae; R-M crossvein placed near or beyond middle of cell dm and

usually below or beyond apex of pterostigma; cell bcu extension distinct

Wing cell m with 2 distinct posterior hyaline indentations; apex of cell

rs hyaline except for a medial dark band or spot from wing margin

[Papua New Guinea (New Britain); 1 sp., illustrated by Hardy 1986] ......

LN Peete OTe SOR En Oe Tir Anchiacanthonevra Hardy, 1986

Wing cell m largely hyaline to subhyaline or with 1 distinct posterior

indentation; apex of cell 1445 NOt aS above .................sssisaaasisssaasaani 5

Upper orbital seta vestigial or absent; 1 pair of frontal setae; wing apex

hyaline across posterior half of cell 1243 and all of cell 1745 [Fiji; 1 sp,

illustrated in Fig. 1] .....0....00 00. eee Parachlaena Hering, 1944

Upper orbital seta normally distinct; 1 or 2 pairs of frontal setae; wing

apex not hyaline across posterior half of cell r2+, and all of cell 1445 ...... 6

Australian Entomologist, 2015, 42 (3) 111

6 Upper orbital seta weak, much shorter than lower seta; prescutellar

acrostichal setae absent; R-M crossvein placed below middle of

pterostigma; wing pattern largely hyaline to subhyaline with a narrow,

dark transverse band from costa over DM-Cu crossvein to posterior

margin [Papua New Guinea and Solomon Islands; 1 sp., illustrated by

Hardy 1986] ..........................u................ Mimoeuphranta Hardy, 1986

— Upper orbital seta well developed, almost as long as lower seta;

prescutellar acrostichal setae present or absent; R-M crossvein placed

below or beyond apex of pterostigma; wing pattern dark, usually with

numerous hyaline spots and indentations .................sssasssssssaaaaaani 7

7 Anepimeral seta vestigial or absent; usually only 2 pairs of scutellar setae,

if 3 then medial pair weak and rudimentary; presutural setae present or

absent, if present then wing pattern distinctly sexually dimorphic; aculeus

apically dentate and bifid, the preapical setae minute [eastern Australia; 7

spp, keyed and illustrated by Permkam and Hancock 1995] ..................

abiha bitentats dA e R RA Puke Micronevrina Permkam & Hancock, 1995

— Anepimeral seta distinct; 3 pairs of scutellar setae, the medial pair at last

half length of apicals, presutural setae present and wing pattern not

sexually dimorphic; aculeus apically blunt and non-serrate, the preapical

sëtae Wwelldeyeloped sei etie hh hús, ae eet aa creed E a eon ii et 8

8 Thorax and scutellum with a black medial stripe; fore femur densely

setose ventrally in males; wing cell ry; with an elongate hyaline

indentation posteroapically [eastern Australia (seQld-neNSW); 1 sp.,

illustrated by Permkam and Hancock 1995 and in Fig. 2] ....................

PAAPA NWA d BOREAS ELM) RRO SIAP Lumirioxa Permkam & Hancock, 1995

— Thorax and scutellum fulvous, without a black medial stripe; fore femur

not densely setose ventrally in males; wing cell r4,5; entirely dark apically

[eastern Indonesia (Papua Province), Australia, New Caledonia and

Vanuatu; 3 spp, keyed below] .......................... Dirioxa Hendel, 1928

9 Wing cell m with at most a single posterior hyaline indentation ......... 10

— Wing cell m with 3 posterior hyaline indentations .............................

ie NAN eee AA ah r eke ASAA edn aan Tete tos Neothemara subgroup ... 11

10 Mid tibia with 2 long and subequal apical spines; scutellum bare or

densely setulose; scutum often with 2 pairs of supra-alar setae; wings

usually with hyaline spots or indentations, if largely brown and scutellum

not densely setulose then with an additional 2 pairs of secondary scutellar

Beta Sak AL Mot Nk OE Bee MeN! Themaroides subgroup ... 16

— Mid tibia normally with 1 long and 1 or more shorter apical spines;

scutellum bare or at most weakly setulose marginally; scutum with only 1

pair of supra-alar setae; wings largely brown without hyaline spots or

112 Australian Entomologist, 2015, 42 (3)

indentations in at least anterior half; scutellum without additional pairs of

Secondary setae ................sissssssaaaiaaaasaa Clusiosoma subgroup ... 25

Fig. 2. Lumirioxa araucariae (Tryon): female from SE Queensland. Photograph by

Andy Wang (Queensland Museum).

11 Vein R+, curved sharply from level of DM-Cu crossvein to meet costa at

an angle of about 60°; wing pattern with numerous brown rays extending

to wing margin in cells 11, 1243 and 144; [Papua New Guinea (New Britain);

1 sp., illustrated by Hardy 1986]......... Quasirhabdochaeta Hardy, 1986

— Not as above; vein R»; not sharply curved towards costa and pattern

without dark marginal rays ..............iaiiaianasaaasasaanaaasaaaasaskassaaaa 12

12 Face vertical in profile, the epistome receding ........................ 2.2... 13

— Face concave in profile, the epistome projecting ........................... 14

13 Ocellar setae distinct and well developed; vein R4s distinctly curved

subapically then straightening to costa [eastern Indonesia (Papua

Province), Papua New Guinea and Australia (nQId); 2 spp, illustrated by

Hardy 1986 and Permkam and Hancock 1995] l..

RV Pak N E Benak K R TERNA Pseudacanthoneura Malloch, 1939

Australian Entomologist, 2015, 42 (3) 113

— Ocellar setae weak and hair-like; vein Ry; curving evenly to costa

[eastern Indonesia (West Papua and Papua Provinces), Papua New Guinea

and Australia (nQId); 2 spp, keyed by Hardy 1986; N. trigonifera Hering,

1951 was placed as a synonym of N. formosipennis (Walker, 1861) by

Permkam and Hancock 1995; illustrated by Hardy 1986 and Permkam

and Hancock 1995] ..............2.2.ssssssssssviiii Neothemara Malloch, 1939

14 Anepimeral seta vestigial; scutum yellow with only a small black spot

behind each wing base; mid tibia with apical spine scarcely longer than

small secondary spinules [Papua New Guinea; | sp., illustrated by Hardy

LIZO nin Sweats Ae khi bansa b ðD t Alloeomyia Hardy, 1986

— Anepimeral seta well developed; scutum with distinct dark markings; mid

tibia with 1 long and 1 shorter apical spines .....................2222aaniiaai 15

15 Vein Cu with basal portion along cell bcu setulose; wing with an oblique

hyaline band from apex of vein Cu2+A; to vein R4+s above line of DM-Cu

crossvein; scutellum with short black setulae at sides [eastern Indonesia

(Batjan), Papua New Guinea (New Britain, New Ireland, Bougainville)

and Solomon Islands; 2 spp, illustrated by Hardy 1986] ......................

STI R Matte ald ee Mea E VT Bask sted Pseudoneothemara Hardy, 1986

— Vein Cu bare, including basal portion; wing without an oblique hyaline

band from apex of vein Cu+A; to vein R4s above line of DM-Cu

crossvein; scutellum without marginal black setulae [eastern Indonesia

(Sulawesi, Aru, West Papua, Papua), Papua New Guinea and Solomon

Islands; 2 spp, illustrated by Hardy 1986; Lyronotum Hering, 1941, syn.

n. and Hyponeothemara Hardy, 1986 are both regarded as synonyms;

Acanthoneura seriata de Meijere, 1915, syn. n., the type species of

Lyronotum, is inseparable from Trypeta multistriga Walker, 1859, the

type species of Hexarestd] ...............sssssiaaaaii Hexaresta Hering, 1941

16 Wing with a row of long costal spines between humeral vein and apex of

Cel as, «Fei oe gS), r ok a Behe s oom Sel T Mos hehe 4. T 17

— Wing without a row of long costal spines between humeral vein and apex

GL Cel n eet a nh atte ha eats cette ae an ee e EOC L a te AS 18

17 Vein Cu base and Cu, both setulose; wing pattern not distinctly reticulate

[Papua New Guinea; 1 sp., illustrated by Hardy 1986] ........................

bee rac am RM, SU Sal R Tr. Jóla HR VASK JE Buloloa Hardy, 1986

— Vein Cu base and Cu, both bare; wing pattern distinctly reticulate on disc

[eastern Indonesia (Papua Province) and Papua New Guinea (mainland

and New Britain); 3 spp, keyed and illustrated by Hardy 1986] ............

sonia te act haste thas PROS a Saas reich Enoplopteron de Meijere, 1915

18 Wing with a complete hyaline transverse band from middle of costal cell

c and enclosing BM-Cu crossvein to posterior margin of cell cui, plus

incomplete transverse bands in cells r,+r243+br and 14+5+dm; apex of cell

114

Australian Entomologist, 2015, 42 (3)

r4s and cell m with elongate hyaline indentations; vein R4s strongly

curved anteriorly beyond R-M crossvein; vein M strongly curved

posteriorly before R-M crossvein so that cell dm is distinctly petiolate in

basal half [eastern Indonesia (West Papua Province); | sp., illustrated by

Hardy 1986] .......iiaiiiiisiiaaaiaasssaaadsaaaaakaaaa Walkeraitia Hardy, 1986

Wing pattern not as above, if banding evident then BM-Cu crossvein lies

within the dark area; veins Ry,; and M not both strongly curved ......... 19

19 Wing pattern dark with base largely hyaline and 5 narrow hyaline bands

not arising from costa: 2 longitudinal in cells rj+1o43 and 14; and 3

obliquely transverse in cells m, br+dm+cu, and cu,;; DM-Cu crossvein

inwardly oblique and curved towards wing base in anterior half [Australia

(seQld); 1 sp., illustrated by Permkam and Hancock 1995] ..................

ues Neen edt Bey Bic bal Wee Pee ce ale, Taeniorioxa Permkam & Hancock, 1995

Wing pattern not as above ...........isiiiaaniiassaaaaasaaasssaaaaaksssaaaaaaaa 20

20 Wing pattern with costal cell c hyaline medially and 4 incomplete,

obliquely transverse hyaline bands: an indentation in cells r)+143, a

curved band in cells r4,;+m, an apical band in cell dm and a narrow basal

band along vein Cuj+A, in cell cu, DM-Cu crossvein strongly outwardly

oblique, its apex situated below the apex of vein R+ at costa; scutellum

black and swollen, with 4 scutellar setae and dense setulae [Australia

(swQld); 1 sp., illustrated by Permkam and Hancock 1995] .................

cen ntbee aca Eð Ta Aridonevra Permkam & Hancock, 1995

Wing pattern not as above; DM-Cu crossvein not strongly outwardly

oblique, its apex situated basad of the apex of vein R»+3; if only 4 scutellar

setae then scutellum yellow medially ..................ssssssaassssaaaaaanaai 21

Vein M setulose along cell dm ...............niiniaiiiaaaaaaasaaaaaaaassaaaaaa 22

Vein M Dare gh Lc ee JL aE SR A A, a E e NE B ae. BE 23

22 Veins Cu base and Cu, setulose; wings normally dark brown with no

more than the posterior area hyaline; if base broadly hyaline then hyaline

patch from cell dm into cell 145 not extending into cell rə}, [eastern

Indonesia (Misool, West Papua, Papua) and Papua New Guinea; 9 spp,

keyed and illustrated by Hardy 1986]....... Themarohystrix Hendel, 1914

Veins Cu base and Cu, bare; wing base broadly hyaline; if hyaline patch

from cell dm into cell r4.; present then not extending into cell 12; [eastern

Indonesia (Papua Province) and Papua New Guinea (mainland and

Bougainville); 4 spp, keyed and illustrated by Hardy 1986] ...............

EE SLL LEO! Ok OE LE k Pind Oe Themaroidopsis Hering, 1941

23 Wing with anterior half brown beyond cell sc, if a quadrate hyaline

indentation present in cells 1;+1243, then cells br and r4+s without medial

hyaline spots and scutellum entirely setulose; head with lower orbital

Australian Entomologist, 2015, 42 (3) 115

setae placed below middle of frons [eastern Indonesia (Ambon, West

Papua, Papua) and Papua New Guinea (mainland and New Britain); 6

spp, keyed below; Rioxina Hering, 1941 is a synonym (Hardy 1986)] ...

MT M A E a bags M onia Themaroides Hendel, 1914

— Wing with a triangular hyaline indentation from costa in cell rı or ry+1243;

cells br and r4,; with medial hyaline spots present, if not then scutellum

only laterally setulose; head with lower orbital setae placed above middle

of frons, usually in upper quarter ...... 2.006... cece eee cece eens 24

24 Wing pattern blackish brown; scutellum black with a medial yellow band,

swollen and densely setulose [Australia; 5 spp, keyed and illustrated by

Permkam and Hancock 1995 and in Fig. 3; the South Australian A.

bicolor (Macquart, 1855) is a synonym of A. basalis (Walker, 1853)

(Hancock 1998)] ............ Acanthonevroides Permkam & Hancock, 1995

— Wing pattern brown and yellow-brown; scutellum pale with at most basal

dark patches, not swollen and setulose only laterally [eastern Indonesia

(Timor, Misool and Papua Province), Papua New Guinea and Australia);

11 spp, keyed below; Kertesziola Hering, 1941 is a synonym (Permkam

and Hancock 1995)] ..........iiiiissiasaaasaaaaii Termitorioxa Hendel, 1928

1mm

Fig. 3. Acanthonevroides jarvisi (Tryon): male from SE Queensland. Photograph by

Andy Wang (Queensland Museum).

116 Australian Entomologist, 2015, 42 (3)

25 Wing veins Cu base and Cu, bare; anepisternum with a prominent black

seta near lower medial margin [Papua New Guinea and Australia (neQld);

2 spp, keyed and illustrated by Permkam and Hancock 1995 and

illustrated by Hardy 1986] ..................... Paedohexacinia Hardy, 1986

— Wing vein Cu base setulose; vein Cu, bare or setulose; anepisternum

without a prominent black seta near lower medial margin ................ 26

26 Wing vein Ri extending to beyond level of DM-Cu crossvein, the

pterostigma much longer than cell c [eastern Indonesia (Papua Province)

and Papua New Guinea; 1 sp., illustrated by Hardy 1986] ...................

seh E ð O Bul iy eRe beh note EEO Cheesmanomyia Malloch, 1939

— Wing vein R; ending near level of R-M crossvein, the pterostigma no

longer than cell c and normally much shorter ........................... a... 27

27 Face concave in profile, the epistome protruding; male fore tibia with a

densely white-pilose, pad-like posteroapical process [2 subgenera] ......

sae dy Tre eara da Abe detest pets Clusiosoma Malloch, 1926 ... 28

— Face vertical or convex in profile, the epistome not protruding .......... 29

28 Scutum black; face black; parafacials yellow; male fore femur with a

spinose basoventral tubercle but without ventral spines [Papua New

Guinea and Australia (nQld); 1 sp., illustrated by Hardy 1986 and

Permkam and Hancock 1995] ..... subgenus Paraclusiosoma Hardy, 1986

— Scutum partly fulvous; if face black then parafacials also black; male fore

femur with long ventral spines but no tubercle [eastern Indonesia

(Ambon, Waigeo, West Papua and Papua), Papua New Guinea (mainland,

New Britain, New Ireland), northern Australia and Solomon Islands; 14

spp, keyed and illustrated by Hardy 1986 (non-Australian species) and

Permkam and Hancock 1995 (Australian species); C. minutum (de

Meijere, 1913) appears to have the first and second antennal segments

black, not yellow as implied by Hardy (1986) and C. semifuscum

Malloch, 1926, syn. rev., separated by Permkam and Hancock 1995, is

returned to synonymy (as in Hardy 1986); for separation of pullatum

group males, see key below] ......... subgenus Clusiosoma Malloch, 1926

29 Face yellow with a pair of brown lateral spots on ventral half; scutum

yellow with a pair of submedial brown vittae extending onto scutellum

laterally; abdomen yellow with broad brown lateral margins; male fore

tibia with 2 preapical setae; male fore basitarsus with an apically

spinulose anteroventral process [eastern Australia; 1 sp., illustrated by

Permkam and Hancock 1995] .................. Clusiosomina Malloch, 1939

— Notas above; if face with dark spots or band then this is placed medially,

not in lower half; male fore basitarsus without an apical process ........ 30

Australian Entomologist, 2015, 42 (3) 117

30 Wing vein R; ending beyond line of R-M crossvein, the pterostigma as

—

long as cell c; scutum yellow with 6 black vittae not extending onto

scutellum [eastern Indonesia (Papua Province) and Papua New Guinea; 1

sp., illustrated by Hardy 1986] ............... Nothoclusiosoma Hardy, 1986

Wing vein R; ending at or before line of R-M crossvein, the pterostigma

shorter than cell c; scutum variable, if yellow with 6 black or brown vittae

then submedial pair extending over scutellum .......................00.005. 31

Face with a black medial spot; male with fore femur ventrally spinose,

fore tibia with a pad-like posteroapical process, fore basitarsus without a

ventral comb and surstyli long and slender [Papua New Guinea; 1 sp.,

illustrated by Hardy 1986] ..................... Hemiclusiosoma Hardy, 1986

Face black, yellow or with a pair of usually confluent lateral spots; male

with fore femur often not ventrally spinose, fore tibia without a pad-like

posteroapical process, fore basitarsus often with a ventral comb and

surstyli short and thick ...................... Trypanocentra ‘complex’ ... 32

32 Head with vertex developed into a sharp, thin, posteriorly directed keel;

face distinctly convex in profile, highest just below middle and receding

to epistome and often with a black, transverse medial band [eastern

Indonesia (West Papua), Papua New Guinea (mainland and New Britain),

Australia (neQId) and Solomon Islands; 3 spp, keyed and illustrated by

Hardy 1986; ‘R. fascifacies’ of Permkam and Hancock 1995 is a

misidentification of R. nigrotibia Hering (Hancock et al. 1998)] ...........

etn RNa Sy ote Ane A robs A EL os: ALEC a haters tut te! Rabaulia Malloch, 1939

Head with vertex at most weakly keeled, not posteriorly directed; face

normally vertical, if convex then all yellow and highest in middle ...... 33

Face gibbose, the convexity highest in the middle; face yellow and

unspotted; scutum with a pair of submedial black vittae; male fore

basitarsus with black ventral comb [eastern Indonesia (Papua Province)

and Papua New Guinea (mainland and New Britain); 1 sp., illustrated by

Hardy TIBG] hereunto deb ees Rad Rabauliomorpha Hardy, 1970

Face vertical in profile or almost so; colour of face and scutum variable;

male fore basitarsus with or without a ventral comb [2 subgenera] .........

ee ee ee eee ee ted eae Trypanocentra Hendel, 1914 ... 34

34 Male fore basitarsus with 1 or 2 ventral combs [eastern Indonesia (Papua

Province) and Papua New Guinea (mainland, New Britain, New Ireland);

5 spp, keyed and illustrated by Hardy 1986] ....................s...2asassa.sa

TE PEDO. he ee I TS E L subgenus Clusiomorpha Hering, 1941

Male fore basitarsus without a ventral comb [eastern Indonesia (Aru),

Papua New Guinea and Australia (neQld); 6 spp, keyed and illustrated by

Hardy 1986; additional illustrations of T. nigrithorax Malloch, 1939 in

Permkam and Hancock 1995] ...... subgenus 7rypanocentra Hendel, 1914

118

Australian Entomologist, 2015, 42 (3)

Key to Aethiothemara species

Pterostigma very elongate, the costal section in cell r; shorter than that in

cell 1213; vein M with last section shorter than the second last; hyaline

indentation in cell r; oval; mid femur with 4 long ventral setae [Equatorial

Guinea (Biok0)] ..........iisiissaasssssssaaaaaaaaan A. transiens Hendel, 1928

Pterostigma less elongate, the costal section in cell rı longer than that in

cell r233; vein M with last section longer than the second last; hyaline

indentation in cell r; quadrate; mid femur with 2 long ventral setae ....... 2

Pterostigma with apex hyaline, included within the hyaline indentation in

cell rı, which ends at vein Rə; hyaline spots in cells br and 14+5 narrow

and linear [Cameroon, Equatorial Guinea (Bioko), Gabon, Congo,

Democratic Republic of Congo and Uganda; var. trispila (Bezzi, 1923)

has an additional hyaline spot in cell dm below R-M crossvein; illustrated

by Enderlein 1911] .......................... A. fallacivena (Enderlein, 1911)

Pterostigma with apex brown or black; hyaline indentation in cell rı

sometimes extends beyond vein R; hyaline spots in cells br and r4+5

often broad and rounded 2.00000... 000. c ccc cee cee ceeceeeeeeeeaeeueeees 3

Hyaline indentation in cell r; ends at vein R23; hyaline spots in cells br

and r4+5 elongate or broadly Oval ..............isisassasaaaaassaaaaaaaaasaaaaa 4

Hyaline indentation in cell r; extends beyond vein R>,3; hyaline spots in

cells br and 14+5 broadly rounded .............iisnasaisssasassasssssassassasasssssaaaai D

Scutum with dorsocentral setae placed just before the supra-alar setae;

hyaline spots in cells br and 145 about 1.5 times as high as broad

[Cameioon i n 2 ain SSA Ø, JE onlara A. trigona Hendel, 1928

Scutum with dorsocentral setae placed just behind the supra-alar setae;

hyaline spots in cells br and 1445 broadly oval [Uganda and Tanzania]

Mee mS TO a ð Mga ee A. graueri Hendel, 1928

Scutum and scutellum without (?) a pair of dark submedial vittae [“West

Africa’; only partially described in a key] .... A. speiseriana (Bezzi, 1924)

Scutum and scutellum with a pair of dark submedial vittae [Uganda;

probably synonymous with A. speiseriana] ........ A. striata Hendel, 1928

Key to Diarrhegma species

= new records based on examined material in the Natural History

Museum, London (*) or photographs submitted for identification (^).

Abdomen with dark transverse bands narrower than yellow bands; wing

of male with a distinct hyaline spot in cell 14+5 (female with 2 spots) and

hyaline spots in cells 1243 and dm distinct [India and Sri Lanka*; Trypeta

incisa Wiedemann, 1824 and Trypeta viana Walker, 1849 are synonyms

Australian Entomologist, 2015, 42 (3) 119

(Hardy 1986, Hancock 1998); illustrated by Bezzi 1913 and Hancock and

Drew:-19994 DD iristeari heia Seat D. modestum (Fabricius, 1805)

— Abdomen with dark transverse bands broader than yellow bands; wing of

male without a hyaline spot in cell r4+s (female with 2 spots) and hyaline

spots in cells 12+3 and dm vestigial or absent [S China (Yunnan, Guangxi,

Guangdong*, Hong Kong*, Macau, Hainan), Vietnam’, Thailand, West

Malaysia*, Singapore”, Borneo, Indonesia (Java, Sumba) and southern

Philippines (Mindanao, Palawan); the citation of Ambon in the Moluccas

as the type locality (Bezzi 1913, Hardy 1986) appears to be an error, since

Doleschall (1856) mentioned only Jogjakarta in central Java in his

original description (Hancock and Drew 1994b) and the specimen is

possibly mislabelled; Diarrhegma eburata Zia, 1963 is a synonym

(Hancock and Drew 1994b); illustrated by Wang 1998 (both sexes) and

Zia 1963 COM ising epee ohne vacenep eprepean as gaecctnan D. paritii (Doleschall, 1856)

Key to Dirioxa species

Wing largely fuscous, without a hyaline costal indentation in cell r; above

R-M crossvein or a hyaline band across apex of cell dm; abdomen fulvous

to red-brown with or without dark transverse bands on terga IHI, III and

IV [Vanuatu; illustrated by Hancock and Drew 2003] ........................

Sa barð ab ahs taeda D. fuscipennis Hancock & Drew, 2003

Wing with a broad, quadrate hyaline costal indentation in cell r; above R-

M crossvein and a hyaline band across apex of cell dm; abdomen fulvous

on terga IHI, mostly or entirely black on terga I-V ........................ 2

Wing with posterior hyaline indentation in cell cu, extending broadly into

cell dm medially; hyaline apical band in cell dm distinctly broadened

anteriorly and often united posteriorly with medial indentation [Australia

and New Caledonia (introduced); Trypeta musae Froggatt, 1899 and

Rioxa confusa Hardy, 1951 are synonyms (Permkam and Hancock 1995);

illustrated by Hardy 1951 and Permkam and Hancock 1995] ...............

EE TAN E ETR E BERIETE atlece de D. pornia (Walker, 1849)

Wing with posterior hyaline indentation in cell cu, not extending into cell

dm; hyaline apical band in cell dm not broader anteriorly than posteriorly

[eastern Indonesia (Papua Province); illustrated by Hardy 1986; placed in

Dirioxa by Hancock and Drew 2003] ............. D. incerta (Hardy, 1986)

Key to Termitorioxa species

Wing cell 145 with a curved longitudinal hyaline band from spot above

DM-Cu crossvein to apex [Australia (Qld); illustrated by Permkam and

Hancock 1995] aaan T. bicalcarata (Hering, 1944)

— Wing cell 1445 without a curved longitudinal hyaline band at apex ........ 2

120 Australian Entomologist, 2015, 42 (3)

2 Wing cell dm almost entirely hyaline or with an elongate medial hyaline

band along vein CAT ....aiiiiiiiaaaiiiaiiaaasssaasaasaasakaassaaskakassakkaaaaaaa 3

— Wing cell dm mostly brown and without an elongate hyaline band along

WE O R 2 JÁ Jo's að Weide) Cho al a ede stat Darren maaa a aT Aa emer

3 Wing cell dm almost entirely hyaline; large hyaline area in cell dm united

with large hyaline spots in cells br and r4+5; a second large hyaline spot in

cell r445 united with indentation in cell m; postnotum fulvous [Papua New

Guinea; illustrated by Hardy 1986] ................... T. flava (Hardy, 1986)

— Not as above; cell dm with an elongate medial hyaline band along vein

Cu,; cell r4, with second hyaline spot, if present, small and isolated;

postnotum largely blackish brown .............0. 0.000000 cece ee ec ec ee eee ees 4

4 Cell 145 with hyaline spot above DM-Cu crossvein united with

indentation in cell m; scutum fulvous [Australia (nQld); illustrated by

Permkam and Hancock 1995] .................... T. testacea (Hendel, 1928)

— Cell 145 with hyaline spot above DM-Cu crossvein not united with

indentation in cell m; scutum with a brown to black posterior border

[eastern Indonesia (Misool and Papua Province) and Papua New Guinea;

Ptilona lateralis Kertész, 1901 is a synonym (Hardy 1986); illustrated by

Hardy: 1986] ..: 2123 eresas rupnik T. meritoria (Walker, 1864)

5 Cell cu, with a hyaline indentation at apex below DM-Cu crossvein;

costal cell c hyaline [northern Australia; illustrated by Permkam and

Hancock 1995] ..................22.2 T. exleyae Permkam & Hancock, 1995

— Cell cu, without a hyaline indentation at apex below DM-Cu crossvein;

costal cell c hyaline or at least partly yellowish .............................. 6

6 Head with 1 pair of orbital setae; wing base pale yellowish, including

cells be MÀ ese RN PLE A I YOR YE RSS PST A 7

— Head with 2 pairs of orbital setae; wing base hyaline or with pale

yellowish markings in cell c or cells ctbr o.n 8

7 Scutum normally with a pair of brown patches adjacent to scutellum;

scutellum normally with a pair of dark basal patches; postnotum blackish

brown laterally; wing with hyaline spot in cell 145 not united with

indentation in cell m and pattern dark brown with distinct yellowish areas

in most of cell dm and around R-M crossvein; male with a tuft of fine

hairs at apex of arista [northern Australia; illustrated by Permkam and

Hancock 1995]... 0.0... T. termitoxena (Bezzi, 1919)

— Scutum and scutellum entirely fulvous; postnotum fulvous; wing with

hyaline spot in cell r4.; united with indentation in cell m; male without a

tuft of fine hairs at apex of arista [eastern Indonesia (Timor); illustrated

by Hardy 1986] ..........iisssaiiaaanasaaasaaaaaaaað T. timorensis Hardy, 1986

Australian Entomologist, 2015, 42 (3) 121

Cell r; with hyaline indentation small, not crossing into cell 1243; cell c

LTR ech NE ag 2 TB, each, Se ESOL SR Re ire Ne Let ONL LRT OI. fe AA 9

Cell rı with hyaline indentation large, crossing into cell 1243; cell c at least

partly yellowish nda hertan edna es da eet aa aa scarey a sie home 10

Cells br and 145 each with a distinct medial hyaline spot; hyaline spot at

apex of cell dm anteriorly placed [Australia (NT); illustrated by Permkam

and Hancock 1995] ............... T. inconnexa Permkam & Hancock, 1995

Cells br and 14+5 without distinct hyaline spots; hyaline spot at apex of cell

dm posteriorly placed [eastern Indonesia (Papua Province); illustrated by

Hardy 1986] aoa T. acanthoneurides (Hering, 1953)

10 Postnotum red-brown laterally; cells c+br yellowish, cell c sometimes

with only a complete yellow costal band [Australia (NT); illustrated by

Hancock 1996] ..................................... T. cobourgensis Hancock, 1996

Postnotum entirely fulvous; cell c with a yellow costal band over basal

two-thirds [northern Australia; illustrated by Permkam and Hancock

TOIS cig rie ve sabes Sara rnein T. laurae Permkam & Hancock, 1995

Key to Themaroides species

Pleura with a longitudinal black or brown vitta from proepisternum to

anatergite; wings largely brown, including costal cells ..................... 2

Pleura pale with at most a dark line along dorsal margin of anepisternum;

wings often with large hyaline indentations and base, including costal

cells, subhyaline or yellowish ............siiaiiasnanaaasssaasasaassaaaaaaaaaan 4

Scutum and scutellum with a complete medial black vitta; face with a

medial black vitta on lower half; additional small secondary scutellar

setae absent [eastern Indonesia (West Papua and Papua) and Papua New

Guinea; Rioxa debeauforti de Meijere, 1906 is a synonym (Hardy 1986);

illustrated by Hardy 1986] .................... T. abbreviatus (Walker, 1865)

Scutum and scutellum without a complete medial black vitta; face wholly

yellow; 2 pairs of additional small secondary scutellar setae present ...... 3

Scutum and scutellum yellow, without distinct dark vittae; scutellum

entirely densely setulose [Papua New Guinea; illustrated by Hardy 1986]

ad es a peel cect r abc ðe has Ser Meath T. robertsi Hardy, 1986

Scutum with 5 narrow, dark postsutural vittae; scutellum with dark lateral

vittae and only densely setulose laterally, bare medially [Papua New

Guinea; illustrated by Hardy 1986] .................. T. vittatus Hardy, 1986

Wings pale brown to subhyaline (possibly teneral) but without distinct

large hyaline marginal indentations; vein R4s straight, not curving

posteriorly in apical portion, the wing narrow and elongate [Papua New

Guinea; illustrated by Hardy 1986]............ T. xanthosoma Hardy, 1986

122 Australian Entomologist, 2015, 42 (3)

— Wings dark brown, subhyaline to yellow basally and with large hyaline

marginal indentations in at least cells m and cu,;+dm; vein R4+s curving

posteriorly in apical portion, the wing normal in shape ..................... 5

5 Wing with a large quadrate hyaline indentation in cells rı and 1243 from

costa to vein R4s; posterior hyaline indentation confined to cells cu;+dm;

scutellum without an additional pair of small secondary setae [eastern

Indonesia (Ambon and West Papua) and Papua New Guinea; Helomyza

optatura Walker, 1865 is a synonym (Hardy 1986; illustrated by Hardy

162516 Ta hbase, MER ENT CA Bet asl tay T. quadrifer (Walker, 1861)

— Wing without an anterior hyaline indentation; posterior hyaline

indentation in cells cu;+dm extending into cell br; scutellum with an

additional pair of small secondary subapical setae [Papua New Guinea

(New Britain); illustrated by Hancock and Drew 2003] ......................

EA T. Me Bday TEs A A ae eae ema T. bicolor Hancock & Drew, 2003

Key to Clusiosoma pullatum group males

The pullatum group keys to couplet 14 in Hardy (1986). Males of the

Australian C. macalpinei Permkam & Hancock, 1995 (females unknown)

may be separated using the following key.

1 Face and genae yellow; occiput yellow posteroventrally; fore femur

broadly swollen and with a subbasal clump of ventral spinules in basal

third [Papua New Guinea; illustrated by Hardy 1986] .......................

ee ee Ee oe Oe Ye C. subpullatum Hardy, 1986

— Face and genae black; occiput black posteroventrally; fore femur without

a clump of ventral spinules in basal third .................i2ii iii aaa asa eee 2

2 Fore femur narrowly swollen and with long spines restricted to middle

third, absent from apical third [Papua New Guinea; illustrated by Hardy

DESC] A Aeae tad bates adn C. pullatum Hering, 1941

— Fore femur broadly swollen and with long spines continuous over middle

and apical thirds [Australia (nQld); illustrated by Permkam and Hancock

1995] ee ee TT D C. macalpinei Permkam & Hancock, 1995

Discussion

The Afrotropical Aethiothemara group of genera have mid tibiae with 1 long

and several short apical spines. One of the secondary spines is about half as

long as the main spine in Themarictera and all are about 1/5 as long in the

other genera (A. Freidberg pers. comm.). This is similar to the condition seen

in the Neothemara and Clusiosoma subgroups but cell m has only a single

marginal hyaline indentation and the pattern is not almost entirely brown,

suggesting a closer relationship with Diarrhegma and the Themaroides

subgroup and a homoplasious reduction in the size of the secondary spine.

Australian Entomologist, 2015, 42 (3) 123

It appears likely that the Aethiothemara, Diarrhegma and Themaroides

groups represent radiations in Africa, Asia and Australasia respectively, the

presence of tree-trunk breeders in all three groups suggesting that decaying

tissue beneath the bark is the ancestral larval food source. Fruit-feeding in

Themarictera, Dirioxa and the Clusiosoma subgroup, which infest different

fruit types, are therefore likely to represent homoplasious apomorphies.

All six African genera lack anepimeral (= pteropleural) setae and the R-M

crossvein is placed within the apical third of cell dm, supporting monophyly

of the group. Afrocneros, Labeschatia, Ocnerioxa and Ptiloniola have only 4

scutellar setae (apomorphy within the group) and appear to form a related

series; Aethiothemara and Themarictera have the usual 6 scutellar setae. In

Themarictera the pterostigma is shorter and the secondary midtibial spine

longer than in the other genera and the male head is expanded, similar in

shape to some Themara Walker species in the Acanthonevra group.

In the Southeast Asian genus Diarrhegma, the short, broad epandrium and

surstyli in males and long-setose, apically broadly rounded aculeus are

typical of the groups treated here. Although host records are few, it is likely

to be a non-generalist breeder beneath decaying tree bark.

In the Australasian 7Themaroides group, the Themaroides subgroup most

resembles the Diarrhegma group in having two long, subequal midtibial

spines. The male epandrium and surstyli are short and broad, narrowed

slightly in Termitorioxa and some Themarohystrix. The aculeus is apically

blunt or broadly rounded except in Themarohystrix, where the apex is

subtriangular and tapered to a blunt point (apomorphy). In Buloloa,

Themarohystrix and Themaroidopsis veins M and/or Cu+Cu, are setulose, a

character also seen in most Cl/usiosoma subgroup species. Buloloa and

Enoplopteron have a row of long setae on the costa (apomorphy). In the

Australian genera Acanthonevroides, Aridonevra and Taeniorioxa the

scutellum is distinctly swollen (apomorphy); these three genera, and possibly

also Walkeraitia, lack the additional supra-alar seta that is present

(apomorphy) in all the other genera, although in Termitorioxa the additional

seta is small and sometimes more than one are present. Walkeraitia and

Aridonevra share an unusually-shaped cell dm, narrowed in basal half and

broadly expanded in distal half, together with elements of the wing pattern,

that suggest a reasonably close relationship.

The Neothemara subgroup is characterised by the numerous hyaline spots or

indentations on the wing, including three marginal spots in cell M. Where

known, all have a short and broad epandrium and surstyli in males except in

Hexaresta multistriga (Walker), where the surstyli are distinctly narrowed

(apomorphy). The aculeus is apically blunt or broadly rounded except in

Neothemara digressa Hardy, where it is short and apically triangular

(apomorphy); this species is possibly generically distinct.

124 Australian Entomologist, 2015, 42 (3)

Within the C/usiosoma subgroup, the male epandrium and surstyli are short

and broad in Nothoclusiosoma, Paedohexacinia, Rabaulia, Rabauliomorpha

and Trypanocentra, or long and slender (apomorphy) in Cheesmanomyia,

Clusiosoma, Clusiosomina and Hemiclusiosoma. An apical pad-like process

on the fore tibia (apomorphy) links Nothoclusiosoma with Clusiosoma and

Hemiclusiosoma; in Clusiosomina the pad is absent but the fore basitarsus

has a prominent anteroapical process (apomorphy) and in Cheesmanomyia

both are absent. The face is concave in Clusiosoma (apomorphy), convex in

Rabaulia and Rabauliomorpha (homoplasious apomorphies) and vertical or

nearly so in the other genera. The female aculeus is tapered and apically

acute (apomorphy) in Paedohexacinia and broadly blunt apically in all the

other genera. Rabauliomorpha and Trypanocentra (Clusiomorpha) both have

a ventral comb (apomorphy) on the fore basitarsus; its absence in subgenus

Trypanocentra suggests that Clusiomorpha should be raised to genus level

but further study is required. Additionally, T. (Trypanocentra) tricuneata

Hardy resembles Rabaulia in facial and scutal markings and has a distinct

keel on the vertex of the head, also suggesting that 7rypanocentra is

paraphyletic and in need of revision.

The Dirioxa group contains both rotting wood (e.g. Lumirioxa) and rotting

fruit infesters (Dirioxa) and, with its single midtibial spine, is possibly related

to the Acanthonevra group (Korneyev 1999). The epandrium and surstyli of

males are short and broad. The aculeus is normally long-setose and apically

blunt or broadly rounded (including in Parachlaena: see Hancock and Drew

1994a) except in Griphomyia, where it is short-setose and apically produced

to a blunt point (apomorphy) and in Micronevrina, where it is microsetose

and apically bifurcate (apomorphy). Griphomyia and Mimoeuphranta have

small secondary scutellar setae and similar wing pattern elements, including

the R-M crossvein placed below or before the midpoint of the pterostigma; in

the other genera it is placed near or beyond its apex. In Anchiacanthonevra

and Parachlaena cell r; has two distinct hyaline indentations from costa and

the apex of cell 1445 is extensively hyaline. The occasional use of fallen

Araucaria cunninghamii cones by Dirioxa pornia larvae links that genus with

Lumirioxa, which breeds beneath the bark.

Acknowledgements

I thank Susan Phillips for Figure 1, Andy Wang and Geoff Monteith

(Queensland Museum) for Figures 2 and 3, Amnon Freidberg (Tel Aviv

University, Israel) for information on the African genera and Daniel

Whitmore (Natural History Museum, London) for access to specimens in his

care.

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Tephritidae) from Australia and the south Pacific. Australian Entomologist 30(2): 65-78.

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distribution and host plants of fruit flies (Diptera: Tephritidae) in Australia. Queensland

Department of Primary Industries Information Series Q199067; iii + 75 pp.

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HANCOCK, D.L., OSBORNE, R. and McGUIRE, D.J. 1998. New hostplant and locality

records for Ceratitidinae and Trypetinae in northern Queensland (Diptera: Tephritidae). General

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HARDY, D.E. 1951. The Krauss collection of Australian fruit flies (Tephritidae—Diptera).

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HARDY, D.E. 1986. Fruit flies of the subtribe Acanthonevrina of Indonesia, New Guinea, and

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Australian Entomologist, 2015, 42 (3): 127-128 127

FIRST RECORD OF PETRELAEA TOMBUGENSIS (ROBER)

(LEPIDOPTERA: LYCAENIDAE) FROM WESTERN AUSTRALIA

MICHAEL F. BRABY

Research School of Biology, Australian National University, Canberra, ACT 0200

Abstract

Petrelaea tombugensis (Rober, [1886]) is recorded from Western Australia for the first time in

the northern Kimberley. Adults were recorded in riparian evergreen monsoon vine forest during

May 2015. This new location, together with previous records from the “Top End’, NT and Cape

York Peninsula, Qld, suggest the species is distributed widely but sporadically throughout the

Australian monsoon tropics.

The lycaenid butterfly Petrelaea tombugensis (Rober, [1886]) has a sporadic

distribution within Australia. It was first recorded from the Australian

mainland only as recently as 1977, in northeastern Queensland near Iron

Range (McEvey 1977). It has since been recorded at a few other locations,

including Cobourg Peninsula, NT (Common and Waterhouse 1981), Torres

Strait (Muller et al. 1998), Lockerbie on Cape York Peninsula (Braby 2000),

and the Wet Tropics between Cairns and El Arish (Muller et al. 1998),

Queensland. Adults have mainly been recorded in monsoon vine thicket or

lowland rainforest in riparian areas where they are believed to breed on

Terminalia spp. with which they are strongly associated (Miller et al. 1998,

Braby 2000). The butterfly appears to be highly seasonal in its timing of

appearance.

In northwestern Australia, P. tombugensis is considered to be rare, known

previously only from two specimens (both males) collected from Black Point

on Cobourg Peninsula in the ‘Top End’ of the Northern Territory, on 26-27

January 1977 by E.D. Edwards (specimens examined in the Australian

National Insect Collection, Canberra). On 19 May 2015, I collected six

specimens (2 males, 4 females) of P. tombugensis from the Carson River

crossing on the Kalumburu Rd, approximately 18 km SSE of Kalumburu,

Western Australia (14.45277°S, 126.66373°E). They were associated with

riparian evergreen monsoon vine forest and were either flying around

vegetation 2-4 m above the ground or puddling from moist sand along the

edge of a stream. The specimens were all in relatively good condition,

according to the extent of wing wear, and had probably emerged shortly

before their capture.

The new location from the northern Kimberly in Western Australia extends

the range of this species 700 km southwest from its former western limit in

the Northern Territory. This new location, together with previous records

from the ‘Top End’ and Cape York Peninsula (and elsewhere in northeastern

Qld), suggest that P. tombugensis is distributed widely throughout the

monsoon tropics of northern Australia. Presumably, its apparent rarity is a

128 Australian Entomologist, 2015, 42 (3)

consequence of its temporal and spatial distribution, being highly seasonal

and restricted to patches of riparian monsoon forest. Further work on the life

history and larval food plant specialisation is needed to determine the

species’ ecological requirements.

Acknowledgments

I thank G.J. Paras for his pleasant company in the field and E.D. Edwards for

access to material in the ANIC.

References

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

CSIRO Publishing, Collingwood, Melbourne; xx + 976 pp.

COMMON, LF.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Revised edition.

Angus and Robertson, Sydney; xiv + 682 pp.

McEVEY, S.F. 1977. Notes from a collecting trip to Cairns and Iron Range including a new

record for the Australian mainland. Victorian Entomologist 7: 59-62.

MULLER, C.J., OLIVE, J. and LAMBKIN, T.A. 1998. New records for Petrelaea tombugensis

(Röber) (Lepidoptera: Lycaenidae) in Queensland. Australian Entomologist 25: 61-63.

Australian Entomologist, 2015, 42 (3): 129-132 129

TAGIADES NESTUS KORELA MABILLE, 1891

(LEPIDOPTERA: HESPERIIDAE: PYRGINAE) FROM

MER ISLAND, TORRES STRAIT, QUEENSLAND

S.S. BROWN’, C.E. MEYER’, R.P. WEIR? and C.G. MILLER“

119 Kimberley Drive, Bowral, NSW 2576 (Email: stnac@bigpond.com)

729 Silky Oak Avenue, Moggill, Old 4070 (Email: cmameyer@bigpond.com)

390 Horne Road, Bees Creek, NT 0822 (Email: Richard. Weir@nt.gov.au)

137 Alison Ave, Lennox Head, NSW 2478 (Email: cgrantmiller@bigpond.com)

Abstract

Tagiades nestus korela Mabille, 1891 is recorded from Mer (Murray) Island in Torres Strait for

the first time. One male specimen was collected on 12 April 2015 flying on a track in monsoon

forest near the centre of the island. T. n. korela is now known from three Torres Strait islands,

viz. Dauan, Erub (Darnley) and Mer.

Introduction

Tagiades Hübner contains twelve species distributed from Africa to northern

Australia (Parsons 1998, Braby 2000). In the Australian Region it is

represented by three taxa, viz. T. trebillius canonicus Fruhstorfer, 1910,

which is restricted to Papua New Guinea (Evans 1949), T. nestus korela

Mabille, 1891 and T. japetus janetta Butler, 1870, which are additionally

found in Torres Strait and eastern tropical Queensland (Parsons 1998).

Tagiades species prefer shaded areas and tracks beneath and adjacent to the

forest canopy or mangroves, usually around areas where their larval host

plant occurs (pers. obs.).

The first Australian specimen of 7. nestus (C. Felder) was collected on Erub

(Darnley) Island, Torres Strait in May 1910 by H. Elgner (Fig. 7) and is

housed in the Australia Museum, Sydney. It was illustrated as T. neira Plotz,

1885 by Waterhouse and Lyell (1914). Although not illustrated in his later

work, Waterhouse (1932) did make reference to this specimen as T. neira,

belonging in the T. nestus species group. It was assigned to T. nestus korela

by Common and Waterhouse (1972).

After this initial collection, there were no further records of this species from

Australia until it was rediscovered, in April 2004, on Dauan Island, Torres

Strait (Valentine and Johnson 2005). On that occasion the immature stages

were also recorded, on climbing yam vines (Dioscorea sp.: Dioscoraceae). In

April 2015, we conducted a butterfly survey on Mer, Dauan and Saibai

Islands. While on Mer Island, a male T. n. korela (Figs 1-2) was observed

and collected in monsoon forest in the central part of the island.

Mer Island lies in the far east of Torres Strait, roughly 120 km SE of Papua

New Guinea at its closest point. Roughly circular in shape and approximately

2.8 km by 1.7 km in size, it is the result of Pleistocene era volcanic activity,

with the major land formation behind the village consisting of a volcanic vent

composed of poorly consolidated scoria and ash (Torres Strait Regional

130 Australian Entomologist, 2015, 42 (3)

Authority 2013). Thus, this part of the island is extremely porous and, with

its associated severe drainage, limits the nature and abundance of vegetation

growing on the western half of the island. The eastern half of the island is

predominantly monsoon forest mixed with many introduced species and it is

in this area that most butterflies were collected.

Discussion

Tagiades nestus korela is now known to occur on three islands in Torres

Strait, viz. Dauan in the north and Mer and Erub in the east. Due to the

widespread planting of yam vines on many of the Torres Strait islands and

hundreds of years of inter-island trade between them and Papua New Guinea

(Doug Passi and Barbara Waterhouse pers. comms.), it is feasible that

individuals were transported to the islands as immatures on yam plants. It is

also possible that the insect has existed on the islands since they were

connected by a land bridge, when Torres Strait was a part of the Sahul shelf

(Parsons 1998). Tagiades japetus janetta (Figs 3-6) is not known from Mer

Island but the two species are sympatric on Dauan Island. In January 2011,

one of us (CEM) collected a pupa on Dauan Island in a typical Tagiades

shelter on Dioscorea sp., which emerged as T. n. korela. At the same time,

within 300 metres of that spot, one of us (SSB) collected a T. j. janetta male

(Figs 3-4). Based on our records, T. j. janetta is more widespread and

observed more frequently on Dauan Island than T. n. korela.

The original Elgner specimen of T. n. korela is labelled as having passed

through the C.W. Wyatt Theft Collection. This refers to the activities of Colin

Wyatt, an English amateur entomologist who stole specimens from a number

of Australian Museums (and the Natural History Museum in London) in the

1930s and 1940s. Wyatt appeared at Westham Magistrates Court in London

in 1947 and was fined £100 when he pleaded guilty to the unlawful

possession of a large number of Australian specimens. These were returned

to Australia, where it was found that labels on a number of specimens had

been removed or altered, presumably to conceal their provenance. All such

Specimens now bear yellow labels alerting researchers to the fact that they

passed through Wyatt’s hands and that labels may be inaccurate (John

Tennent pers. comm. ).

E.D. Edwards (pers. comm.) also provided information regarding the veracity

of the label data on Elgner’s historical specimen of T. n. korela, stating: ‘The

specimens stolen by Wyatt were returned to the South Australian Museum

(which was also robbed of specimens by Wyatt) and the specimens belonging

to the different Australian institutions sorted out by Norman Tindale.

Waterhouse was too ill to travel to Adelaide to do it. While Waterhouse may

have complained about some details of Tindale’s sorting, he would have

made sure such an historic specimen was returned to the Australian Museum

with the right labels. The Elgner label is genuine, the date is right, the sex is

right’.

Australian Entomologist, 2015, 42 (3) 131

Australian Museum

TORRES STRAIT IS. K 455977

[Darney L£. )

Figs 1-7. Tagiades spp: (1-2) T. nestus korela from Mer Island: (1) upperside; (2)

underside (forewing length 21 mm). (3-4) T. japetus janetta (Papua New Guinea

form) from Dauan Island: (3) upperside; (4) underside (forewing length 22 mm). (5-6)

T japetus janetta from Thursday Island, Torres Strait: (5) upperside; (6) underside

(forewing length 21 mm). (7) T. nestus korela from Darnley Is, 15.v.[19]10, H. Elgner

(Image source: Australia Museum: Russell Cox).

132 Australian Entomologist, 2015, 42 (3)

The tornus of the left hind wing has been repaired at some stage. This repair

seems to have been done after the specimen was illustrated in Waterhouse

and Lyell (1914) as the illustrated specimen is lacking the third black spot on

the tornus of the left hind wing.

In spite of the Elgner specimen’s journey and repairs, it seem safe to assume

that the specimen in the Australian Museum is genuine as it was illustrated

only four years after it was collected and the label (Fig. 5) appears consistent

with other Elgner labels.

Thus, given the fresh condition of our specimen, the isolation of Mer Island

and the abundance of the larval host plant on the island, the species is most

likely established on the island.

Acknowledgements

The authors thank Mr Doug Passi (Chairman of the Prescribed Bodies

Corporate on Mer Island), Mr Joey Tapau (Biosecurity Officer, Mer Island)

and the people of Mer Island for permission to visit and conduct the butterfly

survey there. We thank Mr E.D. Edwards for information regarding the

history and validity of the Elgner record of 7. n. korela from Erub (Darnley)

Island, John Tennent (Natural History Museum, London) for information

regarding the C.W. Wyatt Theft Collection and Dr David Britton, Russell

Cox and Jacquie Recsei of the Australian Museum, Sydney for providing the

digital image and contextual information relating to the 1910 H. Elgner

record of 7. n. korela. We also thank Dr Trevor Lambkin for his comments

on the manuscript.

References

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

CSIRO Publishing, Collingwood; xxvii + 976 pp.

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

Sydney; 498 pp, 41 pls.

EVANS, W.H. A catalogue of the Hesperiidae from Europe, Asia and Australia in the British

Museum (Natural History). British Museum (Natural History), London; 502 pp.

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

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

TORRES STRAIT REGIONAL AUTHORITY. 2013. Profile for management of the habtats and

related ecological and cultural resource values of Mer Island. Available at: http:/Avww.tara.gov.

au/__data/assets/pdf_file/0008/4499/Mer_Biodiversity_Profile_January_2013.pdf

VALENTINE, P.S. and JOHNSON, S.J. 2005. The rediscovery of Tagiades nestus (C. Felder)

(Lepidoptera: Hesperiidae: Pyrginae) in Australia. Australian Entomologist 32: 155-156.

WATERHOUSE, G. 1932. What butterfly is that? Angus & Robertson, Sydney; viii + 291 pp,

34 pls.

WATERHOUSE, G.A. and LYELL, G 1914 The butterflies of Australia. A monograph of the

Australian Rhopalocera. Angus & Robertson, Sydney; vi + 239 pp.

Australian Entomologist, 2015, 42 (3): 133-147 133

A NEW SUBSPECIES OF HASORA HURAMA (BUTLER, 1870)

(LEPIDOPTERA: HESPERITDAE: COELIADINAE) FROM THE

NORTHERN TERRITORY, AUSTRALIA

C.E. MEYER’, R.P. WEIR’ and S.S. BROWN?

129 Silky Oak Avenue, Moggill, Old 4070 (Email: cnameyer@bigpond.com)

790 Horne Road, Bees Creek, NT 0822 (Email: richard.weir@nt.gov.au)

319 Kimberley Drive, Bowral, NSW 2576 (Email: stnac@bigpond.com)

Abstract

Hasora hurama territorialis subsp. n. is described and figured from the ‘Top End’, Northern

Territory, Australia. Distinct differences in adult phenotype separate it from populations of the

nominate subspecies H. hurama hurama (Butler, 1870) from Queensland, Australia and from

other subspecies in the Moluccas, mainland New Guinea and adjacent islands eastwards through

the Bismarck Archipelago to the Solomon Islands. The two subspecies in Australia are

geographically isolated and are consistent in their character differences. Hasora h. hurama is

newly recorded from Dauan Island, northern Torres Strait.

Introduction

Hasora hurama (Butler, 1870) ranges from the Moluccas, Aru, Biak,

Maluku, through mainland New Guinea and adjacent islands and northern

Australia to the Bismarck Archipelago and Solomon Islands (Evans 1949,

Parsons 1998, Braby 2000, Tennent 2002). Evans (1949) recognised four

subspecies but subsequently described a fifth, H. h. diana Evans, 1958, from

Rennell Island (Evans 1958). Two subspecies occur in Papua New Guinea,

with Æ. h. hurama (Butler, 1870) widespread throughout the mainland and its

adjacent islands and H. h. kieta Strand, 1921 occurring in the Admiralties and

most of the Solomon Islands (Evans 1949, Parsons 1998, Tennent 2002). In

eastern Indonesia, H. h. mola Evans, 1949 occurs on the Bacan Islands in

northern Maluku and H. h. arua Evans, 1934 occurs on the Aru Islands in

eastern Maluku (Evans 1949). Hasora h. diana is restricted to Rennell and

Bellona Islands in the Solomon Islands (Evans 1958, Tennent 2002). Only H.

h. hurama has been recognised previously from the Australian subregion

(Common and Waterhouse 1981, Braby 2000).

The type locality of H. h. hurama is Cape York, Queensland (Edwards et al.

2001). In Australia it has been recorded from the Torres Strait islands of

Dauan (CEM in April 2015, T.A. Lambkin pers. comm. and here noted as a

new northern record for Australia), Mer (Murray), Dauer and Hammond

(T.A. Lambkin pers. comm.), Moa and Thursday, and on the mainland at

Cape York, from Cooktown to Townsville and from the Mackay district

(Common and Waterhouse 1981, Braby 2000), including the Conway Ranges

near Proserpine and the Broken River near Eungella, all in eastern

Queensland (Braby 2000). In the Northern Territory it has been recorded

from Buffalo Creek, near Lee Point, Darwin, south through the Elizabeth

River Bridge on the Stuart Highway near Noonamah, east-south-east to Fogg

Dam, the Beatrice Hill region on the Adelaide River flood plain, and further

134 Australian Entomologist, 2015, 42 (3)

east on the Blyth River (Peters 1969, Braby 2000, Meyer et al. 2006) and

new Nangnak outstation (Bisa 2013). Braby (2000) noted that H. h. hurama

shows little variation throughout its range.

For many years we have suspected that there are significant differences in

adult phenotype between H. hurama specimens from the Northern Territory

and those from Queensland, warranting further examination. We consulted

relevant literature and compared specimens held in our private collections

with the type specimens of H. h. hurama (Figs 1-3), H. h. mola (Figs 4-6), H.

h. arua (Figs 7-9) and H. h. diana (Figs 10-12) held in the Natural History

Museum, London (BMNH). We concluded that the Northern Territory

population warrants recognition as a distinct subspecies.

All specimens held in the author’s private collections of the nominate

subspecies from Queensland examined are detailed at Appendix 1. Appendix

2 details the voucher specimens of H. h. territorialis subsp. n. examined and

used in the description of this subspecies.

Abbreviations for specimen repositories. AMC — Australian Museum

Collection, Sydney; ANIC — Australian National Insect Collection, Canberra;

BMNH — Natural History Museum, London; CEMC — C.E. Meyer

Collection, Brisbane; DALC — D.A. Lane Collection, Atherton; MFBC —

M.F. Braby Collection, Canberra, NTMC — Museum and Art Gallery of the

Northern Territory Collection, Darwin; RPWC — R.P. Weir Collection, Bees

Creek; SSBC — S.S. Brown Collection, Bowral.

Abbreviations for collectors. CEM — C.E. Meyer; DNW — D.N. Wilson;

MFB — M.-F. Braby; RPW — R.P. Weir; SSB — S.S. Brown.

Hasora hurama territorialis subsp. n.

(Figs 17-20)

Types: Holotype 6, NORTHERN TERRITORY: Adelaide River Bridge, Bred/larva,

em[erged] 4.11.1994, C.E. Meyer (in ANIC). Paratypes: 3 99, Fogg Dam, 22.iv.1991,

D.N. Wilson; 1 ĝ, Buffalo Creek, 3.iii.1992, C.E. Meyer; 1 4, Adelaide River, em.

22.ix.1992, C.E. Meyer; 1 Æ, 1 Q, same data except em. 24.ix.1992; 1 Æ, same data

except em. 27.ix.1992; 2 SÆ, same data except em. 4.ii.1994 and 5.ii.1994; 19, same

data except em. 10.11.1994; 1 9, Adelaide River Bridge on Arnhem Hwy, em.

16.iv.1995, C.E. Meyer; 3 Æ, same data except em. 18.11.1995, 23.iv.1995 and

6.ix.1998; 1 Æ, Adelaide River Hunting Reserve, Arnhem Hwy, via Darwin, em.

10.iv.2009, C.E. Meyer & R.P. Weir; 1 ĝ, Adelaide River Bridge environs, Arnhem

Hwy, em. 3.v.2013, C.E. Meyer & R.P. Weir; 1 ĝ, same data except, em. 14.v.2013;

3 22, same data except em. 14.v.2013 and 15.v.2013 (all in CEMC); 1 Q, Adelaide

River, 8.v.1989, D.N. Wilson; 2 99, Fogg Dam, 22.iv.1991, D.N. Wilson; 2 22,

Berrimah Research Farm, 8.iv.1994, R.P. Weir; 1 Q, same data except 15.1v.1994;

1 Q, 10 km East, Adelaide River, 7.viii.1996, R.P. Weir, 1 Æ, Adelaide River, em.

6.11.2008, R.P. Weir & S.S. Brown; 1 ĝ, 1 Q, Adelaide River, em. 5.iv.2009, R.P.

Weir & C.E. Meyer; 5 bo, 3 22, Beatrice Hill, em. 1.iii.2000, R.P. Weir, 2 29,

Adelaide River, em. 5.iii.2013, R.P. Weir & M.J. Griffiths (all in RPWC); 2 G,

Australian Entomologist, 2015, 42 (3) 135

4 29, Fogg Dam, 22.iv.1991, S.S. Brown (all in SSBC); 1 Æ, Adelaide River, em.

2.v.1995, D.A. Lane & C.E. Meyer (in DALC); 1 9, Adelaide River, 10km NE

Arnhem Hwy, 9.v.1991, D.N. Wilson (NTM 1004069); 1 Æ, Beatrice Hill (Window

on the Wetlands Visitor Centre), Adelaide River floodplain, M.F. Braby (NTM

1004726) (both in NTMC); 2 Æ, 1 Q, 12.64956°S, 131.31851°E, Beatrice Hill,

17.1.2009, M.F. Braby; 1 Æ, same data except 23.1.2010; 1 Q, 12°39736”S,

131°20°11”E, Adelaide River crssng, Arnhem Hwy, em. 15.x1.2010, M.F. Braby;

2 dh, 2 99, same data except em. 17.xi.2010; 1 3, 1 Q, same data except em.

18.xi.2010; 1 Q, same data except em. 19.xi.2010; 1 Æ, same data except em.

21.xi.2010; 1 Q, same data except em. 27.xi.2010; 1 Æ, same data except em.

7.xii.2012; 1 4, 12°39°36”S, 131°20’11”E, Adelaide River crossing, Arnhem Hwy,

em. 12.vii.2013, M.F. Braby & L. A. Lilleyman; 1 9, same data except em.

16.vii.2013; 1 Æ, 12.40329°S, 132.96107°E, East Alligator, near Ubirr, Kakadu NP,

em. 13.x.2013, M.F. Braby; 1 Æ, same data except em. 16.x.2013; 1 2, same data

except em. 20.x.2013; 2 9 Q, same data except em. 28.x.2013; 1 9, same data except

em. 30.x.2013 (all in MFBC); 1 £, Blyth River, 21.ix.1968, F. Omer-Cooper (Reg.

No: K.446984); 1 2, Adelaide River, 13.1.1962, T.E. Moulds (Reg. No: K.446985)

(both in AMC); 1 Q, Fogg Dam, 22.1v.1991, D.N. Wilson (in ANIC).

Description. Male (Figs 17-18). Upperside: dull dark chocolate-brown, with

lighter brown basal hairs; forewing with an irregular dull brown-black sex-

brand between vein M; and dorsum: head and thorax with light brown hairs;

antennae black with strongly curved apiculus. Underside: ground colour

lighter chocolate-brown than above, with forewing margins lighter brown;

hindwing darker brown than forewing and generally with a purplish sheen

especially in newly emerged specimens; hindwing with an uninterrupted

broad white postmedian band extending from mid-costa towards the tornus

turning sharply at vein 1A+2A to the inner margin; vein 1A+2A sometimes

suffused with brown scales; the white band is narrowest at the costa and at its

broadest between veins M; and CuA,; hindwing with a dark brown-black

tornal patch and a white line along the termen from the tornus to vein Mi.

Measurements (forewing length / wingspan). holotype 21.0 / 37.5 mm. Mean

20.9 / 37.6 mm (n = 32).

Male genitalia (Figs 24-26). The genitalia of H. h. hurama (Figs 21-23) and

H. h. territorialis are overall similar. In H. h. territorialis the valva is not as

humped posteriorly as in H. h. hurama and the apical dorsal margin is very

angular and produced, with a short projection at the end of the inner margin,

whereas in H. h. hurama it is more rounded. In H. h. territorialis, the saccus

is shorter and club-shaped compared with the thin and extended condition in

H. h. hurama. The uncus of H. h. territorialis is also broader and more

curved and talon-like than in H. h. hurama. The tegumen is also more pointed

in H. h. territorialis than in H. h. hurama. The aedeagi are similar in both

subspecies.

Female (Figs 19-20). Upperside: similar to male but without the forewing

sex-brand. Underside: similar to male but with a slightly broader white

136 Australian Entomologist, 2015, 42 (3)

postmedian band and hindwing ground colour with a brighter purplish sheen.

Measurements (forewing length / wingspan). 21.2 / 37.9 mm (n = 44).

Etymology. The subspecific epithet, an adjective, refers to it being found in

the Northern Territory, Australia.

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B.M, 1954-222.

PITE PEETER u TVI I á aS OO PVA l Í BIMINH E) 1623547

Figs 1-12. Hasora hurama subspecies type specimens from BMNH: (1) H. h. hurama

male upperside; (2) H. h. hurama male underside; (3) H. h. hurama label data; (4) H.

h. mola male upperside; (5) H. h. mola male underside; (6) H. h. mola label data; (7)

H. h. arua male upperside; (8) H. h. arua male underside; (9) Æ. h. arua label data;

(10) H. h. diana male upperside; (11) H. h. diana male underside; (12) H. h. diana

label data. All images @ Natural History Museum, London.

Australian Entomologist, 2015, 42 (3) 137

Figs 13-20. Hasora hurama subspecies from Australia: (13-16) H. h. hurama adults

from Queensland: (13) male upperside; (14) male underside; (15) female upperside;

(16) female underside. (17-20) H. h. territorialis subsp. n. adults from the Northern

Territory: (17) holotype upperside; (18) holotype underside; (19) paratype female

upperside; (20) paratype female underside.

138 Australian Entomologist, 2015, 42 (3)

25 26 N

Figs 21-26. Hasora hurama male genitalia: (21-23) H. hurama hurama from Innisfail,

Queensland: (21) lateral view of tegumen, uncus and valva; (22) lateral view of valva;

(23) aedeagus. (24-26) H. hurama territorialis subsp. n. from Fogg Dam, Northern

Territory: (24) lateral view of tegumen, uncus and valva; (25) lateral view of valva;

(26) aedeagus.

Diagnosis

Hasora hurama territorialis subsp. n. phenotypes (Figs 17-20) can be

separated from H. h. hurama (Figs 1-3, 13-16), H. h. arua (Figs 7-9), H. h.

diana (Figs 10-12) and H. h. kieta (Tennent 2002, plate 1, figs 28-29)

principally by the narrower white band of the hindwing underside. It is

closest to H. h. mola (Figs 4-6) but the white band on the hindwing underside

of H. h. territorialis is more uniformly narrow and not as pointed at the costa.

Additionally, specimens of H. h. territorialis generally have a purplish sheen

over the hindwing underside.

Biology

Food plant. Derris trifoliata Lour. (Fabaceae).

Life history. The life history of H. h. hurama was first described by Manski

(1940) and updated by Braby (2000). The immature stages of H. h.

territorialis are similar to those of H. h. hurama as described by Braby

(2000), with some minor morphological differences in larval and pupal

colouring and markings. In the final instar larva of H. h. territorialis (Figs

27-28) the head is paler brown and the body ground colour paler pinkish-

brown and lacking the prominent dorsolateral black patches on abdominal

segments found in H. h. hurama (Fig 27). The pupa of H. h. territorialis

(Figs 29-30) has paler abdominal segments with the spiracles pale brown as

opposed to brown-black in H. h. hurama (Fig 29). The black mid-dorsal line

on the thorax of H. h. hurama is lacking in the pupa of H. h. territorialis.

Australian Entomologist, 2015, 42 (3) 139

Figs 27-30. Hasora hurama larvae and pupae: (27) H. hurama hurama final instar

larva dorsal view; (28) H. hurama territorialis final instar larva dorsal view; (29) H.

hurama hurama pupa dorsolateral view; (30) H. hurama territorialis pupa lateral

view.

Ecology and behaviour. Eggs are laid on the new growth of the food plant,

which grows adjacent to mangroves or within mixed monsoon forest-

mangrove associations. Breeding areas are in estuarine habitats along river

banks in flood plains. Adults have been more commonly observed flying

along the mangrove margins, with a quick jerky flight, frequently settling on

the tendrils of the food plant or sticks protruding from the foliage. Once

disturbed the adults quickly take flight. The larvae make a series of shelters,

initially in the immature leaves, by joining the sides of the leaves together

with silk and then later in the mature leaves where pupation takes place.

Larvae of H. h. territorialis, like those of H. h. hurama, frequently consume

all available new growth of the food plant with the availability of new growth

affecting the size of the emergent adult. To date, adults have been recorded in

the Northern Territory in all months of the year, with the life cycle of the

butterfly being triggered by new growth on the food plant.

Discussion

Evans (1949) noted that H. h. mola was smaller and paler than H. h. hurama,

having a male forewing length of 22 mm as opposed to 23-25 mm in H. h.

hurama, and with no purple gloss on the hindwing underside. Evans (1949)

140 Australian Entomologist, 2015, 42 (3)

also noted that the white band on the hindwing underside in H. h. mola was

very narrow, tapering to 1⁄2 mm at the costa. Specimens of H. h. territorialis

are also smaller than specimens of H. h. hurama and also slightly smaller

than those of H. h. mola, having an average male forewing length of 20.9 mm

(n = 32), although this is probably not statistically significant. Specimens of

H. h. territorialis generally have a purplish sheen over the hindwing

underside that is absent in H. h. mola. The white band on the hindwing

underside in H. h. territorialis is a different shape to that of H. h. mola, being

more uniformly narrow, less swollen medially and not as pointed at the costa.

Hasora h. mola is also geographically isolated from H. h. territorialis,

occurring some 1300 km to the north of the Northern Territory.

Hasora hurama territorialis and H. h. hurama are allopatric and as such are

reproductively isolated. Hasora h. territorialis is distinct in its adult

phenotype and, although it shares the same food plant as H. h. hurama, its

larval and pupal stages are morphologically different. The food plant, Derris

trifoliata, has not been recorded from the Kimberley of Western Australia or

from the Gulf Country of southwestern Queensland (AVH 2015). The male

genitalia of H. h. territorialis are significantly different from those of H. h.

hurama.

In Australia, adults of H. h. territorialis (Figs 17-20) can be separated from

those of H. h. hurama (Figs 13-16) by the following characters. Size: H. h.

territorialis is generally smaller with an average forewing length / wingspan

of 20.9 / 37.6 mm (n = 32) in males and 21.2 / 37.9 mm (n = 44) in females,

compared with 22.9 / 41.9 mm (n = 17) and 22.5 / 41.5 mm (n = 22)

respectively in H. h. hurama. Colour: the upperside wing colour of H. h.

territorialis is dull chocolate-brown, whereas in H. h. hurama it is a rich dark

reddish-brown. The underside colour is also paler in H. h. territorialis than in

H. h. hurama, giving the appearance of a paler insect. Wing, head and thorax

hairs in H. h. territorialis are light brown and rarely have the greenish-bronze

colouration noted by Braby (2000) in specimens of H. h. hurama. Hindwing

underside: the white band in H. h. territorialis is approximately half the

width of the band in H. h. hurama, being on average 2.3 mm (n = 32) in

males and 2.6 mm (n = 44) in females at its widest, compared with 4.7 mm (n

= 17) and 4.6 mm (n = 22) respectively in H. h. hurama. The white band in

H. h. territorialis is, however, broader than in either H. chromus chromus

(Cramer, [1780]) or H. khoda haslia (Mabille, 1876), both also found in

Australia. The male forewing sex brand is similar in both subspecies.

The geographic isolation of the two populations of H. hurama in Australia

has undoubtedly contributed to the observed differentiation within each

region. Future surveys in coastal areas in the Kimberley region of Western

Australia and in the Gulf Country of northwestern Queensland would help to

understand the full extent of the butterfly’s distribution across northern

Australia.

Australian Entomologist, 2015, 42 (3) 141

Acknowledgements

We would like to thank Mark Hopkinson, Ted Edwards, You Ning Su, Gavin

Daly, Jared Archibald, Dr Michael Braby, Dr Dave Britton, John Chainey,

Dave Lane and Dr Trevor Lambkin: Mark Hopkinson, for the provision of

nominate race immature stages; Ted Edwards, for access to literature,

provision of ANIC specimen data, genitalia dissections and for his support in

the development of this paper; You Ning Su, for the genitalia photographs;

Gavin Daly and Jared Archibald, for the provision of data for specimens in

the NTM; Dr Michael Braby, for the provision of data for specimens in his

private collection and for constructive comments on an initial draft of the

manuscript; Dr Dave Britton, for the provision of data for specimens in the

AM; John Chainey, for the provision of photographs of the type specimens

held in the BMNH; Dave Lane for provision of data for specimens in his

private collection; and Dr Trevor Lambkin for permission to include his

unpublished Torres Strait records.

References

AVH [Australia’s Virtual Herbarium]. 2015. Atlas of Living Australia [Accessed 20 May 2015].

<http://www.avh.ala.org.au/occurrences/search?q=Isid%3Aurn%3Alsid%3A biodiversity.org.au

%3Aapni.taxon%3A684717&qc=data_hub_uid:dh2&fq=species:%22Derris%20trifoliata%22#

tab_mapView>

BISA, D. 2013. New locations of butterflies from northern Arnhem Land, Northern Territory.

Northern Territory Naturalist 24: 2-13.

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

CSIRO Publishing, Melbourne; xxvii + 976 pp.

COMMON, I.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Revised edition.

Angus & Robertson, Sydney; xiv + 682 pp.

EDWARDS, E.D., NEWLAND, J. and REGAN, L. 2001. Lepidoptera: Hesperioidea,

Papilionoidea. /n Wells, A.A. and Houston, W.W.K. (eds), Zoological Catalogue of Australia.

Vol. 31.6. CSIRO Publishing, Melbourne.

EVANS, W.H. 1949. A catalogue of the Hesperiidae from Europe, Asia, and Australia in the

British Museum (Natural History). British Museum (Natural History), London; xix + 502 pp, 53

pls.

EVANS, W.H. 1958. A new sub-species of Hasora hurama Butler (Lepidoptera: Hesperiidae).

Natural History Rennell Island, British Solomon Islands 2: 113, 1 pl.

MEYER, C.E., WEIR, R.P. and WILSON, D.N. 2006. Butterfly (Lepidoptera) records from the

Darwin region, Northern Territory. Australian Entomologist 33: 9-22.

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

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

PETERS, J.V. Notes on the distribution of Australian Hesperioidea and Papilionoidea.

Australian Zoologist 15: 178-184.

TENNENT, W.J. 2002. Butterflies of the Solomon Islands. Systematics and biogeography.

Storm Entomological Publications, Dereham, Norfolk; xxiii + 413 pp.

142 Australian Entomologist, 2015, 42 (3)

Appendix 1. Collection and external character data for Hasora hurama

hurama specimens examined.

No. = Voucher specimen number; A = Forewing length (mm); B = Wingspan (mm),

measured apex to apex; C = Hindwing underside band width (between veins CuA,

and M3); Y = yes; N = no; CEMC = CE Meyer Collection; SSBC = SS Brown

Collection, RPWC = RP Weir Collection. Illustrated male (Figs 13-14) = Table 1, No.

3. Illustrated female (Figs 15-16) = Table 2, No. 5.

Table 1. Male specimens of Hasora hurama hurama examined.

No. Collection data Reared A B C

(Y/N)

Cooktown, N QLD 4 111.1987 CEMC Y 22.55 42.0 4.5

Quarantine Bay via 19.vi.2008 CEMC Y 22.0 41.0 45

Cooktown, QLD

3 Cooktown, QLD 11.11.2009 CEMC Y 220 420 5.0

4 Cooktown, QLD 18.11.2009 CEMC Y 23.0 ` 42.0 5.0

5 Cooktown, QLD 18.11.2009 CEMC Y 21.5 40.5 4.5

6 Cooya Beach via Port 29.1.2009 CEMC N 240 440 5.0

Douglas, QLD

7 Cooya Beach via Port 29.1.2009 CEMC N 22.0 40.0 4.0

Douglas, QLD

8 Scott Street, Cairns, 30.1.2009 CEMC Y 22.0 41.0 4.0

QLD

9 Scott Street, Cairns, 30.1.2009 CEMC Y 21.5 39.0 4.0

QLD

10 ` Lockerbie, QLD 5-9.v.2003 RPWC N 23.0 ` 42.0 4.4

11 Quarantine Bay, QLD 24.v1.2006 RPWC Y 22.0 39.0 4.0

12 ` Ingham, QLD (Figured ` 27.111.1987 SSBC Y 240 42.0 5.0

in Braby 2000)

13 Ingham, QLD 27.i11.1987 SSBC Y 23.0 43.0 5.0

14 ` Ingham, QLD 27.111.1987 SSBC Y 25.0 45.0 6.0

15 Lockerbie Scrub, Cape 10.v.2003 SSBC N 25.5 46.0 5.2

York QLD

16 Quarantine Bay, 16.vi.2006 SSBC Y 23.0 43.0 5.0

Cooktown, QLD

17 Quarantine Bay, 25.v1i.2006 SSBC Y 23.0 41.0 5.0

Cooktown, QLD

Average sizes (n = 17): 22.9 41.9 4.7

Australian Entomologist, 2015, 42 (3)

Table 2. Female specimens of Hasora hurama hurama examined.

No.

Nn B L N =e

O won AD

Collection data

Cooktown, QLD

Quarantine Bay via

Cooktown, QLD

Quarantine Bay via

Cooktown, QLD

Cooktown, N. QLD

Cooktown, QLD

Scott Street, Cairns,

QLD

Scott Street, Cairns,

QLD

Scott Street, Cairns,

QLD

Scott Street, Cairns,

QLD

Scott Street, Cairns,

QLD

Cooktown, QLD

Quarantine Bay, QLD

Quarantine Bay,

Cooktown, QLD

Quarantine Bay,

Cooktown, QLD

Ingham, QLD

Average sizes (n = 22):

22.ix.1997

23.1x.1997

3.x. 1997

16.vi.2006

20.vi.2006

4 111.2007

12.11.2009

17.11.2009

30.1.2013

3.11.2013

4.xi.1987

24.vi.2006

1.v11.2006

28.1.2007

17.111.1987

CEMC

RPWC

SSBC

Reared

(Y/N)

143

144 Australian Entomologist, 2015, 42 (3)

Appendix 2. Collection and external character data for types of Hasora

hurama territorialis subsp. n. used for the description of the subspecies.

No. = Voucher specimen number; R’red or R’d = Reared; A = Forewing length

(mm); B = Wingspan (mm), measured apex to apex; C = Hindwing underside band

width (between veins CuA, and M3); Y = yes; N = no; ANIC = Australian National

Insect Collection; CEMC = CE Meyer Collection; DALC = DA Lane Collection;

NTMC = Museum and Art Gallery of the Northern Territory Collection, Darwin;

MFBC = MF Braby Collection, RPWC = RP Weir Collection; SSBC = SS Brown

Collection. Illustrated holotype male (Figs 17-18) = Table 1, No. 5. Illustrated

paratype female (Figs 19-20) = Table 2, No. 4.

Table 3. Male specimens of Hasora hurama territorialis subsp. n.

No. Collection data R’red A B C

(Y/N)

1 Buffalo Creek, NT 3.111.1992 CEMC N 21.0 390 2.5

2 Adelaide River, NT 22.13.1992 CEMC Y 210 390 25

3 Adelaide River, NT 23.1x.1992 CEMC Y 195 39.0 2.5

4 Adelaide River, NT 24.1x.1992 CEMC Y 20.0 355 2.0

5 Adelaide River, NT 4.11.1994 ANIC Y 21.0 375 2.5

(Figs 17-18)

6 Adelaide River, NT 5.11.1994 CEMC Y 20.0 340 2.5

7 Adelaide River Bridge on 23.iv.1995 CEMC Y 21.0 365 2.0

Arnhem Hwy, NT

8 Adelaide River Bridge on 6.1x.1998 CEMC Y 20.0 360 2.0

Arnhem Hwy, NT

9 Adelaide River hunting 10.iv.2009 CEMC Y 21.5 380 2.0

reserve, Arnhem Hwy via

Darwin, NT

10 Adelaide River Bridge 3.v.2013 CEMC Y 21.5 385 20

environs, Arnhem Hwy,

11 Adelaide River Bridge 14.v.2013 CEMC Y 20.5 385 30

environs, Arnhem Hwy,

12 Beatrice Hill, NT 1.111.2000 RPWC Y 20.0 350 20

13 Beatrice Hill, NT 1.111.2000 RPWC Y 20.5 355 2.0

14 Beatrice Hill, NT 1.111.2000 RPWC Y 22.0 370 2.5

15 Beatrice Hill, NT 1.111.2000 RPWC Y 225 385 25

16 Beatrice Hill, NT 1 111.2000 RPWC Y 215. 395, 20

17 Adelaide River, NT 6.11.2008 RPWC ¥ 20.0 345 2.0

18 Adelaide River, NT 5.1v.2009 RPWC Y 21.5 375 2.0

19 Fogg Dam, NT 22.iv.1991 SSBC N 215 420 3.5

20 Fogg Dam, NT 22.iv.1991 SSBC N 215 41.5 3.2

21 Beatrice Hill (Window 14.11.2009 NTMC N 221 359 28

on the Wetlands Visitor

Centre), Adelaide River

floodplain, NT

Australian Entomologist, 2015, 42 (3)

No. Collection data

22 12.64956°S, 131.31851°E 17.1.2009

Beatrice Hill, NT

23 12.64956°S, 131.31851°E 17.1.2010

Beatrice Hill, NT

24 ` 12.64956°S, 131.31851°E 23.1.2010

Beatrice Hill, NT (MFBC

00116)

25 12°39°36”S, 131°20°11”E 17.x1.2010

Adelaide River crssng,

Arnhem Hwy, NT

26 =12°39°36”S, 131920” 11”E 17.x1.2010

Adelaide River crssng,

Arnhem Hwy, NT (MFBC

00582)

27 129%39'36”S, 131920? 11”E 18.x1.2010

Adelaide River crssng,

Arnhem Hwy, NT (MFBC

00583)

28 12°3936”S, 131220 11”E 21.1x.2010

Adelaide River crssng,

Arnhem Hwy, NT (MFBC

00584)

29 12939*36”S, 131220 11”E 7.xii.2012

Adelaide River crossing,

Arnhem Hwy, NT (MFBC

00824)

30 12°3936”S, 131920 11”E 12.vii.2013

Adelaide River crossing,

Arnhem Hwy, NT (MFBC

00582)

31 12.40329°S, 132.96107°E 13.x.2013

East Alligator, near Ubirr,

Kakadu NP, NT (MFBC

00896)

32 12.40329°S, 132.96107°E 16.x.2013

East Alligator, near Ubirr,

Kakadu NP, NT (MFBC

00896)

33 Blyth River' 21.ix.1968

Average sizes (n = 32):

MFBC

AMC

21.6

21.3

20.5

20.0

19.3

21.1

20.2

20.9

37.8

39.3

37.2

36.8

35.1

37.5

38.3

36.6

37.6

Note 1: specimen measurement data not used in calculation of average sizes.

145

2.2

2.3

2.2

2.0

1.7

1.8

2.0

2.9

2.3

146

Australian Entomologist, 2015, 42 (3)

Table 4. Female specimens of Hasora hurama territorialis subsp. n.

No.

A UNS

Collection data

Fogg Dam, NT

Adelaide River, NT

(Figs 19-20)

Adelaide River, NT

Adelaide River Bridge on

Arnhem Hwy, NT

Adelaide River Bridge on

Arnhem Hwy, NT

Adelaide River Bridge on

Arnhem Hwy, NT

Adelaide River Bridge on

Arnhem Hwy, NT

Adelaide River Bridge

environs, Arnhem Hwy, NT

Adelaide River Bridge

environs, Arnhem Hwy, NT

Adelaide River Bridge

environs, Arnhem Hwy, NT

Adelaide River, NT

Fogg Dam, NT

Berrimah Research

Farm, NT

Berrimah Research

Farm, NT

Berrimah Research

Farm, NT

10 km East, Adelaide

River, NT

Beatrice Hill, NT

Adelaide River, NT

Fogg Dam, NT

Adelaide River, NT

Adelaide River, 10 km NE

on Arnhem Hwy, NT

22.1v.1991

24.13.1992

10.11.1994

16.1v.1995

18.1v.1995

18.1x.1998

13.v.2013

14.v.2013

15.v.2013

8.v.1989

22.1v.1991

8.1v.1994

15.1v.1994

7.v111.1996

1.111.2000

5.iv.2009

5.111.2013

22.1v.1991

2.v.1995

9.v.1991

CEMC

RPWC

SSBC

DALC

NTMC

R'd

qe Zaz

ZZ ZZ

ZZZ Z ZKKK

22.0

21.0

22.0

21.0

20.0

20.5

20.0

21.0

20.0

21.0

22.0

22.5

23.0

21.5

23.0

21.5

22.0

21.0

19.0

21.0

23.0

21.0

22.0

22.5

21.2

22.3

38.5

39.0

35.0

35.5

36.0

38.0

36.0

38.0

39.0

38.0

38.5

41.5

35.0

40.0

36.0

38.5

37.0

38.5

36.5

33.0

37.0

44.0

40.0

40.2

42.5

37.0

41.2

3.0

2.5

3.0

2.0

3.0

2.0

3.0

2.5

3.0

2.0

3.5

3.0

3.5

2.1

3.0

Australian Entomologist, 2015, 42 (3)

No.

Collection data

12.64956°S 131.31851°E

Beatrice Hill, NT (MFBC

00428)

12939”36”S 131°20°11”E

Adelaide River crssng,

Arnhem Hwy, NT (MFBC

00585)

12°39°36”S 131220? 11”E

Adelaide River crssng,

Arnhem Hwy, NT (MFBC

00586)

12°39°36”S 131°20°11”E

Adelaide River crssng,

Arnhem Hwy, NT

12°39°36”S 131°20°11”E

Adelaide River crssng,

Arnhem Hwy, NT (MFBC

00587)

12°39°36”S 131°20°11”E

Adelaide River crssng,

Arnhem Hwy, NT (MFBC

00588)

12°39°36"S 131°20°11”E

Adelaide River crssng,

Arnhem Hwy, NT (MFBC

00589)

12°39°36"S, 131920 11”E

Adelaide River crossing,

Arnhem Hwy, NT

12.40329°S, 132.96107°E

East Alligator, near Ubirr,

Kakadu NP, NT

12.40329°S 132.96107°E

East Alligator, near Ubirr,

Kakadu NP, NT

12.40329°S 132.96107°E

East Alligator, near Ubirr,

Kakadu NP, NT

12.40329°S 132.96107°E

East Alligator, near Ubirr,

Kakadu NP, NT (MFBC

00897)

Fogg Dam, NT

Adelaide River'

Average sizes (n = 44):

17.1.2019

15.x1.2010

17.xi.2010

17.x1.2010

18.x1.2010

19.x1.2010

27.x1.2010

16.v11.2013

20.x.2013

28.x.2013

30.x.2013

22.iv.1991

13.1.1962

MFBC

ANIC

AMC

R’d

19.9

20.9

21.9

19.9

20.9

20.2

21.4

19.4

19.7

20.7

20.3

20.5

22.3

21.2

36.8

39.4

40.6

34.9

38.7

37.5

38.9

36.1

35.7

37.4

36.5

37.9

Note 1: specimen measurement data not used in calculation of average sizes.

147

2.2

2.7

2.8

2.5

XT]

2.2

2.0

2.2

2.3

1.7

2.0

2.5

2.6

148 Australian Entomologist, 2015, 42 (3)

MISCELLANEOUS NOTES

The following notes are abstracted from the News Bulletin of the

Entomological Society of Queensland and appeared, with illustrations, in the

Volumes and Parts indicated.

Anthene lycaenoides godeffroyi (Semper) (Lepidoptera: Lycaenidae). — Three worn

females were collected on 24 January 2015 in southern Queensland, at two different

locations separated by about 10 km, approximately 800 km south of the previous

southern recorded limit [Cannonvale near Airlie Beach]. Several others were seen at

close quarters and positively identified but not collected.

Two specimens were collected and several others identified 5 km east of Pomona and

another collected 2 km south of Cooroy. All specimens seen or collected were

females. The Pomona specimens were all active late morning, flying around and

landing on a flowering Albizia julibrissin (Mimosaceae), a recorded food plant for this

species. The specimens appeared to be coming from a northerly direction. No further

specimens were encountered until 30 January 2015, when a further female was

collected and released at the Pomona site.

All specimens collected have a prominent white patch on the forewing upperside and

appear typical of specimens from Townsville and Cairns.

While A. lycaenoides godeffroyi is a relatively strong flier for a lycaenid, it is

considered unlikely that these specimens originated from within its previously known

range, given the significant distance involved. It appears more likely that the range of

this species has been expanding incrementally without previous detection.

MAYO, R. — Interesting new locations for Anthene lycaenoides godeffroyi (Semper)

in southern Queensland. — Volume 42(10): 189 (February 2015).

Hypolimnas alimena lamina Fruhstorfer (Lepidoptera: Nymphalidae) — A single

female was observed 5 km east of Pomona on 20 April 2015. This is the first

specimen observed in the area since moving there in 2002. It was of the ‘brown form’

and appeared to be in relatively fresh condition.

Junonia orithya albicincta Butler (Lepidoptera: Nymphalidae) — Three males were

observed 5 km east of Pomona on separate single occasions, between 6 and 20 April

2015. These are the first specimens observed in the area since moving there in 2002.

All flew rapidly close to the ground without stopping and appeared to be travelling in

a generally southern direction. Also encountered regularly over the past decade near

Imbil, where it appears to be established with males exhibiting a rapid low flight but

often alighting on the ground (unlike the Pomona specimens).

Anthene lycaenoides godeffroyi (Semper) (Lepidoptera: Lycaenidae). — A further

female was observed and positively identified at Buderim on 3 May 2015.

MAYO, R. — Recent observations of butterfly species rarely encountered in southern

Queensland. — Volume 43(3): 50 (May 2015).

Australian Entomologist, 2015, 42 (3): 149-159 149

ADDITIONAL POLYMORPHISM IN FEMALES OF

HYPOLIMNAS BOLINA PALLESCENS (BUTLER)

(LEPIDOPTERA: NYMPHALIDAE) FROM THE ISLANDS

OF TAVEUNI AND VANUA LEVU, FIJI

R.B. LACHLAN

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

Abstract

Numerous additional polymorphic Hypolimnas bolina pallescens (Butler, 1874) female forms

were collected on the central west coast of Taveuni during May 1982 and from a small area

approximately 5 km southwest of Savusavu on the south central coast of Vanua Levu, Fiji,

during November-December 2014. Their polymorphism is discussed and the morphs not

previously collected on Viti Levu by the author are illustrated.

Introduction

In Fiji, Hypolimnas bolina pallescens (Butler) is the subspecies of H. bolina

(L.) assigned to this region of the Pacific. Adults of H. bolina are sexually

dimorphic but the females in Fiji, Samoa and Guam (the largest island in

Micronesia, in the Western Pacific), in particular, display an extraordinary

range of polymorphism apparently not seen to the same degree anywhere else

across its very extensive range (Clark and Sheppard 1975).

The primary purpose of this paper is to illustrate additional female morphs

not previously collected and illustrated (Lachlan 2014) from Fiji. Once again,

all specimens collected on both islands were within small areas of forest and

coconut plantations measuring just a few hectares in area.

Hypolimnas bolina pallescens adults are generally widespread and locally

common in Fiji and can best be found along forest trails, clearings and

roadsides where they usually fly close to the ground and often rest on low

vegetation. The author has noted that, if disturbed, females will often fly for a

short period and look for a resting place. Having collected and examined 244

females from three different islands in Fiji, it is clear that, while most morphs

can be placed with reasonable accuracy into the four main forms given by

Poulton (1924) to describe the four basic phenotypes, placing the large

number of intermediate morphs the author has collected into the three

additional intermediate forms given by Clark and Sheppard (1975) has

proved, with many morphs, much more difficult and rather subjective at

times. There are also a small number of uniquely coloured and patterned

morphs that are difficult to place in any of the seven current groupings.

Surveys

The brief survey on the central west coast of the island of Taveuni (the third

largest island in Fiji) was done from 11 to 20 May 1982. Being at the end of

the wet season, most specimens collected showed signs of wear and no fresh

Specimens were taken. An intensive survey was also conducted near

Savusavu on the south central coast of Vanua Levu, the second largest island

150 Australian Entomologist, 2015, 42 (3)

in Fiji, from 30 November to 13 December 2014. Unfortunately, most days

were overcast with rain periods and only intermittent hours of sunshine. H. b.

pallescens adults were commonly encountered not only along the forest trails

but also in a nearby coconut plantation, where they flew close to the ground

generally landing frequently. Good rains had fallen about two weeks prior to

the commencement of the survey and most specimens collected and observed

showed wear and wing damage, indicating they had been on the wing for

some time. As further rain fell regularly during the survey period more fresh

specimens appeared. Numerous morphs not taken by the author earlier in

2014 on Viti Levu were collected. Vanua Levu is approximately 63 km NE

of Viti Levu (the main island in Fiji) while Taveuni is just 8 km SE of the

eastern side of Vanua Levu.

Hypolimnas bolina pallescens (Butler, 1874)

(Figs 1-22)

Material examined. Taveuni survey: 14 GG, 25 99, FIJI: Central west coast of

Taveuni Island, between 11-20.v.1982, R.B. Lachlan. (All in RBL collection).

Vanua Levu survey: 4 SZ, 112 9 Q, FIJI: 5 km SW of Savusavu, south central coast

of Vanua Levu, 16°48°58”S, 179°17°20”E, between 30.x1.-13.x11.2014, R.B. Lachlan.

(All in RBL collection).

The various female forms collected during both surveys and their numbers

are recorded in Table 1. As the Taveuni survey was brief and done over 30

years ago, an accurate comparative analysis of any differences between the

two islands is not worthwhile, other than to note that no morphs of the forms

euploeoides-pallescens or pallescens were collected on Taveuni.

Specimen sizes. All measurements are set wingspans. The largest from

Taveuni was 82 mm, the smallest was 68 mm and the average (n = 25) was

73.4 mm. The largest from Vanua Levu was 83 mm (this is very large for the

species) and the smallest was 61 mm.

Table 1. Results for Hypolimnas bolina pallescens females from Taveuni and

Savusavu, Vanua Levu.

Form Numbers collected / %

Taveuni Savusavu, Vanua Levu

euploeoides 6 / 24 28 / 25

euploeoides-naresi 6 / 24 10/9

naresi 3/12 15 /13.3

euploeoides-nerina 3/12 43 / 38.4

nerina 7/28 6/5.3

euploeoides-pallescens 0/0 1/0.8

pallescens 0/0 9/8

Australian Entomologist, 2015, 42 (3) 151

Figs 1-3. Females of Hypolimnas bolina from Fiji: f. naresi.

152 Australian Entomologist, 2015, 42 (3)

Figs 4-6. Females of Hypolimnas bolina from Fiji: f. euploeoides-nerina.

Australian Entomologist, 2015, 42 (3) 153

Figs 7-9. Females of Hypolimnas bolina from Fiji: f. euploeoides-nerina.

154 Australian Entomologist, 2015, 42 (3)

Figs 10-12. Females of Hypolimnas bolina from Fiji: f. euploeoides-nerina.

Australian Entomologist, 2015, 42 (3) 155

Figs 13-15. Females of Hypolimnas bolina from Fiji: f. euploeoides-nerina.

156 Australian Entomologist, 2015, 42 (3)

Figs 16-18. Females of Hypolimnas bolina from Fiji: (16-17) f. euploeoides-nerina;

(18) f. euploeoides-pallescens.

Australian Entomologist, 2015, 42 (3) 157

SS SS

Ë ` \ i

21 4

Figs 19-21. Females of Hypolimnas bolina from Fiji: f. pallescens.

158 Australian Entomologist, 2015, 42 (3)

Fig 22. Female of Hypolimnas bolina from Fiji: f. pallescens.

Discussion

Having now examined 244 female H. b. pallescens adults as a result of field

work conducted in Fiji over a number of years, it has become increasingly

obvious to the author that they display such an astonishing degree of

polymorphism that it was described by Vane-Wright et al. (1977) as being

quasi-continuous — without ceasing. Given that in many parts of the Pacific

this is the case, relying on changes in individual wing patterns and colours to

describe subspecies or particular forms is not possible as there are so many

intermediates between the four basic phenotypes. Vane-Wright and Tennent

(2011) correctly noted that, in the case of Junonia villida (Fabricius),

‘systematists working from short series collected during rare visits to remote

islands could be misled into thinking that they were dealing with stable,

locally adapted subpopulations worthy of subspecific status.’ The same must

also be said for the past use of form names that have been applied to various

H. bolina females.

Acknowledgements

I am grateful to John Tennent (Natural History Museum, London) for

providing important reference material. I would also like to thank Russel Cox

(Entomology Department, Australian Museum, Sydney) for his help in

producing the digital images used in the plates.

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160 Australian Entomologist, 2015, 42 (3)

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