Australian Entomologist, 2015, 42 (2): 53-58 53

A NEW SPECIES OF POLYRHACHIS (HAGIOMYRMA) WHEELER

FROM THE NORTHERN TERRITORY, AUSTRALIA

(HYMENOPTERA: FORMICIDAE: FORMICINAE)

BENJAMIN D. HOFFMANN

CSIRO Land and Water Flagship, Tropical Ecosystems Research Centre, PMB 44, Winnellie,

NT 0822 (Email: Ben. Hoffmann@csiro.au)

Abstract

Polyrhachis kohouti, a new species of the penelope species-group of the subgenus Hagiomyrma

Wheeler, is described from Arnhem Land in Australia’s Northern Territory. The new species is

illustrated and characters distinguishing it from other described species of the group are

provided.

Introduction

The subgenus Hagiomyrma Wheeler of the genus Polyrhachis Fr. Smith was

recently revised by Kohout (2013). He recognised 48 species as bona fide

members of the subgenus and organised them into six species-groups, the

ammon, metella, penelope, schenkii, trapezoidea and tubifera groups. One

species missed by the recent revision was a Hagiomyrma species belonging

to the penelope species-group that I had collected by hand in Arnhem Land in

the Northern Territory. The new species is described and illustrated below

and characters separating it from other members of the pene/ope-group are

provided.

Methods

Photographs of the holotype were taken by Geoff Thompson (QMBA) on a

Visionary Digital BK-plus Lab System using a Canon EOS 5D MkII camera.

The source images were adjusted and exported from Adobe Lightroom, focus

stacked with Zerene Stacker software and formatted with Adobe Photoshop

(Adobe Systems Inc., USA).

Abbreviations for Australian institutions and depositories (with the names of

co-operating curators): ANIC — Australian National Insect Collection,

CSIRO, Canberra, ACT (Dr Beth Mantle); QMBA — Queensland Museum,

Brisbane, QLD; TERC — Tropical Ecosystems Research Centre, CSIRO,

Darwin, NT (Prof. A.N. Andersen).

Standard measurements and indices follow those of Kohout (2013): TL —

Total length (the necessarily composite measurement of the entire ant

measured in profile); HL — Head length (the maximum measurable length of

the head in perfect full face view, measured from the anterior-most point of

the clypeal border or teeth, to the posterior-most point of the occipital

margin); HW — Head width (width of the head in perfect full face view,

measured immediately in front of the eyes); CI — Cephalic index (HW x

100/HL); SL — Scape length (length of the antennal scape, excluding the

condyle); SI — Scape index (SL x 100/HW); PW — Pronotal width (greatest

width of the pronotal dorsum, including the pronotal teeth, or across the

54 Australian Entomologist, 2015, 42 (2)

humeri in species without teeth); MW — Mesonotal width (minimum width of

the mesonotal dorsum measured along the metanotal groove when viewed

from behind); PMI — Promesonotal index (PW x 100/MW); MTL -

Metathoracic tibial length (maximum measurable length of the tibia of the

hind leg). All measurements were taken using a Zeiss stereomicroscope with

an eyepiece calibrated against a stage micrometer. All measurements are

expressed in millimetres (mm).

Systematics

Genus Polyrhachis Fr. Smith, 1857

Polyrhachis Fr. Smith, 1857: 58. Type species: Formica bihamata Drury, 1773: 73,

pl. 38, figs 7, 8, worker; by original designation.

Subgenus Hagiomyrma Wheeler, 1911

Hagiomyrma Wheeler, 1911: 860 (as subgenus of Myrma Billberg, 1820 [sensu

Wheeler, 1911: 859] = Polyrhachis Fr. Smith, 1857). Type species: Formica ammon

Fabricius, 1775: 394, worker; by original designation.

(For full synonymy citations see Kohout, 2013: 489.)

Polyrhachis kohouti sp. n.

(Figs 1-4)

Types. Holotype worker: AUSTRALIA (NORTHERN TERRITORY): Arnhem Land,

Balkpalkbuy, 60 km SW of Nhulunbuy, 12°35'55"S, 136°31'25"E, elevation 56 m.

22.111.2006, B.D. Hoffmann. Paratype: 1 worker, data as for holotype. Holotype in

ANIC; paratype in TERC.

Description. Worker. Dimensions (holotype cited first): TL c. 6.95, 5.80; HL

1.65, 1.50; HW 1.40, 1.25; CI 85, 83; SL 1.75, 1.59; SI 125, 127; PW 1.34,

1.20; MW 0.75, 0.72; PMI 179, 167; MTL 1.93, 1.78 (1+1 measured).

Median flange of anterior clypeal margin with two distinct acute teeth

medially, laterally flanked by acute, somewhat laterally directed teeth.

Clypeus with median, anteriorly distinct, longitudinal carina; straight in

profile, posteriorly rounding into moderately impressed basal margin. Frontal

triangle poorly indicated. Frontal carinae weakly raised; central area

relatively wide, rather flat, with only anteriorly indicated frontal furrow.

Sides of head in front of eyes converging towards mandibular bases in

virtually a straight line; behind eyes sides rounding into convex occipital

margin. Eyes moderately convex, in full face view marginally exceeding

lateral cephalic outline. Ocelli lacking, position of median ocellus indicated

by a shallow pit in cephalic structure. Pronotal dorsum distinctly wider than

long with humeri widely rounded, dorsally shallowly concave; lateral

pronotal margins weakly raised, converging posteriorly towards clearly

impressed promesonotal suture. Mesonotal dorsum with margins converging

posteriorly towards indistinct metanotal groove. Propodeal dorsum with

lateral margins terminating in moderately long, rather strongly divergent

Australian Entomologist, 2015, 42 (2) 55

spines with tips weakly turned outwards. Propodeal spiracles situated on

moderately projecting tubercles. Petiole with dorsum deeply concave

medially, armed with pair of divergent, rather massive, bull horn-shaped

spines with tips turned downwards and slightly outwards (fig. 2). Anterior

face of first gastral tergite widely rounding onto dorsum.

Mandibles finely, longitudinally striate with numerous piliferous pits. Head,

mesosoma and petiole finely reticulate-punctate; somewhat semipolished on

vertex of head and dorsum of mesosoma; sides of head and dorsum of spines

more coarsely sculptured. Gaster finely shagreened.

Figs 1-4. Polyrhachis (Hagiomyrma) kohouti sp. n., holotype worker: (1) head in full

face view; (2) petiole in dorsal view (not to scale); (3) dorsal view; (4) lateral view.

Mandibles with numerous golden hairs, longer and curved at masticatory

border, shorter and anteriorly inclined towards mandibular bases. Anterior

clypeal margin with several anteriorly directed setae medially and a few

shorter setae fringing margin laterally. Several, mostly paired, relatively short

hairs on clypeus, along frontal carinae and on sides of head, with distinctly

longer, anteriorly inclined hairs on vertex and along occipital border. Dorsum

of mesosoma and petiole, including spines, with numerous, variously

inclined, mostly silvery hairs, longest more than half greatest diameter of

eyes. Gaster with abundant, rather long, silvery or pale golden, posteriorly

inclined hairs. Closely appressed, silvery pubescence, very sporadic over

56 Australian Entomologist, 2015, 42 (2)

dorsal body surfaces; pubescence somewhat denser on sides of mesosoma

and posterior face of petiolar dorsum and spines. Dorsum of gaster with

somewhat longer and more abundant silvery pubescence, almost hiding

underlying sculpturation.

Body black; appendages very dark reddish-brown or black.

Etymology. Named after Rudy Kohout in recognition of his outstanding

contribution to Polyrhachis taxonomy.

Remarks. Polyrhachis kohouti can be easily recognised by the form of its

petiole which has a deeply medially concave dorsum and is armed with rather

massive, bull horn-shaped spines (Fig. 2). The petiole in lateral view is

relatively slender, in contrast to other species of the penelope-group where

the posterior face of the petiole is usually more-or-less convex or distinctly

swollen. Other distinguishing characters of P. kohouti include the strongly

transverse pronotal dorsum with widely rounded humeri and strongly

posteriorly converging lateral margins that are shallowly emarginate before

terminating at the distinctly impressed promesonotal suture. The presence of

numerous setae on most of the body surfaces also separates P. kohouti from

other species of the penelope-group, which lack hairs, except P. hoffmanni

which has distinctly longer and sparser hairs on the body. In the key to

Hagiomyrma species in Kohout (2013), Polyrhachis kohouti runs to couplet

26 and can be identified by the following modification and insertion of a

couplet. Figure numbers in square brackets refer to illustrations in the

original article (Kohout 2013).

26 Outline of head in full face view with numerous bristle-like hairs fringing

margin between eyes and mandibular bases ...... 0.. 27a

— Outline of head in full face view without hairs, or at most with only a

few inconspicuous, very short hairs fringing margin between eyes and

mandibular bases 2.0.0.0... cece cece cccccecceceessseseessseeeseesesesesssaeeesseeeensees 27

27 Head, mesosoma and petiole very distinctly and evenly, rather coarsely,

reticulate-punctate; petiole in profile with posterior face distinctly convex

and swollen towards base [Figs 19G-H] (Nth Qld) ..... P. tenebra Kohout

— Head, mesosoma and petiole finely reticulate-punctate with sculpture

somewhat longitudinally rugulate-striate on vertex of head; petiole in

profile with posterior face only weakly convex [Figs 9A-B] (New

Ireland, New Guinea, Cape York Peninsula) .........000...cccccccceecceeseeeeeenes

bes Shean Died baru thy: EVE AA ETA EO hy ch Me P. semiobscura Donisthorpe

27a Petiole armed with rather massive, bull horn-shaped spines (Fig. 2) (NT

ROLES ELO CPLL O AE P E E IA TE ON P. kohouti sp. n.

— Petiole armed with slender, acute spines, mostly curved into shape of

first gastral segment or downcurved [e.g. Figs 9C, 19A] ....... 28

Australian Entomologist, 2015, 42 (2) 57

Polyrhachis kohouti is currently known only from the type locality, with both

specimens collected when their nest was disturbed by overturning a log. The

species appears to be uncommon because extensive ant collections

throughout NE Arnhem Land conducted since 2003, including in the type

locality, have failed to find this species again. The vegetation of the type

location is open savanna woodland dominated by Eucalyptus oligantha with

an understorey of dense grasses, on gently sloping, seasonally waterlogged,

loamy soil (Fig. 5).

Fig. 5. Seasonally waterlogged savanna woodland in Arnhem Land, the type locality

of P. kohouti sp. n.

Acknowledgements

I am very grateful to the Yolngu of NE Arnhem Land for providing me

access to their lands, especially BaluPalu Yunupingu who took me to the

location where P. kohouti was found. I thank Geoff Thompson (QMBA) for

producing the digital images used for illustrations. Thanks also to Dr Chris

Burwell (QMBA) for reading and commenting on a draft manuscript.

References

BILLBERG, G.J. 1820. Enumeration Insectorum in Museo Gust. Joh. Billberg: Stockholm; 138

Pp.

DRURY, D. 1773. Illustrations of Natural History. Wherein are exhibited upwards of two

hundred and twenty figures of exotic insects 2. London; 90 pp.

FABRICIUS, J.C. 1775. Systema Entomologiae, systens insectorum classes, ordines, genera,

species, adiectis, synonymis, locis, descriptionibus, observationibus. Flensburgi et Lipsiae; 832

Pp.

58 Australian Entomologist, 2015, 42 (2)

KOHOUT, R.J. 2013. Revision of Polyrhachis (Hagiomyrma) Wheeler, 1911 (Insecta:

Hymenoptera: Formicidae: Formicinae). Memoirs of the Queensland Museum — Nature 56(2):

487-577, 14 figs.

SMITH, FR. 1857. Catalogue of the hymenopterous insects collected at Sarawak, Borneo;

Mount Ophir, Malacca; and at Singapore, by A.R. Wallace. Journal of Proceedings of the

Linnean Society of London, Zoology 2: 42-88.

WHEELER, W.M. 1911. Three formicid names which have been overlooked. Science (N.S.) 33:

858-860.

Australian Entomologist, 2015, 42 (2): 59-62 59

THE FEMALE OF DEUDORIX CLEORA MILLER & MILLER, 1986

(LEPIDOPTERA: LYCAENIDAE)

CHRIS J. MULLER

Honorary Associate, Australian Museum, 6 College Street, Sydney, NSW 2010

(Address for correspondence: PO Box 3228, Dural, NSW 2158) (Email:

chrismuller999@gmail.com)

Abstract

The hitherto unknown female of Deudorix cleora Miller & Miller, endemic to Sulawesi,

Indonesia, is described and figured. New records for the species from central and southern

Sulawesi are presented and the male is illustrated in colour for the first time.

Introduction

Deudorix Hewitson, 1863 is a large genus belonging to the tribe Deudorigini,

with representatives in the Afrotropical, Oriental and Australian regions. A

number of new species have been described in recent years from the latter

region, including the south-west Pacific (Tennent 2000, Yagashita 2006,

Tennent 2008, Tennent et al. 2010).

Deudorix cleora Miller & Miller, 1986, arguably one of the most striking

members of the genus, was previously known only from the holotype male,

collected by Robert G. and Cleora M. Wind in June, 1940. It was among

material presented to the Allyn Museum of Entomology (now incorporated

into the McGuire Centre, Florida State Museum) by the couple following

their trip to the Indonesian Archipelago in 1939-40 and is held in that

institution (Miller and Miller 1986). Robert requested that the new butterfly

be named after his wife, a request that was honoured after both had died. A

female of D. cleora, in the private collection of Akira Yagashita (AYC),

Ibaraki, Japan, appears to be the only one known and is described here.

Vein description follows that of the numerical notation of Corbet and

Pendlebury (1993).

New records

Deudorix cleora was previously only known from the type locality at Lake

Dano, near Tondana, north Sulawesi. Additional new records include one

male from Palopo, central Sulawesi, taken by a local collector (in Chris J.

Muller reference collection) as well as the female described here from the

same locality. There are also a few males of D. cleora in the reference

collections of the author (CJMC) and that of Ken Thorne (KTC), Canada,

from southern Sulawesi, implying that the species has a relatively wide

distribution in mainland Sulawesi.

Although no colour illustration of D. cleora appears to have ever been

published, one of the latter male specimens appears on the Wikispecies url

(species. wikimedia.org/wiki/Deudorix_cleora).

60 Australian Entomologist, 2015, 42 (2)

Deudorix cleora Miller & Miller

(Figs 1-4)

Material examined. INDONESIA (SULAWESI): 1 ĝ, Palopo, central Sulawesi, Feb.

2009 (in CJMC); 1 9, Palopo, central Sulawesi, Aug. 2007 (in AYC). 6 ¢4, Maaki,

southern Sulawesi, 1995 (in KTC); 1 3, Maaki, southern Sulawesi, 1995 (in CJMC).

Description of female (Figs 3-4). Measurements: forewing length 25 mm;

antenna length 12 mm. Head dark brown, frons white; labial palpus white,

eye ringed narrowly with white; antennae black, ringed weakly with grey and

tipped with orange-brown. Thorax dark brown above, beneath grey-brown.

Abdomen dark brown-black above, pale yellow-brown beneath.

Forewing with costa bowed near base, termen convex, inner margin nearly

straight; upperside medium brown, broadly darker brown along costa and

termen; a pale yellow-brown median patch extending from sub-basal to

postmedian area and occupying part of spaces 1b, 2 and 3 and narrowly into

the cell and space 4; cilia medium brown; underside background colour

medium grey-brown, apex and termen darker brown; a faint narrow

submarginal brown band; a subapical band of four sub-rounded dark brown

spots between 1 and 2 mm in diameter, one each in spaces 4, 5, 6 and 9; a

postmedian band of three sub-rounded dark brown spots between 1 and 3 mm

in diameter, one each in spaces 1b, 2 and 3; a sub-rounded dark brown spot

approximately 2 mm in diameter at end of cell and discocellulars; cilia

medium brown.

Hind wing with thread-like tail (9 mm long) at vein 2; termen slightly

produced at vein 3; a prominent lobe adjacent to vein 1b; upperside medium

brown, veins dark brown; tail black, tipped with cream-brown; lobe light

blue, brown basally and with broad black border; cilia medium brown;

underside background colour medium grey-brown; termen darker brown; a

faint narrow submarginal brown band; a large subtornal black spot between

veins 2 and 3, rimmed with light orange and black crescents basally; entire

ground colour in tornal area washed with iridescent blue-green; a postmedian

band of elongated brown-black spots extending from inner margin to vein 7,

roughly parallel to inner margin and termen, a large dark brown median spot

in space 7; tail black, tipped with cream-brown; lobe black, dusted with light

blue at margins; cilia light brown at apex and inner margin, light grey-white

in tornal area.

Discussion

Deudorix cleora is a large, highly distinctive taxon that shows some

relationship to the Sundaic D. hypargyria Elwes [1893] and Philippine D.

philippensis Schroder, Treadaway & Hayashi, 1981. However, the males of

both the latter species are predominantly orange and brown above and both

sexes are comparatively unmarked on the undersides.

Australian Entomologist, 2015, 42 (2) 61

Figs 1-4. Deudorix cleora adults: (1) male upperside; (2) female upperside; (3) male

underside; (4) female underside. Scale bar = 10 mm.

As with most members of the tribe Deudorigini, males and females of D.

cleora differ markedly on their uppersides, while the undersides are similar.

However, the underside ground colour of the female is much darker than that

of the male, being grey-brown rather than the grey-white of the male. The

tornal spot is larger in the female and the tornal iridescence is a deeper blue-

green, rather than the light turquoise colouration of the male. The female

upperside of D. cleora resembles closely that of the orange female form of

Rapala ribbei Rober, 1886, another lycaenid endemic to Sulawesi, being

comparable in size and upperside pattern and colouration.

Butterfly endemism in Sulawesi is pronounced, being approximately 40% at

the species level (Vane-Wright and de Jong 2003). Reconstructed

phylogenies of several Indo-Pacific butterfly genera demonstrate that certain

62 Australian Entomologist, 2015, 42 (2)

endemic Sulawesi taxa represent ancient, early diversifying lineages (Vane-

Wright & de Jong 2003, Müller et al. 2010, Müller and Beheregaray 2010,

Müller et al. 2013). Based on the distinctive characters of D. cleora, it is

possible that this taxon has also been long isolated on Sulawesi and may well

belong to a primitive lineage within the genus.

Acknowledgements

Akira Yagashita, Ibaraki, Japan, kindly allowed the author to work on the

female described in this paper. Alan Cassidy, Berkshire, England, located

additional male specimens of D. cleora. Ken Thorne, Canada, generously

provided additional D. cleora male material to the author.

References

CORBET, A.S. and PENDLEBURY, H.M. 1993. The butterflies of the Malay Peninsula.

Malayan Nature Society, Kuala Lumpur; 4th edition, revised by J.N. Eliot; 597 pp.

MILLER, L.D. and MILLER, J.Y. 1986. A New thecline from Sulawesi (Lepidoptera:

Lycaenidae: Theclinae). Insecta Mundi 1(3): 143-145.

MULLER, C.J. and BEHEREGARAY, L.B. 2010. Palaeo island-affinities revisited —

biogeography and systematics of the Indo-Pacific genus Cethosia Fabricius (Lepidoptera:

Nymphalidae). Molecular Phylogenetics and Evolution 57: 314-326.

MULLER, C.J., MATOS-MARAVIL, P.F. and BEHEREGARAY, L.B. 2013. Delving into Delias

Hubner (Lepidoptera: Pieridae): fine-scale biogeography, phylogenetics and systematics of the

world’s largest butterfly genus. Journal of Biogeography 40(5): 881-893.

MULLER, C.J, WAHLBERG, N. and BEHEREGARAY, L.B. 2010. ‘After Africa’ — The

evolutionary history and systematics of the genus Charaxes Ochsenheimer (Lepidoptera:

Nymphalidae) in the Indo-Pacific Region. Biological Journal of the Linnean Society 100: 457-

481.

TENNENT, W.J. 2000. Notes on Deudorix Hewitson in the Solomon Islands, the Bismarck

Archipelago and New Guinea, with description of nine new taxa (Lepidoptera: Lycaenidae).

Australian Entomologist 27(1): 9-26.

TENNENT, W.J. 2008. A new Deudorix Hewitson, 1863, from New Caledonia and the Loyalty

Islands (Lepidoptera: Lycaenidae). Butterflies (Teinopalpus) 50: 19-23.

TENNENT, W.J., MULLER, C.J. and RAWLINS, A. 2010. A new Deudorix from Irian Jaya

(West Papua), Indonesia (Lepidoptera, Lycaenidae), with notes on Deudorix epirus Felder, 1860.

Australian Entomologist 37(2). 33-43.

VANE-WRIGHT, R.I. and DE JONG, R. 2003. The butterflies of Sulawesi: annotated checklist

for a critical island fauna. Zoologische Verhandelingen, Leiden 343: 3-267.

YAGISHITA, A. 2006. A new species of genus Deudorix (Lepidoptera, Lycaenidae) from

Morotai Island, Indonesia. Futao 52:18-19.

Australian Entomologist, 2015, 42 (2): 63-66 63

THE FIRST RECORDS OF ANTHENE SELTUTTUS SELTUTTUS

(ROBER, 1886) AND ANTHENE LYCAENOIDES SUTRANA

(FRUHSTORFER, 1916) (LEPIDOPTERA: LYCAENIDAE:

POLYOMMATINAE) FROM NORTHERN TORRES STRAIT,

QUEENSLAND, AUSTRALIA

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

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)

Abstract

Anthene seltuttus seltuttus (Rober, 1886) and Anthene lycaenoides sutrana (Fruhstorfer, 1916)

are recorded from northern Torres Strait, Queensland, Australia for the first time. Specimens of

both were collected on Dauan Island and a single female of Anthene seltuttus seltuttus was also

collected on neighbouring Saibai Island.

Introduction

The genus Anthene Doubleday, 1847 contains about 100 species occurring in

the Afrotropical Region and throughout the Indo-Australian Region (Parsons

1998). In Australia, it is represented by two species: Anthene seltuttus affinis

(Waterhouse & R.E. Turner, 1905) and A. lycaenoides godeffroyi (Semper,

[1879]). Anthene seltuttus affinis has been recorded from Moa Island, Torres

Strait and throughout the northeastern mainland as far south as Rockhampton

in Queensland, plus the Northern Territory and its neighbouring islands

(Braby 2000). Anthene lycaenoides godeffroyi has been recorded sporadically

from Moa Island to Cannonvale in Queensland and from northern Western

Australia and the Northern Territory (Braby 2000).

Discussion

Adult male and female specimens of a dark Anthene species were collected

by the authors during field trips to Dauan Island in 2001, 2002, 2011 and

2015 (Figs 1-4). A single female was also collected on Saibai Island in 2002.

The specimens are darker and larger than any mainland Australian specimens

of A. seltuttus affinis held by the authors (Figs 5-8). During the 2002 trip to

Dauan Island, a single male A. lycaenoides (C. Felder, 1860) (Figs 9-10) was

also collected by one of us (CEM) and it also appeared larger and more

conspicuously marked underneath than any Australian specimens contained

in the authors’ collections. This led to a review of the literature and a

comparison of Anthene specimens held in the Brandt Collection in the

Australian National Insect Collection, Canberra, in order to determine the

status of the specimens collected.

Tite (1966) recognised five subspecies of A. se/tuttus (Röber, 1886), two of

which occur in Papua New Guinea, with A. se/tuttus seltuttus occurring on

Daru Island, approximately 50 km to the east of Saibai Island (Parsons 1998).

Tite (1966) also recognised four subspecies of A. /ycaenoides, only one of

which, A. /. sutrana (Fruhstorfer, 1916), occurs in Papua New Guinea

64 Australian Entomologist, 2015, 42 (2)

(Parsons 1998). Anthene seltuttus seltuttus and A. lycaenoides sutrana have

not previously been recorded from within the Australian faunal region.

Abbreviations: ANIC — Australian National Insect Collection, Canberra;

CEMC — C.E. Meyer Collection, Brisbane; RWPC — R.P. Weir Collection,

Darwin; SSBC — S.S. Brown Collection, Bowral; PNG — Papua New Guinea.

Anthene seltuttus seltuttus (Röber, 1886) (Figs 1-4)

Material examined: QUEENSLAND (TORRES STRAIT): 4 do, 5 99, Dauan

Island, 26.iv.2002, 1 4, Dauan Island, 17.iv.2015, S.S. Brown (all SSBC);); 6 3, 15

2°, Dauan Island, 11-17.iv.2001, C.E. Meyer & S.S. Brown; 3 63, 5 99, Dauan

Island, 26.1v-2.v.2002, C.E. Meyer, S.S. Brown & R.P. Weir; 1 2, Dauan Island, 7-

14.1.2011, C.E. Meyer, S.S. Brown & A.I. Knight; 1 9, Saibai Island, 19-20.iv.2001,

C.E. Meyer & S.S. Brown (all CEMC); 1 3, 1 2, Dauan Island, 17-20.iv.2015, C.E.

Meyer (both RPWC)..

The authors initially suspected that these specimens were Anthene paraffinis

emoloides Tite, 1996 but, upon examination of the Brandt specimens of this

species and specimens of Anthene seltuttus seltuttus in that collection (in

ANIC), this was clearly not the case. The black spots on the margin of the

hindwing upperside in A. s. se/tuttus are larger, darker and confluent with the

dark marginal band, whereas in A. paraffinis emoloides these spots are

smaller, rounded and separated from the marginal band. Anthene paraffinis

emoloides is a paler, more translucent purple in colour and less robust than A.

s. Seltuttus. Genitalia dissections were carried out on male specimens of A. s.

affinis from Queensland, A. s. se/tutus from Dauan Island and A. paraffinis

emoloides from Woodlark Island (PNG) and, as stated by Tite (1966), the

genitalia were identical and provided no means of separating these species.

As Daru Island (PNG) is only some 50 km from Saibai Island and both are

near the PNG coast, it is reasonable to expect that this subspecies would

occur on the mainland between the two islands and, therefore, it is not

surprising that it also occurs on Saibai and Dauan Islands.

Anthene lycaendoides sutrana (Fruhstorfer, 1916) (Figs 9-10)

Material examined: QUEENSLAND (TORRES STRAIT): 1 ĝ, Dauan Island,

29.iv.2002, C.E. Meyer, S.S. Brown & R.P. Weir (CEMC).

A review of Parsons (1998) led the authors to believe that the A. lycaenoides

specimen (Figs 9-10) collected on Dauan Island might be of subspecies A. /.

sutrana, since it was larger than any A. l. godeffroyi specimens held in our

collections and the underside ground colour is a darker brownish grey,

making the white markings more conspicuous. Examination of A. I. sutrana

specimens held in the Brandt Collection (in ANIC) confirmed that the Dauan

specimen was most closely aligned with that subspecies. It also closely

resembles the phenotype of A. I. sutrana illustrated by Parsons (1998, pl. 64.

1759-1760) and it is therefore tentatively placed with this subspecies.

Australian Entomologist, 2015, 42 (2) 65

Ne

Figs 1-8. Anthene seltuttus subspp. Upper and undersides of adults: (1-2) Anthene

seltuttus seltuttus, male; (3-4) Anthene seltuttus seltuttus female; (5-6) Anthene

seltuttus affinis male; (7-8) Anthene seltuttus affinis female.

66 Australian Entomologist, 2015, 42 (2)

i A r }

b 1a ¥ =

Figs. 9-12. Anthene lycaenoides subspp. Upper and undersides of adults: (9-10)

Anthene lycaenoides sutrana, male from Dauan Island, Torres Strait. Forewing length

/ wingspan — 16 mm / 26 mm; (11-12) Anthene lycaenoides godeffroyi, male from

Green Hill, Thursday Island, Torres Strait.

Acknowledgements

The authors thank the respective Chairpersons of Dauan and Saibai Islands

for permission to collect there, Mr Ted Edwards for access to specimens in

the ANIC, Canberra, the genitalia dissections and for comments on the draft

manuscript, and Drs Michael Braby, Grant Miller and the late Steve Johnson

for assistance with the identification of these specimens.

References

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

CSIRO Publishing, Melbourne; xxvii + 976 pp.

COMMON, IF.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Angus and

Robertson, Sydney; xiv + 682 pp.

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

Academic Press, London; xvi + 736 pp.

TITE, G.E. 1966. A revision of the genus Anthene from the Oriental Region (Lepidoptera:

Lycaenidae). Bulletin of the British Museum of Natural History (Entomology) 18: 255-275.

Australian Entomologist, 2015, 42 (2): 67-72 67

CETHOSIA CYDIPPE DAMASIPPE C. & R. FELDER, 1867

(LEPIDOPTERA: NYMPHALIDAE: HELICONIINAE) IN TORRES

STRAIT, QUEENSLAND, AUSTRALIA

TREVOR A. LAMBKIN! and STEPHEN J. JOHNSON?’

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

(Email: trevor.lambkin@uqconnect.edu.au)

?Vale 17 November 2014

Abstract

Cethosia cydippe damasippe C. & R. Felder, 1867 has previously been recorded only from New

Guinea, while C. c. chrysippe (Fabricius 1775) is endemic to Queensland, Australia, the two

subspecies being separated by Torres Strait. We report two specimens of C. c. damasippe

collected from two of the three northernmost islands of Torres Strait that are within 10 km of the

southern Papua New Guinea coastline, viz. Saibai and Dauan. Although both specimens are

possibly vagrants from Papua New Guinea, it is suggested that they might have originated from a

resident population on Dauan Island, which has extensive areas of tropical semi-deciduous

monsoon forest growing among granite boulders, a likely habitat for the as yet unrecorded larval

host plant Adenia heterophylla (Passifloraceae). This record is thus the first for C. c. damasippe

in Australian waters.

Introduction

The genus Cethosia Fabricius, 1807 is a widespread group of tropical

nymphalid butterflies commonly known as lacewings. The genus consists of

15 species (Küppers 2006) and occurs throughout the Indo-Australian

Region, ranging from India and China, throughout Southeast Asia and its

associated islands, to New Guinea and Australia (Woodhouse 1949, Tsukada

1985, Parsons 1998, Bascombe et al. 1999, Braby 2000, Küppers 2006).

Cethosia cydippe (Linnaeus, 1767) is an Australian Region species occurring

in the Moluccas, Tanimbar, Kai, Aru, Waigeo, Misool, Biak, Yapen, New

Guinea, Woodlark, Goodenough, D’Entrecasteaux, Trobriand, Rossell and

Misima islands and in northern Queensland, Australia (Parsons 1998,

Kuppers 2006). Across this region, 15 subspecies are presently recognised

(Küppers 2006), although Parsons (1998) considered a population from

Misima Island to represent an additional, undescribed subspecies. Corbet

(1949) believed that the holotype of C. cydippe originated from Ambon

Island in the Southern Moluccas of eastern Indonesia but Parsons (1998) later

expressed some doubt surrounding the type locality.

In Australia, C. c. chrysippe (Fabricius 1775) is endemic to Queensland,

while C. penthesilea paksha Fruhstorfer, 1905 occurs in the Northern

Territory (Braby 2000). The latter taxon is also known from Timor, Wetar,

Western Daya Islands and Tanimbar in Indonesia (Küppers 2006).

In Queensland, C. c. chrysippe is considered common in lowland rainforest,

including riparian forest (Braby 2000). It occurs along the east coast from

Cape York to the Rocky River on Cape York Peninsula, and from Mt Webb,

68 Australian Entomologist, 2015, 42 (2)

south-west of Cooktown, south to Ollera Creek, north-west of Townsville

(Braby 2000).

Directly to the north of Australia, C. c. damasippe C. & R. Felder, 1867

occurs on mainland New Guinea, including its close islands of Daru, Yule

(Parsons 1998) and Yapen (Jobi) (Küppers 2006), which are all within 14 km

of the New Guinea mainland. This subspecies is separated from C. c.

chrysippe in Queensland by the expanse of Torres Strait, a body of tropical

water approximately 150 km wide containing around 150 islands distributed

across the Strait (Stanton et al. 2008). Within New Guinea, C. c. damasippe

is widespread in the Western Province of Papua New Guinea (Parsons 1998),

where it is recorded from Waidoro and Daru Island (Parsons 1998). These

two locations are not a great distance (42 and 79 km respectively) from

Dauan Island, which is one of the three northern islands of the Torres Strait

(viz. Boigu, Saibai and Dauan) that are no more than 10 km from the southern

Papua New Guinea coastline. Parsons (1998) indicated that, in Papua New

Guinea, C. c. damasippe can be locally common and occurs in rainforest and

marginal secondary vegetation from sea level to 2300 m.

Considering the proximity of the northern islands of Torres Strait to New

Guinea, it is not surprising that two specimens of C. c. damasippe have been

collected recently from these northern islands. These collections represent a

new butterfly record for Australia and Torres Strait. Here we document and

illustrate these specimens, provide field observations and comments for both,

and discuss the taxon’s diagnostic facies and its possible distribution in

Torres Strait.

The following abbreviations refer to repositories from which material has

been examined: ANIC — Australian National Insect Collection, Canberra;

GRFC — GR. Forbes collection, Brisbane; MTQ — Museum of Tropical

Queensland, Townsville; QM — Queensland Museum, Brisbane; TLIKC —

Joint collection of T.A. Lambkin and A.I. Knight, Brisbane.

Abbreviations of collectors’ names are: AIK — A.I. Knight; EJLH — E.J.L.

Hallstrom; GBM — G.B. Monteith; GRF — G.R. Forbes; HR — H. Rauber; IRJ

— I.R. Johnson; LR — L. Radunz; SJJ — S.J. Johnson; TAL — T.A. Lambkin;

TML — T.M. Lambkin; WWB — W.W. Brandt.

Figs 1-8. Cethosia cydippe (all figures not to scale, upperside left, underside right):

(1-6) C. d. damasippe ő (1, 3): (1) Saibai Is., Torres Strait, Qld, 3.v.2000, AIK,

[forewing length 48 mm], (3) Bulolo, PNG, 4.11.1970, LR [45 mm]; 99 (2, 4-6): (2)

Dauan Is., Torres Strait, 22—27.11.2014, SJJ&IRJ [45 mm], (4) Taraka, Lae, PNG,

12.x1.1973, GRF [40 mm], (5) Wau, Morobe Province, PNG, 1.v.1979, GRF [45 mm],

(6) Finschhafen, Morobe Province, 22.1v.1973, GRF [40 mm]; (7-8) C. d. chrysippe:

(7) & Flying Fish Point, via Innisfail, NQ, 16.iv.1987, TAL [45 mm], (8) 9 Flying

Fish Point, via Innisfail, 21.1v.1985, TAL [40 mm].

Australian Entomologist, 2015, 42 (2) 69

70 Australian Entomologist, 2015, 42 (2)

Material examined

Cethosia cydippe damasippe (Figs 1-6)

QUEENSLAND: 1, Saibai Island, Torres Strait, 3.v.2000, AIK (TLIKC); 19,

Dauan Island, Torres Strait, 09°25’S 142°32’E, 22—27.11.2014, SJJ&IRJ (MTQ).

PAPUA NEW GUINEA: 16, Asuar, Madang, 27.viii.1971, GRF (GRFC); 14, Brown

River, near Port Moresby, Central Province, 28.xii.1967, HR (QM); 299, same data

except, 28.1.1968, 13.iv.1968; 14, same data except 19.x.1976, GRF (GRFC); 14,

Bulolo, 4.11.1970, LR (QM); 19, Finschhafen, Morobe Province, PNG, 22.iv.1973,

GRF (GRFC); 28, Kassam Pass, Morobe Province, 16.viii.1977, GRF (GRFC);

243, Kodama Range (Mt Kaindi), 4500’, New Guinea, 22.i1.1952, WWB & EJLH

(ANIC); 14, Kokoda Trail, 2.xii.1968, HR (QM); 388, 299, Kiunga, Fly River,

New Guinea, 2.vii.-31.x.1957, WWB (ANIC); 8433, 229, Lae, New Guinea,

10.vi.1951 (14), 30.vi.1951 (13), 6.vii.1951 (14), 17.vii.1951 (14), 26.vii.1951

(18), 7.x.1951 (13), 12.x.1951 (13), 16.x.1951 (13), 25.viii.1951 (12), 16.x.1951

(12), WWB & EJLH (ANIC); 1, Madang, 1.xi.1971, GRF (GRFC); 19, Sogeri,

Central Province, 8.ix.1974, HR (QM); 788, 19, Subitana (Central District), 1800’,

New Guinea, 20.i1.1949 (13), 27.vi.1949 (18), L.viii.1949 (13), 27.vii.1949 (13), -

x.1949 (13), 14.x.1949 (13), 4.xii.1949 (18, 19), WWB & EJLH (ANIC); 16,

Tapini, Loloipa River Bridge Camp, 2200’, New Guinea, 15.xi.1957-15.11.1958,

WWB (ANIC); 19, Taraka, Lae, 12.xi.1973, GRF (GRFC); 14, Wau, Morobe

Province, 17.11.1967, GBM (QM); 19, same data except 1.v.1979, GRF (GRFC).

INDONESIA: 16, Tot Village, West Irian, 3.xi.1961 (GRFC).

Cethosia cydippe chrysippe (Figs 7-8)

QUEENSLAND: 204, 329, Flying Fish Point, via Innisfail, NQ, 3.v.1978,

TAL&TML (13), 16.iv.1987, TAL (12), 21.iv.1985, TAL (399) (TLIKC); 13, 19,

Coquette Point, Innisfail, NQ, 20.iv.1987, TAL (TLIKC); 18, 229, Jarra Creek, W

of Tully, NQId, -.iv.1978, TAL&TML (TLIKC).

Field observations and comments

In May 2000, a male C. cydippe (Fig. 1) was collected (by AIK) on the

western end of Saibai Island (9°22'46"S, 142°37'37"E) as it flew out of

mangroves. Saibai is predominantly a mud island and its vegetation consists

of primarily halophytic plant species, grassland, Pandanus and mangrove

species. The occurrence of C. cydippe on Saibai was therefore enigmatic as

the species is principally a forest dweller and thus unlikely to have originated

on the island. At that time it was presumed to be a vagrant from either the

Papua New Guinea mainland (just 5 km north), where only limited forest

occurs (predominantly at Mabaduan, 9°16'35"S, 142°44'15"E, 17 km NE of

Saibai), or from nearby Dauan Island (9°24'54"S, 142°32'22"E), which is

approximately 7 km to the east.

In February 2014, a pale-morph female of C. cydippe (Fig. 2) was collected

by one of us (SJJ) on the margin of monsoon forest at the summit of Mt

Cornwallis, Dauan Island. On Dauan, the most dominant feature of the island

is semi-deciduous monsoon forest, which grows extensively amongst hills

composed of piles of exposed granite boulders, with the highest part of this

Australian Entomologist, 2015, 42 (2) 71

boulder stack (275 m) being Mt Cornwallis. Travel to the summit of Mt

Cornwallis is possible but almost exclusively by helicopter.

Examination of the Saibai and Dauan Island specimens (Figs 1-2) showed

that their external facies, and most obviously that of the female, were

different from C. c. chrysippe from Australia and fitted the New Guinean

subspecies, C. c. damasippe (Figs 3-6).

Discussion

The male of C. c. damasippe (Figs 1, 3) differs from that of C. c. chrysippe

(Fig. 7) by its overall darker appearance on both the upper and undersides.

On the upperside, the black margins are much more extensive in C. c.

damasippe than in C. c. chrysippe. On the underside, C. c. damasippe is,

again, more extensively black than C. c. chrysippe and, on the hind wing, the

series of bright black basal bars edged with grey that is typical of C. c.

chrysippe is greatly reduced or absent. The upperside wing bases of females

of C. c. damasippe are variable in colour (Figs 2, 4-6) but are mostly a much

duller red than those of C. c. chrysippe. Rarely, the wing bases in females of

C. c. damasippe are not red but are pale orange (as in the female from Dauan)

to pale yellow or even white, or greenish to greenish-grey (Parsons 1998).

Although Parsons (1998) believed that the unusually coloured females of C.

c. damasippe were predominantly confined to the southern lowlands of New

Guinea, he did comment that the pale form had been recorded from the

northern coastal regions of Papua New Guinea. Certainly the female collected

on Dauan is geographically very close to the southern lowlands of Papua

New Guinea.

Based on the collection of the female of C. c. damasippe from the top of Mt

Cornwallis, it is possible that the male collected on Saibai, and the above

female, originated from a resident population on nearby Dauan. That premise

is supported by the recent collection on Dauan of six specimens of Vindula

arsinoe (Cramer, [1777]), a species that shares at least one larval host plant

with C. cydippe in Papua New Guinea and Australia, viz. Adenia heterophylla

(Blume) Koord. (Passifloraceae) (Parsons 1998, Braby 2000). Within the

region, A. heterophylla is recorded from Koey Ngurtai Island, Torres Strait

(just north of Badu Island) and the Oriomo River near Daru in Papua New

Guinea (Australia’s Virtual Herbarium 2015). However, the Torres Strait

Regional Authority (2013) did not list A. heterophylla in its plant listings

from Dauan Island and thus it also remains a possibility that the Saibai and

Dauan specimens (and perhaps also V. arsinoe) are vagrants from Papua New

Guinea.

If the infrequent sighting of this species on Dauan reflects the host plant’s

restricted distribution on the island, where it might only occur in more

upland, remote and inaccessible parts, rather than vagrancy from Papua New

Guinea, then C. c. damasippe might also be found on islands further south in

Torres Strait. Some of these islands contain tracts of similar forest (Regional

72 Australian Entomologist, 2015, 42 (2)

Ecosystems 2014) but, due to the environments where C. c. damasippe might

fly, it has thus far not been detected. In particular, C. c. damasippe might

occur on Gabba Island (approximately 37 km SE of Dauan), which has some

areas of granite boulders and semi-deciduous monsoon forest similar to

Dauan but is uninhabited and as yet not visited by lepidopterists.

Acknowledgements

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

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

assistance during time spent on their islands. Appreciation is given to C.J.

Burwell (QM), E.D. Edwards (ANIC), G.R. Forbes (GFC) and N. Bruce

(MTQ) for allowing access to specimens in their care. G. Thompson provided

the high quality photograph of the Dauan specimen of C. c. damasippe. This

work forms a partial tribute to the late Stephen J. Johnson for his significant

contribution to the butterfly knowledge of Torres Strait. In addition, this

paper partially fulfils the requirements for a Master of Philosophy degree

undertaken by the senior author at the University of Queensland, Brisbane.

References

AUSTRALIA’S VIRTUAL HERBARIUM. 2015. [Accessed 20 January 2015]. Available from

URL: Attp://avh.ala.org.au/occurrences/search?taxa=adenia-+ heterophylla#tab_mapView

BASCOMBE, M.J., JOHNSTON, G. and BASCOMBE, F.S. 1999. The butterflies of Hong

Kong. Academic Press, San Diego, CA; xiii + 422 pp.

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

CSIRO publishing, Collingwood; xx + 976 pp.

CORBET, A.S. 1949. The Linnaean names of Indo-Australian Rhopalocera. Part 7. Summary of

determinations. Proceedings of the Royal Entomological Society of London, Series B Taxonomy

18: 191-200.

KUPPERS, P.V. 2006. Cethosia. In: Bauer E. and Frankenbach, T. (eds.), Butterflies of the

World. Vol. XI, Nymphalidae, part 24). Goecke and Evers, Keltern. 6 pp + 34 plates.

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

Academic Press, London; xvi + 736 pp, xxvi + 136 pls.

REGIONAL ECOSYSTEMS. 2014. [Accessed 20 January 2015]. Available from URL:

http://environment. ehp.qld. gov.au/regional-ecosystems/details

STANTON, D.J., FELL, D.G. and GOODING, D.O. 2008. Vegetation communities and regional

ecosystems of the Torres Strait islands, Queensland, Australia. Final report to Torres Strait

Regional Authority Land & Sea Management Unit. 3D Environmental®; v + 235 pp.

TORRES STRAIT REGIONAL AUTHORITY. 2013. Profile for management of the habitats

and related ecological and cultural resource values of Dauan Island, January 2013. Prepared by

3D Environmental® for Torres Strait Regional Authority Land & Sea Management Unit; vi + pp

7-123.

TSUKADA, E. 1985. Nymphalidae (I). In: Butterflies of the South East Asian Islands, Vol IV.

Plapac Co, Ltd, Tokyo; 558 pp.

WOODHOUSE, L.G.O. 1950. The butterfly fauna of Ceylon. The Colombo’s Apothecaries’ Co.,

Ltd., Colombo; xxxii + 231 pp + lv pls.

Australian Entomologist, 2015, 42 (2): 73-76 73

REDISCOVERY OF THE RARE COCCINELLID MICRASPIS

FLAVOVITTATA (CROTCH, 1874) IN WESTERN VICTORIA

(COLEOPTERA: COCCINELLIDAE)

REINER RICHTER

PO Box 37, Monbulk, Vic 3793

Abstract

Micraspis flavovittata (Crotch, 1874), a distinctive yellow and black Australian ladybird that has

not been collected for more than 60 years, is reported in numbers from western Victoria. Notes

are provided on pollen-feeding and other aspects of its biology.

Introduction

The distinctive, yellow and black coccinellid beetle Micraspis flavovittata

(Crotch, 1874) was described 141 years ago from one specimen from

‘Melbourne’ in the Natural History Museum, London (BMNH). Since then

only three specimens have been recorded, all collected by the early Victorian

coleopterist F. E. Wilson. Two are from Narbethong in 1949 (in Museum of

Victoria (MV)) and one from Kallista in 1944 (specimen missing); both

localities are a little to the north-east of Melbourne city (Pope 1989, Atlas of

Living Australia 2015, Ken Walker pers. comm.). It is not represented in the

Australian National Insect Collection (ANIC) and is not illustrated in a recent

monograph of the family (Slipinski 2007). The original type is not dated but

was Clearly collected before 1853, which is when its accession was registered

by the Natural History Museum (Pope 1989).

The author visited the Discovery Bay Coastal Park (Fig. 1) in western

Victoria in 2008 and discovered a thriving colony of the endangered Ancient

Greenling damselfly, Hemiphlebia mirabilis Selys, 1869, in Long Swamp

(Richter 2009). Other Odonata rare in Victoria were also found at Discovery

Bay (e.g. Austroagrion cyane (Selys, 1876) and Austrothemis nigrescens

(Martin, 1901)), many being species previously known only much further

east. Subsequent visits have been made once or twice each season since 2008.

In October 2014, I visited Mcfarlanes Swamp within the Discovery Bay CP

(an area only visited once before, in February 2009 when the ground was

very dry). A yellow and black coccinellid was found to be common on

vegetation in standing water (mostly water ribbons, Triglochin cf. procerum

R.Br.: Juncaginaceae) and was photographed (Fig. 2). This was subsequently

identified by Dr Roger Booth, a coccinellid taxonomist at the BMNH, as the

long-lost Micraspis flavovittata. More adults, as well as larvae, were

observed on later visits as detailed below.

There are about 30 species in the Old World genus Micraspis Chevrolat, six

of which occur in Australia (Pope 1989, Slipinski 2007). Micraspis frenata

(Erichson, 1842) is common and widespread and is the only Australian

species that has been studied in detail. Although members of the genus are

considered to be basically aphid feeders, A. frenata adults have been shown

74 Australian Entomologist, 2015, 42 (2)

to feed on nectar (Hawkeswood 1994) and on grass pollen (Hawkeswood and

Turner 2002). Anderson and Hales (1983) observed adult and larval

development of the same species on both aphids and pollen in the field and

were able to rear it in the laboratory on powdered honeybee brood. Thus

there is a tendency to polyphagy in the genus.

Collection and behavioural observations

5 October, 2014. About 40 individual adults were seen at Mcfarlanes Swamp

(38.0720°S, 141.0657°E). Almost all were on the broad leaves of the water

ribbon plant, which was flowering profusely at the time. Larvae were neither

searched for nor noticed.

29-30 November, 2014. Only a few adults were seen at Mcfarlanes Swamp

on the first day but about 20 larvae were seen in the same situation and were

assumed to be M. flavovittata. The following day at Long Swamp

(38.1034°S, 141.1054°E), two adults and about eight larvae were seen in the

same habitat as at Mcfarlanes Swamp.

4-5 April, 2015. Mcfarlanes Swamp was visited in the late afternoon of April

4 but no beetles were observed. On April 5 at Long Swamp no beetles were

active at the original site but a search further west (38.0927°S, 141.0928°E)

located three adults and one larva active and 20-30 were seen sheltering in

between the tightly packed leaves near the base of bull-rush plants (Typha

domingensis Pers.: Typhaceae). Voucher specimens were collected for

lodgement in the MV and ANIC.

Most adult beetles in October were seen on the long leaves of water ribbons.

They were usually stationary but when disturbed would run along the leaf or

hide on the underside. There was no obvious insect prey such as aphids or

scale insects present. It was not clear if they were feeding on material on the

leaves but there might have been scattered pollen present. One adult was

found in the pollen-covered head of another aquatic plant, where it appeared

to be feeding (Fig. 3). Several others appeared covered with some pollen,

indicating that they most likely at least supplement their diet with pollen.

On the return visit in November, larvae were more closely observed. They

were seen on the water ribbons as well as a spike rush (Baumea arthrophylla

Nees (Boeck.): Cyperaceae) that is common in the swamp. Here they

appeared to be feeding on sections of the stem where the rush had been

damaged and was oozing liquid (Fig. 4). One larva that crawled on to my

hand during photography bit the skin actively without puncturing it (Fig. 5).

On the April visit the sky was mostly overcast and the temperature scarcely

above 16°C. Beetle activity was low and many were observed sheltering in

the bull-rush (possibly where they might hibernate over winter). There were

from 1 to 3 adults on some bull-rush plants, particularly older plants with

dead outer leaves but not obviously in contact aggregations as defined by

Hales et al. (1986). No prey was obvious and little pollen was available.

Australian Entomologist, 2015, 42 (2) 75

Figs 1-5. (1), Mcfarlanes Swamp in the Discovery Bay Coastal Park, eastern Victoria.

(2-5), Micraspis flavovittata: (2) adult on leaf of Triglochin cf. procerum, (3) adult

apparently feeding on pollen on flower-spike of an unknown plant; (4) larva

apparently feeding on exudate from injury of a rush stem; (5) larva biting surface of

human skin.

76 Australian Entomologist, 2015, 42 (2)

Discussion

The new records are about 340 km west of the original type locality of

Melbourne and almost 400 km west of the only records within a century from

Narbethong and Kallista. Thus this is a substantial range extension. The

abundance of the species in the coastal swamps at Discovery Bay might

indicate that the widespread draining of lowland swamps around Melbourne

has resulted in the beetle’s disappearance from areas where it once occurred.

The apparent feeding of M. flavovittata on pollen and plant exudates accords

with similar observations on the related M. frenata (see Introduction) and

might indicate that this habit is more widespread in members of Micraspis.

Hales et al. (1986) reported Coccinella transversalis Fab., 1781 feeding on

dead fish exudates. Although numerous adults were seen, no mating was

observed during any of the visits.

Acknowledgements

I am grateful to Ken Walker (Museum of Victoria) for information on the

MV collection, to Roger Booth (Natural History Museum, London) for

identification of the species, Di Crowther (Department of Environment,

Land, Water & Planning) and Bryan Haywood (Nature Glenelg Trust) for

help with identification of flora, and to Geoff Monteith (Queensland

Museum) for help in drafting the manuscript.

References

ANDERSON, J.M.E. and HALES, D.F. 1983. Micraspis lineata (Thunberg) (Coleoptera:

Coccinellidae - seasonality and food. General and Applied Entomology 15: 47-52.

ATLAS OF LIVING AUSTRALIA. 2015. [Accessed March 2015]. Available at:

http://bie.ala.org.aw/species/Micraspis+ flavovittata

CROTCH, G.R. 1874. A revision of the Coleopterous Family Coccinellidae. E.W. Janson,

London; xv + 311 pp.

HALES, D.F., ANDERSON, J.M.E. and van BRUNSCHOT, K.A. 1986. Aggregation in

Australian ladybirds. Pp 205-210, in :Hodek I. (ed.), Ecology of Aphidophaga. Academia, Prague

& Dr W Junk, Dordrecht.

HAWKESWOOD, T.J. 1994. Notes on the Australian ladybird beetle Micraspis frenata

(Erichson) (Coleoptera: Coccinellidae) feeding on nectar from Asclepias and Gomphocarpus

flowers (Asclepiadaceae). Giornale Italiano di Entomologia 7: 67-71.

HAWKESWOOD, T.J. and TURNER, J.R. 2002. Observations on the adults of Micraspis

jrenata (Erichson, 1842) (Coleoptera: Coccinellidae) feeding on the pollen of native and non-

native grasses (Poaceae) in eastern New South Wales. Journal of Entomological Research

Society 4: 21-29.

POPE, R.D. 1989. Revision of the Australian Coccinellidae (Coleoptera) Part 1. Subfamily

Coccinellinae. Invertebrate Taxonomy 2: 633-735.

RICHTER, R. 2009. Discovery of a new population of Hemiphlebia mirabilis (Ancient

Greenling). Victorian Entomologist 39: 27-29.

SLIPINSKI, A. 2007. Australian ladybird beetles (Coleoptera: Coccinellidae): their biology and

classification. Australian Biological Resources Study, Canberra; 286 pp.

Australian Entomologist, 2015, 42 (2): 77-90 77

A NEW SPECIES OF TOXALA MOULDS (CICADIDAE:

CICADETTINAE: CICADETTINI) FROM NORTH QUEENSLAND,

WITH NOTES ON ITS CALLING SONG AND COMPARISONS

WITH A CONGENERIC SPECIES

L.W. POPPLE

Biodiversity Assessment and Management, PO Box 1376, Cleveland, Old 4163

(Email: lindsay.popple@uq.net.au)

Abstract

Toxala Moulds was previously a monotypic genus of small grass cicadas known from central

eastern Australia. A new species is described from the Herberton district in north Queensland,

Australia, extending the distribution of this genus. The new species has been found in open

grassy woodland with sparse heath elements, the adults of which are associated with grass and

have been collected during late December to early January. The single known population appears

to be highly localised and is potentially prone to habitat disturbance. Additional distribution

records are provided for Toxala verna (Distant) from southeastern Queensland. First

documentation of the calling songs of the two Toxala species is also provided, along with

comparative notes.

Introduction

The grasslands of Australia are occupied by a diverse array of small cicadas

from approximately 17 different genera. These have been the subject of

several recent taxonomic studies (e.g. Ewart 2005, Ewart and Marques 2008,

Ewart 2013). Many of the species appear to be grassland specialists, with

adults emerging in large numbers after rainfall and, in most cases, normally

persisting for only two weeks. The ephemeral nature of their appearance has

generally resulted in infrequent opportunities for collection and observation.

As a result, much of their natural history and species diversity remains poorly

understood.

The genus Toxala was introduced by Moulds (2012) as part of a substantial

review of Australian cicada genera. Urabunana verna Distant was included

as the type species of this new genus. Here, I describe a second (new) species

of Toxala from north Queensland and provide additional distribution records

for T. verna. A first description of the calling songs of both species is also

given, along with some discussion on similarities and differences between the

two species.

Methods

Anatomical terminology follows Moulds (2005, 2012) for body structures

and wing characters, Dugdale (1972) and Moulds (2005) for genitalia, de

Boer (1999) for opercula, and Simmons and Young (1978), Dugdale (1972)

and Bennet-Clark (1977) for timbals. The long timbal ribs are referred to

sequentially as long ribs numbered 1 to 5, with long rib 1 being the most

posterior (adjacent to timbal plate). The higher classification adopted in this

paper follows Moulds (2012). Measurements (in mm) are given as ranges and

means (in parentheses) and include all specimens available. Head width spans

78 Australian Entomologist, 2015, 42 (2)

across the outer margins of the compound eyes; pronotum width across the

extremities of the lateral margins (excluding ampliated lateral angles);

abdomen width is measured across the outer edges of the auditory capsules.

Collection Institution abbreviations

AE — private collection of A. Ewart, Caloundra; AM — Australian Museum,

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

private collection of David Emery, Sydney; LWP — private collection of L.W.

Popple, Brisbane; MSM — private collection of M.S. Moulds, Kuranda; QM —

Queensland Museum, Brisbane.

Calling song analysis techniques

The description of calling songs broadly follows Ewart and Marques (2008).

The term phrase describes the highest level repeated sequence in a calling

song. Within each phrase, a number of finer structures are present. These are

described in the following text, in order of increasing duration. A ‘pulse’ is

defined as a single complete movement of the timbals. The term ‘syllable’

applies to the smallest grouping of pulses (typically 5-10 ms duration).

Where multiple syllables occur in succession, without coalescence (i.e. where

the syllables are spaced by short periods of silence), this is termed a ‘syllable

train’. Where 2-9 syllables coalesce, these become a ‘macrosyllable’. Longer

durations of continuous sound (>10 syllables) are known as an ‘echeme’. All

of these features have been identified in the calling songs of Toxala cicadas.

Field recordings have been used preferentially for song analysis. All

recordings were obtained from a distance of at least 20 cm from the calling

cicada to reduce the chances of near-field effects. None of the recordings

displayed signs of amplitude clipping due to microphone overload.

I obtained audio recordings using a Sony MZR700 minidisc recorder with

Sony ECM-MS957 Electret Condenser microphone (frequency response up

to 18 kHz). Container recordings of T. verna by A. Ewart were also examined

for the purposes of this study. These were made using a Sony WM-D6C

Walkman with Sennheiser K6/ME66 microphone (frequency response up to

18 kHz).

Processing and analysis of recordings was undertaken with Cool Edit Pro

(Version 2.1) software. Amplitude spectra were analysed using a linear

frequency axis on a 1024-point Fast Fourier Transform with a Hamming

window function.

Systematics

Genus Toxala Moulds

Diagnosis follows Moulds (2012), except for the following amendment to

accommodate the new species: Hind wing with 3-5 apical cells.

Australian Entomologist, 2015, 42 (2) 79

Toxala mckinnonae sp. n.

(Figures 1A-B, 2-3)

Type material. Holotype &, QM reg. no. T228767, AUSTRALIA (QUEENSLAND):

‘Hornet Hill’, Herberton district, 3.1.2007, L. Popple, M. & A McKinnon, 17°23’22”S

145°21'02”E, 328-0001 (QM). Paratypes: 1 4, same location as holotype,

29.xii.2007, L. Popple, A. McKinnon, 328-0015 (AE); 1 ĝ, same location as

holotype, 29.x1i.2007, L. Popple, A. McKinnon, 328-0014; 1 92, same data as

holotype, 328-0008 (both AM); 1 ĝ, same location as holotype, 29.xii.2007, L.

Popple, A. McKinnon, 328-0016 (DE); 5 ¢, 1 9, same data as holotype, 428-0002

to 428-0006, 428-0009; 4 3", same location as holotype, 29.xii.2007, L. Popple, A.

McKinnon, 328-0010 to 328-0013 (all LWP); 1 ĝ, same location as holotype,

29.x11.2007, L. Popple, A. McKinnon, 328-0017 (MSM); 1 2, same data as holotype,

328-0007, QM reg. no. T228768 (QM).

Description

Male (Figs 1A, 2)

Head. Supra-antennal plate variable, often pale brown anteriorly and dark

brown posteriorly, or predominantly dark ochraceous; vertex and frons black;

mandibular plates black with conspicuous silver pubescence; genae black,

sometimes with light brown colouration adjacent to edges of postclypeus,

covered by silver pubescence; small, dull pale brown median orange-brown

triangular fascia, extending and widening posteriorly from near median

ocellus to pronotal margin along the epicranial suture; ocelli red; compound

eyes brown to dark brown. Postclypeus predominantly black, pale brown in

anterior median area and along margins and between the transverse ridges,

sometimes grading from black through ochraceous to pale brown;

anteclypeus mainly black, with silver pubescence; rostrum brown, dark

brown at apex; antennae dark brown to black.

Thorax. Pronotum mainly brown to dark brown, with a brown to dull pale

brown medial fascia, bordered with dark brown to black colouration that

widens anteriorly of pronotal collar and also towards proximal margin; dorsal

and lateral fissures inconspicuous, brown to dark brown; pronotal collar

brown to dull pale brown, with lateral margins ampliate and tending dark

brown. Mesonotum with submedial sigilla dark brown to black, fused

anteriorly, with rounded posterior terminations; lateral sigilla black,

prominent, elongated and narrowing posteriorly; dorsolateral fasciae brown

to dark brown and narrow along parapsidal suture, broadening conspicuously

to anterior of scutal depressions, fused or black in intervening medial area;

remainder of mesonotum, including lateral edges adjacent to lateral sigilla

and area surrounding scutal depressions, brown, becoming pale brown at

margins; scutal depressions black; cruciform elevation brown to pale brown,

bordered with areas of silver pubescence. Metanotum brown anteriorly, pale

brown along posterior margin.

80 Australian Entomologist, 2015, 42 (2)

Fig. 1. (A-B) Toxala mckinnonae sp. n., “Hornet Hill’ west of Herberton, northern

Queensland (17°23’S 145°21'E), (A) male (body length 10.5 mm), (B) female (body

length 12.0 mm); (C-D) 7. verna (Distant), base of Blackbutt Range, southeastern

Queensland (26°53’S 152°13’E), (C) male (body length 11.8 mm), (B) female (body

length 12.1 mm).

Wings. (Fig. 2A). Forewing costal vein translucent, swollen, pale brown

throughout; other venation brown, darker proximally; pterostigma medium to

dark brown, semi-opaque; basal membrane grey. Hind wing venation brown,

with grey, partly opaque plaga around margins of anal cell 3, adjacent to vein

3A and vein 2A; five apical cells.

Legs. Coxae and trochanters predominantly pale brown with irregular dark

brown fasciae on anterior, lateral and posterior faces of fore and mid coxae;

fore, mid and hind femora pale brown with brown longitudinal fascia

developed on posterior, dorsal and anterior faces; fore tibiae pale brown to

brown; mid tibiae and hind tibiae pale brown; tarsi a mixture of pale brown

and brown; claws brown, darker apically; spines of fore femora dark brown.

Opercula. (Fig. 2B). Predominantly pale brown to grey; dark brown to black

anteriorly; with silver pubescence; meracanthus spike dark brown with paler

lateral margins, barely overlapping opercula.

Timbals. (Fig. 2C). Four long ribs; long ribs 1-4 fused dorsally to basal spur;

ribs 1-2 spanning across timbal membrane, fused ventrally; long ribs 3-4 both

shorter than posterior long ribs 1-2; long rib 3 apparently discontinuous, with

separate ventral section.

Australian Entomologist, 2015, 42 (2) 81

Fig. 2. Toxala mckinnonae sp. n. male from ‘Hornet Hill’ west of Herberton, northern

Queensland: (A) fore and hind wings; (B) left operculum; (C) left timbal (anterior

margin at top, dorsal margin at right); (D) pygofer lateral view; (E) pygofer ventral

view. Scale bars are 1 mm, except wings (2 mm).

Abdomen. Tergite 1 mainly black, brown to dark brown laterally; tergite 2

mainly black, grading through brown to pale brown on posterior dorsolateral

half; remaining tergites mainly black, with areas of brown colouration on

posterior dorsolateral surfaces and along lateral margins; silvery pubescence

most prominent along lateral edges of tergites. Sternite I dark brown to black;

sternite II dark brown medially, pale brown over remainder; sternites HI to

VII brown to pale brown, paler posteriorly, with subtle, brown medial areas;

sternite VIII dark brown; intersegmental membranes yellow-brown.

Genitalia. (Fig. 2D-E). Pygofer with black colouration dorsally, becoming

dark brown along beak and dark brown to brown along anterior lateral edges;

pale brown over the remainder; claspers conspicuously elongated, extending

82 Australian Entomologist, 2015, 42 (2)

posteriorly beyond termination of pygofer, bluntly rounded apically; uncus

vertically narrow, relatively short, with rounded, beak-like termination;

pseudoparameres slightly longer than endotheca and ventral support;

endotheca long and fleshy, extending almost as far as ventral support.

Female (Fig. 1B)

Similar to male in general colouring and patterning, although conspicuously

paler overall.

Head. Supra-antennal plate and vertex mottled brown to pale brown; frons

ochraceous with pale brown area extending medially from anterior of median

ocellus; mandibular plates and genae dark brown; small median pale brown

triangular fascia, extending and widening posteriorly from near median

ocellus to pronotal margin along the epicranial suture; ocelli pink to red;

compound eyes brown to dark brown. Postclypeus dark brown with median

area, margins and area between the transverse ridges pale brown; anteclypeus

dark brown to ochraceous; rostrum brown, darker posteriorly; antennae dark

brown.

Thorax. Pronotum mainly pale brown, with a brown to dull yellow-brown

medial fascia, bordered with brown colouration that widens and darkens

anteriorly of pronotal collar; dorsal and lateral fissures relatively

inconspicuous, brown; pronotal collar pale brown to yellow-brown, with

lateral margins ampliate and tending dark brown. Mesonotum with submedial

sigilla dark brown, unfused, with rounded posterior terminations; lateral

sigilla dark brown, prominent, elongated and narrowing posteriorly;

dorsolateral fasciae brown to pale brown and narrow along parapsidal suture,

broadening conspicuously to anterior of scutal depressions, fused in

intervening medial area; remainder of mesonotum, including lateral edges

adjacent to lateral sigilla and area surrounding scutal depressions, brown to

pale brown; scutal depressions dark brown; cruciform elevation pale brown,

bordered with areas of silver pubescence. Metanotum pale brown.

Legs. Coxae and trochanters predominantly pale brown to brown; fore, mid

and hind femora pale brown with brown longitudinal fascia developed on

anterior faces; fore tibiae mainly brown; mid and hind tibiae pale brown; tarsi

a mixture of pale brown and brown; claws brown, darker apically; spines of

fore femora dark brown.

Abdomen. Tergite 1 mottled pale brown; tergites 2—8 pale brown with lines of

dark brown to brown colouration extending from anterior median dorsal and

lateral surfaces; abdominal segment 9 pale brown, with a pair of brown

dorsolateral fasciae, extending from the anterior margin and not reaching the

posterior margin; stigma brown, inconspicuous. Sternites pale brown, with

subtle brown medial areas extending from anterior margin and fading

towards posterior; gonocoxite IX brown; intersegmental membranes yellow-

Australian Entomologist, 2015, 42 (2) 83

brown. Ovipositor sheath extends <1 mm beyond the posterior termination of

abdominal segment 9.

Measurements. N = 14 33,3 29. Ranges and means (in parentheses), mm;

body length: £8 9.7-10.7 (10.3); 2° 12.0-12.4 (12.2); forewing length: ££

10.6-11.8 (11.2); 22 11.9-12.4 (12.2); forewing width: 6S 4.0-4.4 (4.2):

02 4.0-4.3 (4.2); ratio of forewing length to width: ¢¢ 2.6-2.9 (2.7); 29

2.8-3.1 (2.9); head width: £ 2.4-2.8 (2.6); 29 2.7-2.9 (2.8); pronotum

width: ¢¢ 2.1-2.3 (2.2); 99 2.3-2.5 (2.4); abdomen width: Jo 2.5-2.7 (2.6);

OO 2.6-2.8 (2.7).

Etymology. Named after my wife, Ms Anne-Marie McKinnon, an observant

entomologist and photographer who was the first person to detect the

presence of this small, unique cicada.

Distinguishing characters. Toxala mckinnonae can be distinguished from the

related T. verna by having five apical cells in both hind wings (the latter

species has only three to four cells in each wing, or five in one wing if

aberrant). In addition, T. verna exhibits much more contrasting colouration

(black and orange-brown in males and black and greenish or straw brown in

females), whereas T. mckinnonae males are a subtler combination of brown

and dark brown to black and females are mainly pale brown with dark brown

markings.

Distribution, habitat and seasonality. This species is currently known only

from a single locality, known colloquially as ‘Hornet Hill’, west of the

township of Herberton in northeastern Queensland (Fig. 3). Adults occur in

grass on the edges of grassy woodland and areas of open grassland on sandy

loam soils with heath habitat elements (e.g. Melaleuca nodosa). The single

known population appears to be highly localised and as such may be

vulnerable to fire and other habitat altering practices. Specimens have been

collected at the height of summer between late December and early January

following storms just prior to the onset of the wet season. The appearance of

the species and/or its seasonality may be irregular, as adults were not found

during subsequent visits to the site by the author on 7 January 2008 and 26

December 2010.

Additional specimen records of Toxala verna Distant

The former distribution of T. verna and new distribution records are depicted

in Fig. 3. Additional specimen records are as follows: 1 9, Foothills of Bluff

Range, 13-14.xii.1971, H. Frauca (ANIC); 4 44, SEQ: 27°32’S 153°10’E,

J.C. Trotter Park, Burbank, 7.ii.1998, L.W. Popple; 1 ĝ, same data as

previous, 19.ii.1998; 2 44, 1 2, same data as previous, 24.1.1999: 1 4, same

data as previous, ii.2008, L. Popple and A. McKinnon; 3 ¢4, 1 9, SEQ:

28.927°S 151.567°E, near entrance to Sundown National Park, 3.11.1999, J.

Moss and L. Popple; 3 da’, 4 99, SEQ: 28°50’S 151°28’E, Glenlyon Dam,

1-2.11.1999, J. Moss and L. Popple; 3 04, 2 99, 26°53’13”S 152°12’49”E,

84 Australian Entomologist, 2015, 42 (2)

Base of Blackbutt Range, SEQ, 26.i.2002, L. Popple and R. MacSloy; 7 33,

4 29, Chinchilla SEQ, 17.iii.2002, L.W. Popple (LWP); 1 3, 8.5 km ESE

Kaimkillenbun, SEQ, 21.1.2001, A. Ewart A(E). Cassinia and grassland,

27°04.30’S 151°29.76°E; 1 9, 7.3 km NE Maclagan, SEQ, 21.1.2004, A.

Ewart (AE); 27°03.13’S 151°41.64’E. The specimens from the base of

Blackbutt Range are illustrated in Fig. 1 (C-D).

a;

A T. mckinnonae sp. nov.

= |© T. verna: former distribution

\|@ 7. verna: new records

e 3

A A

140°E Ye 150°E

20°S E

250km te

Queensland 7

30°S

New South Wales

Fig. 3. Map of Queensland and New South Wales showing the distribution of Toxala

Moulds species. T. mckinnonae sp. n. is restricted to a single locality in northern

Queensland and 7. verna (Distant) is scattered in southeastern Qld, northeastern and

central eastern New South Wales. The former distribution of T. verna was sourced

from Moulds (1990). New specimen records are detailed in the text.

Australian Entomologist, 2015, 42 (2) 85

Calling songs of Toxala cicadas

The calling songs of Toxala verna (Distant) and T. mckinnonae exhibit clear

similarities (Fig. 4). Both species produce a relatively simple repetitive

phrase, comprising a train of syllables, each punctuated by brief silences,

followed by an echeme, another short period of silence, and then a

macrosyllable and a final period of silence before the song repeats. Male

wing-clicks are a common component of the song and occur at around the

beginning of each phrase (at the start of the train of syllables). Field

observations of T. verna at Chinchilla and T. mckinnonae west of Herberton

indicate that, in each species, the female response occurs in the period of

silence following each macrosyllable (pers. obs. 2002, 2007); however, no

recordings of this behaviour have been obtained.

In T. verna, the syllable train has a variable duration (249 ms to 1.886 s) and

may be composed entirely of single syllables (8-10 ms duration; Fig. 5) or

predominantly composed of double syllables (17-19 ms, Fig. 6); in both cases

these are followed by short silences (3-56 ms, Figs 5 and 6). A brief

macrosyllable of 22-26 ms occasionally follows, with a silence of 2-22 ms

preceding the echeme. The echeme varies considerably in duration (103-830

ms) and is followed by a longer period of silence (38-86 ms). A

macrosyllable typically comprising 3-4 (rarely 2) syllables (23-40 ms),

followed by another silence (22-165 ms), completes each phrase (Figs 5 and

6). The call has a phrase duration of 0.793 to 2.687 s and a frequency range

that extends from 4 kHz (principally from 8 kHz) to >22 kHz. Available

recordings indicate that the highest amplitude dominant frequency plateau

extends from 16-20.5 kHz. This is at the limits of the recording equipment;

therefore the plateau could conceivably extend to higher frequencies.

In 7. mckinnonae sp. n., the syllable train (581 ms to 1.654 s duration)

characteristically begins with a long sequence of single syllables (each 8-9

ms duration) followed by a shorter sequence of double syllables (each 15-16

ms duration), with the sounds in each case punctuated by short silences (8-45

ms, Fig. 7). A brief silence at the end of the train of double syllables (5-10

ms) precedes the echeme. The echeme is produced for 148—263 ms and is

followed by a longer period of silence (54-60 ms). A macrosyllable

comprising 3-4 syllables (24-36 ms), followed by another silence (21-92 ms),

completes each phrase (Fig. 7). The call has a phrase duration of 1.031 to

1.840 s and a frequency range that extends principally from 8.5 to >24 kHz.

Available recordings indicate that the highest amplitude dominant frequency

plateau extends from 16 kHz to at least 22 kHz; however, as for the T. verna

recordings, limitations to the sensitivity of the recording equipment could

ultimately mean that the true dominant frequency is considerably higher.

Whilst the similarities between the calling songs of T. verna and T.

mckinnonae sp. n. are evident and obvious, a small number of noticeable

differences in gross song structure are apparent.

86 Australian Entomologist, 2015, 42 (2)

0 1 2 3 4 5s

Fig. 4. Waveform plots illustrating the male calling song of Toxala Moulds cicadas

from different localities in Queensland. (A-C): T. verna (Distant) from Chinchilla

(26°44’S 150°38’E) base of Blackbutt Range (26°53’S 152°13’E) and 8.5 km ESE

Kaimkillenbun (27°04.30’S 151°29.76’E) (this last recording by A. Ewart)

respectively. (D): 7. mckinnonae from ‘Hornet Hill’ west of Herberton (17°23’S

145°21'E).

Australian Entomologist, 2015, 42 (2) 87

Phrase (single repeated unit) Syllable train Echeme Macrosyllable

ere - Moe cae en ic oo se | n

l

~~ mL ee ee = pm pale by ed Se eS oR 6 o ms

J y t

er ‘mia aie sae M H-

syllable 0 (10 ms) Macrosyllable A (24 ms) E EE (28 ms)

i 0

Dominant

frequency

20.5 kHz j

36 v

ap di a na E

d a i | =

| A E

À iah lita Vet si = <{

ike.. og anh jaanasi T

Seer A Ahi

0 2 4 6 8 10 12 14 16 18 20 22 kHz

Fig. 5. Male calling song structure of Toxala verna (Distant) illustrated in expanded

waveform plots. This recording shows a syllable train composed almost entirely of

single syllables, which is the typical structure of most recordings of T. verna. The

spectrogram at the bottom of the figure displays song frequency. This specimen was

recorded in the field at Chinchilla (26°44? S 150°38’E) in south-eastern Queensland.

The most obvious difference is in the structure of the syllable trains at the

start of each phrase. In T. verna, these are almost always composed of single

syllables (occasionally with a macrosyllable at the very end; e.g. Fig. 5). The

only exception is in the recording of an individual from the base of the

Blackbutt Range (Fig. 6). In this example, the syllable train is almost entirely

88 Australian Entomologist, 2015, 42 (2)

Phrase (single repeated unit) Syllable train Macrosyllable Echeme

nN TT

0 100! | 200 300 ms0 250 a a 500 ms

rl) a Pea sh |

N peth Hants

l =m i]

Double syllable (19 ms) Macrosyllable (36 ms)

Dominant o

frequency

18.5 kHz

-36 v

©

E

| we | B

| -72 a

0 2 4 6 8 10 12 14 16 18 20 22 kHz

Fig. 6. Male calling song structure of Toxala verna (Distant) illustrated in expanded

waveform plots. This recording shows a syllable train that predominantly comprises

double syllables, which is atypical among the available recordings of 7. verna. The

spectrogram at the bottom of the figure displays song frequency. This specimen was

recorded in the field at base of Blackbutt Range (26°53’S 152°13’E) in southeastern

Queensland.

composed of double syllables. In contrast, T. mckinnonae produces a

characteristic long sequence of single syllables, followed by a shorter

sequence of double syllables in each syllable train (Fig. 7). Another feature

that appears to be unique to T. mckinnonae is the amplitude modulation in

Australian Entomologist, 2015, 42 (2) 89

Phrase (single repeated unit) Syllable train Echeme

TI]

Macrosyllable

n

[ae 5] ee)

Syllable (5ms) Double syllable (11 ms) Macrosyllable (30 ms)

, 0

Dominant

Background cicadas frequency

MM]

i 18 kHz ee -36

` i | a B wer il Aan N å 3

ba wl Ward ‘s/h aN dib PEN pr Boe. fh, nll aaaea ee tal á "N =

-72 Ẹ

-108

2

v

0 2 4 6 8 10 12 14 16 18 20 22 kHz

Fig. 7. Male calling song structure of Toxala mckinnonae sp. n. illustrated in

expanded waveform plots. This recording shows the characteristic structure of T.

mckinnonae, with regular amplitude modulation and a unique pattern in the syllable

train (cf T. verna in Figs 5 and 6). The spectrogram at the bottom of the figure

displays song frequency. This specimen was recorded in the field at “Hornet Hill’

west of Herberton (17°23’S 145°21'E) in northern Queensland.

each phrase, whereby amplitude increases markedly as the syllable train

proceeds, reduces abruptly during production of the echeme and increases

again when the macrosyllable is produced. This distinctive pattern is not

90 Australian Entomologist, 2015, 42 (2)

present in any of the T. verna recordings, which appear to show little change

in amplitude during the production of each phrase.

Acknowledgements

I would like to thank Anne-Marie McKinnon for assistance with finding and

locating the new species in the field. Appreciation is also extended to Tony

Ewart for making his extensive collection of cicada recordings available. In

addition, Tony Ewart and Max Moulds provided helpful comments on the

manuscript and Susan Wright (Queensland Museum) kindly arranged

deposition and registration of the type material.

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Australian Entomologist, 2015, 42 (2): 91-104 91

A REVIEW OF THE INDO-AUSTRALIAN SUBGENUS

PARAZEUGODACUS SHIRAKI OF BACTROCERA MACQUART

(DIPTERA: TEPHRITIDAE: DACINAE)

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

18/3 McPherson Close, Edge Hill, Cairns, Old 4870

?International Centre for the Management of Pest Fruit Flies, Griffith University, Old 4111

Abstract

The Bactrocera Macquart subgenus Parazeugodacus Shiraki, 1933 (= Melanodacus Perkins,

1937, syn. n.) is reviewed and eight species included: B. (P.) abbreviata (Hardy), B. (P.)

bipustulata Bezzi, B. (P.) matsumurai (Shiraki) and B. (P.) pendleburyi (Perkins) from Asia,

B. (P.) terminifer (Walker) from Sulawesi, B. (P.) satanellus (Hering) from Papua New Guinea,

B. (P.) nigra (Tryon) from Australia and B. (P.) fulvifacies (Perkins) from New Caledonia. Four

of these species are newly transferred: B. fulvifacies from subgenus Zeugodacus Hendel, B. nigra

and B. satanellus from subgenus Melanodacus and B. terminifer from subgenus Paradacus

Perkins. Primary larval hosts are the fruit of olives (Oleaceae). Three Asian species currently

included in Parazeugodacus, viz. B. arisanica (Shiraki), B. fulvosterna Drew & Romig and

B. tetrachaeta (Bezzi), are transferred to subgenus Hemizeugodacus Hardy. Morphological

characters suggesting monophyly of genus Bactrocera are discussed, with those (plus host plant

data) used to support a suggested elevation of Zeugodacus to genus all homoplasious.

Introduction

Recognition of subgenera has long been a feature of the economically

important and closely related fruit fly genera Dacus Fabricius and Bactrocera

Macquart. Within Bactrocera two distinct lineages are present (Drew 1989) —

(1) the Bactrocera and Queenslandacus groups of subgenera, characterised

by a deep posterior emargination to the male fifth sternite (apomorphy), and

(2) the Melanodacus and Zeugodacus groups of subgenera, characterised by a

(plesiomorphic) shallow emargination.

Within these lineages, subgenera traditionally have been defined by the

presence or absence of certain setae and male characters such as the length of

the posterior lobe of the surstylus and presence or absence of a pecten (row of

cilia) on the third abdominal tergite. Most of these, particularly the setal and

pecten characters, are now known to be homoplasious and of little value in

determining placement of atypical species (Hancock and Drew 2006).

The publication of a comprehensive revision of Southeast Asian species by

Drew and Romig (2013), together with that of Australian-Oceanian species

by Drew (1989), means it is now possible to reassess subgeneric limits and

define them as (probable) monophyletic units, thereby enabling a better

understanding of their relationships and biogeography. This paper deals with

subgenus Parazeugodacus Shiraki (including Melanodacus Perkins, here

considered inseparable), which is redefined to contain eight described species

distributed from India to New Caledonia. To avoid unnecessary confusion

here and with both previous and future studies, Me/anodacus is retained as

the group name until the relationships of Tetradacus Miyake are better

understood.

92 Australian Entomologist, 2015, 42 (2)

Genus Bactrocera Macquart

Subgenus Parazeugodacus Shiraki

Parazeugodacus Shiraki, 1933: 107. Type species Parazeugodacus matsumurai

Shiraki, 1933, by original designation.

Melanodacus Perkins, 1937: 57. Type species Dacus niger Tryon, 1927, by original

designation. Syn. n.

Definition. Abdominal sternite V of male with a shallow posterior

emargination; posterior lobe of male surstylus short; pecten of cilia present or

absent on abdominal tergite HI of male; postpronotal setae absent; supra-alar

setae present; prescutellar acrostichal setae present; two pairs of scutellar

setae; wing pattern reduced, the costal band usually very narrow or absent

beyond apex of vein R+; and usually weak or absent in cell rı, anal streak

absent or narrow and largely confined to cell bcu, occiput black; scutum

black with lateral presutural and medial postsutural yellow vittae absent and

lateral postsutural yellow vittae short and triangular or absent; when present

the vittae do not extend across the transverse suture anteriorly.

Response to male lures. Cue-lure (2 species) or none known (6 species).

Included species. B. abbreviata (Hardy), B. bipustulata Bezzi, B. fulvifacies

(Perkins), B. matsumurai (Shiraki), B. nigra (Tryon), B. pendleburyi

(Perkins), B. satanellus (Hering) and B. terminifer (Walker).

Host plants. Wild olives (Oleaceae), with a few records from Symplocos

(Symplocaceae) and other families. Four of the eight species have been

reared from wild olives and cultivated olives are a potential host.

Comments. As 1s normal with Bactrocera subgenera, the defining characters

of Parazeugodacus are individually homoplasious and it is their unique

combination that defines it. Subgenus Melanodacus differed solely in the

absence of the pecten of cilia and the two are therefore regarded as

synonyms, supported by their host plant use. The relationship between

Parazeugodacus and the other largely Oleaceae-feeding subgenus, the

primarily Afrotropical Daculus Speiser, requires further investigation. The

inclusion of Parazeugodacus in the Melanodacus group follows the first clear

definition of the subgenus by Drew and Romig (2013).

Included species

For detailed morphological descriptions of all species and illustrations of all

except B. terminifer see Drew (1989) and Drew and Romig (2013). The wing

of B. terminifer was figured by Hardy (1959).

B. (Parazeugodacus) abbreviata (Hardy, 1974)

Dacus (Zeugodacus) abbreviatus Hardy, 1974: 44. Type locality Laguna, Luzon,

Philippines.

Bactrocera (Parazeugodacus) abbreviata (Hardy): Drew and Romig 2013: 243.

Australian Entomologist, 2015, 42 (2) 93

Distribution: Southern China (Hong Kong, Yunnan), northern Thailand and

Philippines (Luzon).

Host plants. Chionanthus ramiflorus and Olea salicifolia (Oleaceae)

(Allwood et al. 1999).

Male lure: None known.

B. (Parazeugodacus) bipustulata Bezzi, 1914

Bactrocera bipustulata Bezzi, 1914: 153. Type locality Mysore, India.

Bactrocera (Parazeugodacus) bipustulata Bezzi. Drew and Romig 2013: 243.

Distribution: Southern India and Sri Lanka.

Host plant: None known.

Male lure: Cue-lure (Drew and Romig 2013).

B. (Parazeugodacus) fulvifacies (Perkins, 1939)

Zeugodacus fulvifacies Perkins, 1939: 32. Type locality Dumbea, New Caledonia.

Bactrocera (Zeugodacus) fulvifacies (Perkins): Drew 1989: 216.

Distribution. New Caledonia (including Lifou and Maré Islands).

Host plant: Olea paniculata (Oleaceae) (Amice and Sales 1997, Leblanc et

al. 2012).

Male lure: Cue-lure (Amice and Sales 1997).

Comments: Lack of information on the male characters of this species (Drew

1972) has resulted in its previous retention in subgenus Zeugodacus, where it

was placed originally by Perkins (1939). However, examination of

photographs of recent material (in Bishop Museum, Honolulu: L. Leblanc

pers. comm.) revealed short, blunt surstylus lobes and a shallow sternite V

emargination. These and other characters typical of Parazeugodacus,

including the lack of a postsutural yellow vitta and its host plant, necessitate

its transfer to the latter subgenus.

B. (Parazeugodacus) matsumurai (Shiraki, 1933)

Parazeugodacus matsumurai Shiraki, 1933: 107. Type locality Ogasawari I., Bonin

Is, Japan.

Dacus boninensis Hardy & Adachi, 1956: 12. Type locality Bonin Is, Japan.

Bactrocera (Parazeugodacus) matsumurai (Shiraki): Drew and Romig 2013: 246.

Distribution: Japan (Bonin Islands).

Host plant. Osmanthus insularis (Oleaceae) (Ito 1983).

Male lure: None known.

Comments: A record of Litsea japonica (Lauraceae) as the host plant (Drew

and Romig 2013) is likely to be a misidentification; the fruits of Oleaceae and

94 Australian Entomologist, 2015, 42 (2)

many Lauraceae are very similar and only separable by cross-sectioning the

seed. Similarly, the record of Osmanthus insularis as a host of the Lauraceae-

feeding B. (Bactrocera) hyalina (Shiraki) (Drew and Romig 2013) is likely to

be a reciprocal misidentification.

B. (Parazeugodacus) nigra (Tryon, 1927)

Dacus niger Tryon, 1927: 211. Type locality Gympie or Cleveland, Queensland,

Australia.

Bactrocera (Melanodacus) nigra (Tryon): Drew 1989: 182.

Distribution: Eastern Queensland, from Mossman to the south-east highlands.

Host plants: Chionanthus ramiflorus and Olea paniculata (Oleaceae)

(Hancock et al. 2000) and a single record from Symplocos thwaitesii

(Symplocaceae) (Tryon 1927).

Male lure: None known.

Comments: As with B. matsumurai, records of Lauraceae as host plants are

misidentifications of Oleaceae, whereas that from Gomphandra australiana

(Icacinaceae) (May 1953) requires confirmation (Hancock et al. 2000). This

species lacks the pecten of cilia on abdominal tergite III in males, a character

shared with B. (P.) satanellus. The synonymy of Melanodacus with

Parazeugodacus results in the transfer of B. nigra from the former subgenus.

B. (Parazeugodacus) pendleburyi (Perkins, 1938)

Zeugodacus pendleburyi Perkins, 1938: 141. Type locality Bukit Kutu, Selangor,

West Malaysia.

Bactrocera (Parazeugodacus) pendleburyi (Perkins): Drew and Romig 2013: 247.

Distribution. Thailand and West Malaysia.

Host plants. Symplocos cochinchinensis, S. racemosa (Symplocaceae) and

Gmelina arborea (Verbenaceae) (Allwood et al. 1999), all based on single

records with the last requiring confirmation.

Male lure: None known.

B. (Parazeugodacus) satanellus (Hering, 1941)

Melanodacus satanellus Hering, 1941: 48. Type locality Kapakapa, Papua New

Guinea.

Bactrocera (Melanodacus) satanellus (Hering): Drew 1989: 184.

Distribution: Papua New Guinea (Central District).

Host plant: None known.

Male lure: None known.

Comments: The synonymy of Melanodacus with Parazeugodacus results in

the transfer of B. satanellus from the former subgenus.

Australian Entomologist, 2015, 42 (2) 95

B. (Parazeugodacus) terminifer (Walker, 1860)

Dacus terminifer Walker, 1860: 152. Type locality Makassar, Sulawesi.

Dacus (Zeugodacus) terminifer (Walker): Hardy and Adachi 1954: 195.

Bactrocera (Zeugodacus) terminifera (Walker): Drew 1989: 219.

Bactrocera (Paradacus) terminifera (Walker): Drew and Romig 2013: 224.

Distribution. Sulawesi.

Host plant. None known.

Male lure: None known.

Comments: The type and only known specimen is in poor condition and the

abdomen is missing. However, the combination of 4 scutellar setae, supra-

alar and prescutellar acrostichal setae present, reduced wing pattern and black

scutum with no medial and two very short lateral postsutural yellow vittae is

characteristic of Parazeugodacus and this species is thus transferred from

Paradacus Perkins. The combination of short lateral postsutural vittae and a

totally black face is also seen in B. (P.) satanellus but the presence or absence

of a pecten is unknown. Hardy and Adachi (1954) also noted its resemblance

to B. (P.) pendleburyi, which has a similar scutellar pattern.

Excluded species

Three species with a medial postsutural yellow vitta, currently included in

subgenus Parazeugodacus, are excluded here. Although resembling species

within three different species-groups of Zeugodacus, their inclusion in that

subgenus would require three separate reductions of the posterior surstylus

lobes, an unlikely occurrence. The short surstylus lobes and presence of a

medial postsutural yellow vitta, plus the presence of four scutellar setae and

both supra-alar and prescutellar acrostichal setae, are consistent with the

type-species of subgenus Hemizeugodacus Hardy, the Australian B. (H.)

aglaiae (Hardy, 1951). These three species are therefore transferred to that

subgenus, as B. (H.) arisanica (Shiraki, 1933) from Tatwan, Ryukyu Islands

and Thailand, B. (H.) fulvosterna Drew & Romig, 2013 from Sarawak and B.

(H.) tetrachaeta (Bezzi, 1919) from the Philippines. For full descriptions and

illustrations see Drew and Romig (2013).

Two subgroups are recognisable within Hemizeugodacus. Group A, which

includes B. (H.) aglaiae and the three species included here, has a medial

postsutural vitta and occurs from eastern Asia to Australia. Host plants,

known only for B. (H.) aglaiae, are Aglaia spp. (Meliaceae) (Hancock et al.

2000). Group B contains four species from Papua New Guinea, Solomon

Islands and Australia that lack the medial vitta and often have some setae

absent, viz. B (H.) abdomininigra Drew, B. (H.) aurea (May), B. (H.)

buinensis Drew and B. (H.) ektoalangiae Drew & Hancock. Known host

plants are subspecies of Alangium villosum (Alangiaceae) (Hancock et al.

2000). These four species possibly belong in a separate subgenus

96

Australian Entomologist, 2015, 42 (2)

Neozeugodacus May, currently placed as a synonym of Hemizeugodacus.

Subgenus Hemisurstylus Drew, with its sole species B. (H.) melanoscutata

Drew, bred from Garcinia xanthochymus (Clusiaceae) in New Britain (Drew

1989), appears to be related to group B. Two of the Asian species are

attracted to cue-lure (Drew and Romig 2013), while the Australian B. (H.)

aglaiae and B. (H.) aurea are attracted to zingerone, a compound similar to

cue-lure (Royer 2015).

Key to Parazeugodacus species

l

Scutum black without postsutural lateral yellow vittae; fore femora and at

least most of mid and hind femora fulvous; anatergite and katatergite both

MOSTLY AVE OA AN E E ENE E. EON EE ET 2

Scutum black with a pair of short, triangular, postsutural lateral yellow

vittae; fore, mid and hind femora fulvous or extensively black; anatergite

and katatergite yellow or black ........000. 0.0 c cco cece ccc cece cece eee cues 3

Face fulvous; anepisternal stripe reaching line of anterior notopleural seta;

pecten of cilia present on male tergite III (New Caledonia) .................

i E ey pte doc AE gs ie et teak yc Gate. ER en A Set B. (P.) fulvifacies (Perkins)

Face with a pair of black spots; anepisternal stripe not reaching line of

anterior notopleural seta; pecten of cilia absent on male tergite II

(PAUS ne Fe oes oc oc AEN E T ET EAEE ETS B. (P.) nigra (Tryon)

Face fulvous with two black spots; all femora fulvous with at most their

apices fuscous; anatergite and katatergite both mostly yellow ............. 4

Face blackish at least on lower half; fore femora and at least apical half of

mid femora black; anatergite and katatergite yellow or black .............. 6

Scutellum with only a black basal band (Thailand and Malaysia) ........

N PA ote at in EAE, DI pn ai beets ae, B. (P. pendleburyi (Perkins)

Scutellum with a broad black band from base to apex ...................... 5

Face with a pair of small, rounded black spots; abdomen mostly fulvous

with transverse basal black bands on tergites II to IV; fore and mid tibiae

mostly pale fuscous (Bonin Is, Japan) ........ B. (P.) matsumurai (Shiraki)

Face with a pair of medium-sized, oval black spots; abdomen mostly

black with a black medial band and fulvous areas on posterior of tergite II

and submedially on at least tergites III and IV; fore and mid tibiae mostly

dark fuscous (SE Asia) aaea B. (P.) abbreviata (Hardy)

Face fulvous on upper half, dark fuscous to black on lower half; mid

femur fulvous on basal half; anatergite and katatergite both black (India

CS TAA pak eas esd hak Leal ce cect buh terti belly B. (P.) bipustulata Bezzi

Face and mid femur wholly black; anatergite mostly yellow ............... 7

Australian Entomologist, 2015, 42 (2) 97

7 Postpronotal lobes black; scutellum with a broad black band from base to

apex; hind femur black; anepisternal stripe ending before postpronotal

lobe; katatergite black (Papua New Guinea) ... B. (P.) satanellus (Hering)

— Postpronotal lobes mostly yellow; scutellum with only a black basal band:

hind femur fulvous on basal half, black on apical half; anepisternal stripe

reaching postpronotal lobe; katatergite mostly yellow (Sulawesi) .........

ihe th Ar ia EIA PER LARET tess) eae. gees B. (P.) terminifer (Walker)

Discussion

Biogeography

We recognise six biogeographic zones within the Indo-Australian Region

(Fig. 1), each with a high degree of endemism within Bactrocera (Table 1).

Fig. 1. Biogeographic zones in the Indo-Australian region. A = Indian Subcontinent;

B = South-East Asia; C = Wallacea; D = New Guinea; E = Australia; F = South

Pacific.

98 Australian Entomologist, 2015, 42 (2)

Table 1. Number of species in genus Bactrocera and subgenus Parazeugodacus in the

six recognised biogeographic zones in the Indo-Australian Region, together with the

percentage of endemic Bactrocera species (all Parazeugodacus species are endemic

to the particular zone).

“Biogeographic Zone No. speciesof %endemic No. species of _

Bactrocera Bactrocera Parazeugodacus

(A) Indian subcontinent 74 72 1

(B) South-East Asia 223 84 3

(C) Wallacea 124 82 1

(D) New Guinea 170 85 1

(E) Australia 76 75 1

(F) South Pacific 59 85 1

The eight described species of subgenus Parazeugodacus occur largely

allopatrically from India and Japan to Australia and New Caledonia and are

represented in all six biogeographic zones (Table 1). The four Oriental

Region [Zones A-B] species (bipustulata, matsumurai, abbreviata and

pendleburyi) are individually distinctive and likely represent vicariant

speciation. The four Australian Region [Zones C-F] species form two distinct

but not necessarily directly related pairs — one with a wholly black scutum

(apomorphy) and largely or entirely fulvous face and femora (nigra and

fulvifacies) and one with the short lateral postsutural vittae present, a wholly

black face (apomorphy) and largely black femora (terminifer and satanellus).

Subgeneric relationships

The Melanodacus group contains the Afrotropical subgenera Daculus Speiser

(= Afrodacus Bezzi) and Gymnodacus Munro, plus the Indo-Australian

subgenera Hemisurstylus Drew, Hemizeugodacus Hardy (= Neozeugodacus

May) and Parazeugodacus Shiraki (= Melanodacus Perkins). Molecular

evidence (Krosch et al. 2012) suggests that typical species of Paratridacus

Shiraki also belong here (i.e. those lacking a medial postsutural vitta and with

a relatively short and very broad posterior surstylus lobe, including the type-

species).

The Bactrocera, Melanodacus and Zeugodacus groups all appear to be

derived from the ancestral subgenus Tetradacus. Within the Dacini (Dacina

of some authors), two pairs of scutellar setae occur in most of the species in

the Melanodacus and Zeugodacus groups and that character is a possible

synapomorphy within the tribe, regained after an initial reduction to one pair

in the ancestral taxon (Jchneumonopsis Hardy or Monacrostichus Bezzi). If

two pairs of scutellar setae were to be considered plesiomorphic for the tribe,

then independent loss of the basal pair would be required in the ancestral

species of each of the five outgroups (Jchneumonopsis, Monacrostichus,

Australian Entomologist, 2015, 42 (2) 99

Dacus, Tetradacus and Bactrocera group), rather than a single reacquisition

in the ancestor of the Melanodacus+Zeugodacus groups followed by some

secondary losses. White (2006) also noted the tendency for ‘lost’ setae to

reappear within the Dacini and the exceptional occurrence of four scutellar

setae in some specimens of B. (Bactrocera) quadrisetosa (Bezzi) is certainly

homoplasious. In two unrelated subgenera, Notodacus Perkins (Bactrocera

group) and Heminotodacus Drew (Zeugodacus group) (and also, weakly, in

B. (Zeugodacus) hatyaiensis Drew & Romig), postpronotal setae reappear.

Ecological and morphological characters used to support a suggested

relationship between Zeugodacus and Dacus (White 2006, Virgilio et al.

2015), viz. shared Cucurbitaceae host plants, presence of a postsutural medial

yellow vitta and presence of notopleural vittae or spots along the anterior

margin of the transverse suture, are all homoplasious and occur also in the

Bactrocera group. Cucurbitaceae (and Passifloraceae, another widespread

Dacus host plant) are major hosts of B. (Bactrocera) bryoniae (Tryon)

(Hancock et al. 2000). A postsutural medial vitta occurs in subgenera

Hemizeugodacus (Melanodacus group), Apodacus Perkins (B. visenda

(Hardy), B. cheesmanae (Perkins) and B. neocheesmanae Drew), Bulladacus

Drew & Hancock, Notodacus Perkins (all Bactrocera group) and Tetradacus

(Drew 1989, Drew and Romig 2013). A notopleural sutural spot occurs in

subgenera Hemizeugodacus (two of the three species here removed from

Parazeugodacus), Bulladacus and Notodacus, in B. (Bactrocera) digressa

Radhakrishnan and in B. (Tetradacus) brachycera (Bezzi), while a

notopleural vitta occurs in B. (Tetradacus) minax (Enderlein) and the

Australian B. (Bactrocera) mendosa (May) (Drew 1989, Hancock 2009,

Drew and Romig 2013). Occurrence of the morphological characters in both

Dacus and Tetradacus (and also in Monacrostichus) suggests that they are

plesiomorphies present in the common ancestor of Dacus + Bactrocera.

Non-cucurbitaceaous host plants also occur in several subgenera within the

Zeugodacus group (Drew 1989, Drew and Romig 2013) and the use of

cucurbits, as in Dacus, is likely to represent a secondary shift away from

tropical fruit (ancestral Zeugodacus group) or asclepiad pods (ancestral

Dacus: see Hancock and Drew 2006). Further evidence for this is provided

by several polyphagous species of Zeugodacus, especially B. (Z.) cucurbitae

(Coquillett) and B. (Z.) tau (Walker), reared from 8-11 plant families other

than Cucurbitaceae (Allwood et al. 1999). Many of these non-

cucurbitaceaous hosts are also utilised by typical Bactrocera species but not

by Dacus, which is recorded only from Apocynaceae (primarily subfamily

Asclepiadoideae), Cucurbitaceae and Passifloraceae (Hancock and Drew

2006). Thus, ecological and morphological characters do not support the

raising of Zeugodacus from subgenus group to genus as proposed by Virgilio

et al. (2015) [who, based on diagnoses prior to Drew and Romig (2013) and

without examining material, included Parazeugodacus within it but left

Melanodacus in Bactrocera s.s.|, leaving only molecular evidence that is

100 Australian Entomologist, 2015, 42 (2)

weakly supported by low Bayesian PP and ML bootstrap support values at all

critical nodes on the resulting (and non-congruent) consensus trees.

Monophyly of genus Bactrocera is suggested by the presence of a distinct,

black, T-shaped marking on the abdomen, generally accompanied by black

lateral markings on tergites II-V that are often divided into discrete

rectangular or triangular anterolateral patches. These markings, contrasting

strongly with the pale areas of the abdomen, occur in many species in the

Bactrocera, Melanodacus and Zeugodacus groups of subgenera and also in

subgenus Tetradacus but are not seen elsewhere in the Dacinae (including the

tribes Gastrozonini and Ceratitidini) and appear to be a synapomorphy for the

genus. Dark abdominal markings in Dacus, where they occur, are diffuse and

not clearly demarcated. Black scutellar apical spots or medial bands are also a

recurring character in the Bactrocera, Melanodacus and Zeugodacus groups,

as is the reappearance of prescutellar acrostichal setae in most of the species

in all three groups; these are also possible synapomorphies within

Bactrocera, being absent in all species of Jchneumonopsis, Monacrostichus,

Dacus and subgenus Tetradacus. Within the genus only the Bactrocera group

(including Queenslandacus Drew) appears to have the deep emargination to

male sternite V and only the Zeugodacus group appears to have a long,

narrow and often finger-like posterior surstylus lobe (each of which is

regarded as a synapomorphy for those groups). However, since both states

also occur within Dacus (Munro 1984) neither is suitable for characterising

genera. The Melanodacus group retains plesiomorphic states of both

characters (i.e. shallow sternite V emargination and short surstylus lobes) and

therefore is not necessarily monophyletic. The subgenera we currently

recognise and the number of included species are listed in Table 2.

Table 2. List of subgenera in genus Bactrocera (sensu Drew 1989, Drew and

Hancock 1999, Drew and Romig 2013, Hancock 2015), with the number of species

we currently include. Note that Paratridacus and Parazeugodacus are included in the

Melanodacus group and Queenslandacus is included in the Bactrocera group, while

B. superba Drew & Romig is of uncertain affinity and left unplaced.

Subgenera African species Asia-Pacific species

Bactrocera Group

Apodacus Perkins 0) 3!

Bactrocera Macquart 2° ca 380

Bulladacus Drew & Hancock 0) 21

Calodacus Hancock 0 6

Notodacus Perkins 0 3

Queenslandacus Drew 0) 1

Semicallantra Drew 0 F

Trypetidacus Drew 0 i

Australian Entomologist, 2015, 42 (2) 101

Subgenera African species Asia-Pacific species

Melanodacus Group

Daculus Speiser 8 0)

Gymnodacus Munro 2 0

Hemisurstylus Drew 0) 1

Hemizeugodacus Hardy 0 8"

Paratridacus Shiraki 0) 7

Parazeugodacus Shiraki 0 8"

Tetradacus — ancestral subgenus?

Tetradacus Miyake 0) 7s

Zeugodacus Group

Aglaodacus Munro j 0)

Asiadacus Perkins 0) 7

Austrodacus Perkins 0 4

Diplodacus May 0) i

Hemigymnodacus Hardy 0 2

Heminotodacus Drew 0 i

Hemiparatridacus Drew 0 1

Javadacus Hardy 0) 8°

Nesodacus Perkins 0) li

Niuginidacus Drew 0 i

Papuodacus Drew 0) 6

Paradacus Perkins 0) 6°

Parasinodacus Drew & Romig 0) is

Sinodacus Zia 0 19

Zeugodacus Hendel p 120

'Three Australian and New Guinea species with narrow, slit-like ceromata: B. visenda

(Hardy), B. cheesmanae (Perkins) and B. neocheesmanae Drew. *The two African

species (B. zonata (Saunders) and B. invadens Drew, Tsuruta & White) are

introductions from India. *Includes B. toxopeusi (Hering) from Papua Province,

Indonesia. “This study. "Four additional Papua New Guinea species belong in either

Austrodacus (B. alampeta Drew, B. atrisetosa (Perkins) and B. unichromata Drew) or

Zeugodacus (B. mesonotaitha Drew). ‘Includes B. mesonotochra Drew, B.

neopagdeni Drew and B. pagdeni (Malloch) from Papua New Guinea and Solomon

Islands. "The Madagascan B. (A.) nesiotes (Munro). *Two additional Australian

species belong in subgenus Bactrocera: B. melanothoracica Drew and B. unirufa

Drew. Three additional Papua New Guinea species belong in Zeugodacus (B.

aurantiventer Drew, B. citroides Drew and B. decipiens (Drew)). ‘Includes B.

abdopallescens (Drew) from Papua New Guinea and B. perpusilla (Drew) from New

Caledonia. ''The sole African species (B. cucurbitae) is an introduction from India.

102 Australian Entomologist, 2015, 42 (2)

With regard to the phylogeny of Bactrocera species presented in Virgilio et

al. (2015), it should be noted that the Indonesian specimen of B. dorsalis

(Hendel) used in their analysis is clearly a misidentified B. papayae Drew &

Hancock (which they placed adjacent to it), since only the latter is known to

occur in Indonesia. The two ‘Afrodacus’ species used in the analysis (taken

from Krosch et al. 2012), B. jarvisi (Tryon) and B. minuta (Drew), were both

transferred to subgenus Bactrocera by Copeland et al. (2004). The specimen

of B. unirufa Drew used in the analysis (also taken from Krosch et al. 2012)

is likely to be a misidentification of B. melanothoracica Drew, a species with

a very variable scutal pattern (Royer and Hancock 2012). Placement of both

these species in subgenus Bactrocera (as inferred by Virgilio et al. 2015) is

supported by the deep emargination to sternite V and the produced but still

relatively short posterior surstylus lobe in males.

Full justification for the subgeneric transfer of species indicated in Table 2

will occur in future papers in this series; most are a result of improved

subgeneric diagnoses in Drew and Romig (2013).

Acknowledgements

Luc Leblanc (University of Hawaii) kindly supplied photographs of the

previously unrecorded surstylus of B. fulvifacies. The comments from several

referees are greatly appreciated.

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