THE AUSTRALIAN ENTOMOLOGIST

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COVER

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

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

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

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

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

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

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

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

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

Australian Entomologist, 2016, 43 (2): 49-54 49

NORFOLK ISLAND'S CADDISFLY IS A NEW ZEALANDER

(TRICHOPTERA: HYDROPTILIDAE)

ALICE WELLS! and KARL KJER?

‘Australian National Insect Collection, CSIRO, PO Box 1700, Canberra, ACT 2601

(Email: alice.wells 2 csiro.au)

^Department of Entomology and Nematology, UC Davis, Davis CA 95616, USA

(Email: kkjer@ucdavis.edu)

Abstract

Triplectides australis Navas, a leptocerid species reported in 1917 from a single specimen but

not collected since, and Oxyethira albiceps (McLachlan), the microcaddisfly species reported

here, are the only caddisflies ever recorded from Norfolk Island. This impoverished trichopteran

fauna is discussed and compared with the notably endemic faunas of Lord Howe Island and New

Caledonia, the two other major islands on the Norfolk Ridge of the southwestern Pacific.

Comparisons of COI data confirm the initial morphology-based identification of the Norfolk

Island microcaddisfly as a New Zealand species.

Introduction

The first record of Trichoptera for Norfolk Island was of a single female

reported by Tillyard (1917) as the *... very common Australian species ...’

Notanatolica magna Walker, 1852. Subsequently, this was considered by

Hawkins (1943) to be a misidentification for Triplectides cephalotes (Walker,

1852). However, in their review of the genus Triplectides Kolenati, 1859,

Morse and Neboiss (1982) identified Tillyard’s specimen as a different, also

common Australian species, Triplectides australis Navas, 1934. Smithers

(1998) also listed it as Triplectides australis. We have been unable to locate

the specimen in collections of either the South Australian Museum or

Museum Victoria, the two most likely depositories. Until now, Tillyard’s

specimen represented the only trichopteran reported from Norfolk Island.

Here we report on another trichopteran species, taken by light trapping. This

was the only species taken at light over the years 2012-2015, which suggests

that the Triplectides species could simply have been an adventive, or may

have become extinct.

On both Norfolk Island and Lord Howe Island, the more southerly and

smaller of the set of islands on the southwestern Pacific Norfolk Ridge,

permanent running water is scarce, in contrast with New Caledonia to their

north. Norfolk Island and Lord Howe Island are approximately 35 km^ and

15 km? in size, respectively; both are eroded volcanoes and have very few

permanent to semi-permanent streams and few still-water bodies. In contrast,

the far larger New Caledonia (area 18,575 km”), with its central massif from

which numerous streams arise, has a rich, highly endemic Trichoptera fauna:

the current total is 239 species, most of which are endemic (Johanson and

Wells, in prep.). Nonetheless, for Lord Howe Island, seven endemic

Trichoptera species have been described and life stages of at least three other

species have been reported (Wells 2011). Yet from Norfolk Island, despite

repeated light trapping recently at several of the permanent lotic and lentic

50 Australian Entomologist, 2016, 43 (2)

water bodies, only a single species was taken. This is a microcaddisfly

(Hydroptilidae), not the leptocerid species that was recorded previously for

the island. In this present study, the identification based on morphological

features is tested by comparisons of COI data from Norfolk Island specimens

with data from New Zealand material (BOLD 2015).

Norfolk Island's environment has a bleak post-European settlement history.

Clearing of vegetation for horticulture and agriculture, pollution of the

ground water systems (see Abel and Falkland 1991, Diatloff 2007), invasion

by weeds (NIQS 2014) and physical damage caused by cattle, have resulted

in the severe degradation of the few waterways (e.g. Figs 1-2). Just how

many of the small number of aquatic species listed by Smithers (1998) as

having been recorded from the island currently live and breed there is

unknown. Among Odonata, Smithers listed two species of Coenagrionidae

and two Anisoptera; among Hemiptera: Hydrometridae (1) and Veliidae (1);

among Coleoptera: Gyrinidae (2) and Hydrophilidae (2); among Diptera:

Tipulidae (at least 5), Culicidae (4), Ceratopogonidae (7) and Simuliidae (1);

and in Lepidoptera one unidentified species of Nymphulinae (2 Crambidae).

If comparisons with Lord Howe Island with its smaller size and considerable

level of endemism among Trichoptera have any validity, it seems probable

that very early following the settlement of the island by Europeans in March

1788, even before the first insects were recorded for Norfolk Island, the

island's aquatic habitats were probably severely impacted, resulting in loss of

species (see Abel and Falkland 1991, Diatloff 2007, Coyne 2011).

Figs 1-2. Cockpit Falls Reserve was one of the collecting sites where, over the years

2012-2015, cattle roamed freely: (1) the stream was choked along some reaches by

water hyacinth (Eichhornia crassipes); (2) elsewhere damaged severely by cattle; and

just a little further upstream of these sections was choked by a thicket of taro

(Colocasia esculenta)and bullrushes (Typha sp.).

Over the years 2012 to 2015, a quarantine survey of plant and animal species

of Norfolk Island was conducted. The survey concentrated primarily on

Australian Entomologist, 2016, 43 (2) 51

invasive species of flora, pathogens and fauna, targeting particularly groups

of agricultural and quarantine concern. For some groups, comprehensive

species lists were developed (e.g. weedy plants - NIQS 2014; Thysanoptera —

Mound and Wells 2015). Mound and Wells (2015) found that around 5096 of

the 66 thrips species they took on the island are adventives; six species were

recorded that were described originally from New Zealand; and 12 species

were recorded as endemics. Among the island's flora of some 183 vascular

plant species, 43 endemics are recorded (Coyne 2011). The collection of

Trichoptera samples was an adjunct to the survey.

Methods

Light-trap specimens were collected into and stored in 95% ethanol.

Specimens are deposited in the Australian National Insect Collection (ANIC),

CSIRO, Canberra, Australia. Initial species determination was made by

examination of slide mounts prepared by maceration in caustic potash,

dehydration and clearing in clove oil, followed by mounting in Canada

balsam. Ten male specimens were selected for barcoding, their genitalia

removed and prepared as Canada balsam-mount vouchers (ANIC).

We collected mitochondrial DNA data from a fragment of cytochrome

oxidase, subunit 1 (COL; the barcode fragment), using standard Folmer

primers (Folmer et al. 1994), as modified in Zhou et al. (2009). DNA was

extracted with the “Hotshot” method (Truett et al. 2000) and PCR was

performed using standard protocols (Zhou et al. 2009). Amplified DNA was

commercially purified and sequenced in both strands at Genewiz

(Piscataway, New Jersey). Trace files were manually trimmed and edited.

Sequences were submitted to the BOLD website for identification (http://

www.boldsystems.org/index.php/IDS_OpenIdEngine). Identifications were

made based on the phylogenetic tree option.

Results

All of the Trichoptera specimens taken (more than 500, including males and

females) were identified as belonging to a single microcaddisfly species,

Oxyethira albiceps (McLachlan, 1862) (identification confirmed by Brian

Smith pers. comm., based on morphological information — Figs 3-6 allow

comparison of the Norfolk Island specimens with copies of the figures that

accompanied the original description, reproduced from Mosely and Kimmins

1953). This species 1s widely distributed and common in New Zealand on the

three main islands (North, South and Stewart Islands) and also occurs on

Snares, Antipodes, Auckland, Campbell and Chatham Islands (McMurtrie et

al. 2014); its larvae feed on filamentous green algae and the species is known

to be tolerant of enriched waters.

Amplification of DNA from Norfolk Island specimens was successful for 7

of the 10 Norfolk Island specimens extracted (Table 1). Comparison of the

resulting sequences revealed that three of the specimens were identical with

52 Australian Entomologist, 2016, 43 (2)

data available through the BOLD site for Oxyethira albiceps from Waikato,

in the upper northern part of the North Island of New Zealand. Similarity of

four others ranged between 99.4% and 99.8%. Thus, our data fail to indicate

any significant difference between the two populations.

3 Pp

Po f

K À

kod

Áo

Figs 3-6. (3-4) Norfolk Island Oxyethira albiceps (McLachlan, 1862) male genitalia

(BOLD voucher £06): (3) ventral view; (4) lateral view. (5-6) New Zealand O.

albiceps holotype male genitalia: (5) ventral view; (6) lateral view (modified after

Mosely and Kimmins 1953).

Discussion

In contrast with the two other islands of the Norfolk Ridge, Norfolk Island

appears to have no endemic Trichoptera species. Conceivably, the only

species in the water bodies sampled, the hydroptilid Oxyethira albiceps,

colonised waterways that were devoid of Trichoptera until efforts were made

quite recently to reduce the severe pollution described by Diatloff (2007).

The species could even have persisted in the degraded systems from earlier

times, since O. albiceps tends to be tolerant of enriched waters (see Lavender

et al. 2004). Comparison of COI data from Norfolk Island and New Zealand

supports a very recent association, not ruling out repeated colonisation. It is

possible that colonisation occurred as it has by other insect species and

possibly still occurs, either by chance dispersal on the south-east trade winds

Australian Entomologist, 2016, 43 (2) 53

(Holloway 1982) or through human traffic. In contrast with New Zealand's

other caddisfly species (around 200 described species), its ubiquity,

abundance and tolerance of high nutrient levels in streams would make O.

albiceps an ideal candidate for establishment on Norfolk Island.

Table 1. Similarity (in %) between the Norfolk Island sample of Oxyethira albiceps

(McLachlan, 1862) and specimens from Waikato in the North Island of New Zealand

(NZ data accessed from BOLD courtesy of Ian Hogg).

Norfolk Island Site BOLD voucher £ Similarity (46)

with NZ Waikato

(Data from

BOLDSYSTEMS

2015)

Cascade Falls (2 Cockpit Reserve) AWNICADISNI-01

Cascade Falls (2 Cockpit Reserve) AWNICADISNI-02

Acknowledgements

The field work was an offshoot of the 2012-15 Norfolk Island Quarantine

Survey of Norfolk Island's flora, pathogens and fauna supported by the

Australian Department of Agriculture. Brian Smith, National Institute of

Water and Atmospheric Research (NIWA), New Zealand, is acknowledged

for initial confirmation of the identity of the specimens, and Ian Baird for

permission to access the New Zealand COI data for the species. The

Australian National Insect Collection, CSIRO, is thanked for provision of

laboratory facilities to AW.

References

ABEL, R.S. and FALKLAND, A.C. 1991. The hydrogeology of Norfolk Island, South Pacific

Island. Bureau of Mineral Resources, Australia, Bulletin 534: 57 pp. (online at

http://www. ga.gov.au/corporate_data/3 1/Bull_234.pdf).

BOLD 2015. BOLDS YSTEMS. Attp://www.boldsystems.org/

COYNE, P. 2011. Norfolk Island's fascinating flora. Petaurus Press, Belconnen; 192 pp.

DIATLOFF, N. 2007. Norfolk Island hydrology study, the background to water & waste

management on Norfolk Island. The Government of Norfolk Island; 24 pp. (online at

http://www. info. gov.nflland &env/Environment/NI%20Hydrology%20Study.pdf).

54 Australian Entomologist, 2016, 43 (2)

FOLMER, O., BLACK, M., HOEH, W., LUTZ, R. and VRISENHOEK, R. 1994. DNA primers

for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan

invertebrates. Molecular Marine Biology and Biotechnology 3: 294-299.

HAWKINS, C.N. 1943. The insects of Norfolk Island, including a preliminary report on a recent

collection. Annals and Magazine of Natural History (11) 9: 865-902.

HOLLOWA Y, J.D. 1982. Further notes on the Lepidoptera of Norfolk Island, with particular

reference to migrant species. Journal of Natural History 16: 351-365.

LAVENDER, R., MEREDITH, A. and ANTHONY, M. 2004. Results of ecosystem health

monitoring in Canterbury region between November 2000 and February 2003. Report U04/113,

Environment Canterbury, Technical Report, Investigations and Monitoring Group; 95 pp. (online

at http://ecan.govt.nz/publications/Reports/results-ecosystem-health-monitoring-canterbury-

region.pdf).

MCLACHLAN, R. 1862. Characters of new species of exotic Trichoptera; also of one new

species inhabiting Britain. Transactions of the Royal Entomological Society of London 11(3):

301-311.

MCMURTRIE, S.A., SINTON, A.M.R. and WINTERBOURN, M.J. 2014. Lucid identification

key to Campbell Island freshwater invertebrates: Oxyethira albiceps information sheet. EOS

Ecology, Christchurch, New Zealand; 2 pp.

MORSE, J.C. and NEBOISS, A. 1982. Triplectides of Australia (Insecta: Trichoptera:

Leptoceridae). Memoirs of the National Museum of Victoria A3: 61-98.

MOSELY, M.E. and KIMMINS, D.E. 1953. The Trichoptera (caddis-flies) of Australia and New

Zealand. British Museum (Natural History), London; 550 pp.

MOUND, L.A. and WELLS, A. 2015. Endemics and adventives: Thysanoptera (Insecta)

biodiversity of Norfolk, a tiny Pacific Island. Zootaxa 3964: 183-210.

NIQS (Norfolk Island Quarantine Survey). 2014. Norfolk Island weeds handbook. Australian

Government, Canberra; 54 pp.

SMITHERS, C.N. 1998. A species list and bibliography of the insects recorded from Norfolk

Island. Technical Reports of the Australian Museum 13: 1-55.

TILLY ARD, R.J. 1917. Odonata, Planipennia, and Trichoptera from Lord Howe and Norfolk

Islands. Proceedings of the Linnean Society of New South Wales 42: 529-544.

TRUETT, G.E., HEEGER, P, MYNATT, R.L., TRUETT, A.A., WALKER, J.A. and

WARMAN, M.L. 2000. Preparation of PCR-quality mouse genomic DNA with Hot Sodium

Hydroxide and Tris (HotSHOT). Biotechniques 29: 52-54.

WELLS, A. 2011. The Trichoptera of Lord Howe Island, including 3 new species, larvae and

keys. Zootaxa 2987: 45-55.

ZHOU, X., ADAMOWICZ, S.J., JACOBUS, L.M., DEWALT, R.E. and HEBERT, P.D. 2009.

Towards a comprehensive barcode library for arctic life — Ephemeroptera, Plecoptera, and

Trichoptera of Churchill, Manitoba, Canada. Frontiers in Zoology 6: 30. http://doi.org/

10.1186/1742-9994-6-30

Australian Entomologist, 2016, 43 (2): 55-68 55

FIRST AUSTRALIAN RECORDS OF EUPLOEA WALLACEI MELIA

FRUHSTORFER, 1904 AND EUPLOEA STEPHENSII JAMESI

BUTLER, 1876 (LEPIDOPTERA: NYMPHALIDAE: DANAINAE)

FROM DAUAN ISLAND, TORRES STRAIT, QUEENSLAND, WITH

NOTES ON THE AGGREGATION HABITS OF EUPLOEA

FABRICIUS SPECIES NEAR FLOWERING MANGROVES

TREVOR A. LAMBKIN

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

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

Abstract

Euploea wallacei melia Fruhstorfer, 1904 and E. stephensii jamesi Butler, 1876 are

recorded from Dauan Island, Torres Strait, Queensland for the first time, bringing the

number of Euploea Fabricius species recorded from the island to ten. A large wet

season roosting of danaines, predominantly Euploea spp, was observed on Dauan

Island in late December 2009 and early January 2010. The aggregation clustered near

flowering trees of the White-flowered Black Mangrove, Lumnitzera racemosa

(Combretaceae). Two humid mangrove glades with large amounts of L. racemosa

flowers were preferred loci for the 10 Euploea species encountered, with several

hundred butterflies observed feeding on blossom or roosting in each of the glades at

any one time. Most of these individuals were newly emerged. Butterflies roosted

overnight in the glades prior to feeding on L. racemosa flowers the following

morning. This congregation of Euploea butterflies within the mangrove blossom

environment was temporary, as butterfly numbers declined markedly by the middle of

January 2010, after cessation of the mangrove blossom. Communal roosting by

Euploea at nectar sources is previously unrecorded in Australia and this phenomenon

is suggested to be the result of newly emerged butterflies at the start of the wet season

migrating from their breeding environments to feed at discrete, nectar-rich areas.

Introduction

The summer monsoon season is a key weather event for much of northern

Australia, including the islands of Torres Strait in northern. Queensland

(Bowman et al. 2010). The onset of the season is characterised by a southerly

moving band of low pressure systems (Fig. 1), which generates moist

northwesterly winds that variably begin between late November and late

January (Jones 1987) and persist until April or May.

On Dauan, one of the most northerly of the Torres Strait islands, the

dominant vegetation types are deciduous monsoon forest and semi-evergreen

mesophyll vine forest (Webb 1959, Torres Strait Regional Authority 2013),

which grow extensively among exposed granite boulders (Fig. 2). The

highest part of this boulder stack is Mt Cornwallis, at approximately 300 m.

Crow butterflies, Euploea Fabricius, 1807, are commonly observed during

the wet (monsoon) season on Dauan Island. Euploea species are essentially

tropical in distribution, occurring in the Oriental and Australian regions

(Ackery and Vane-Wright 1984, Scheermeyer 1999), with the greatest

diversity in the Indo-Australian region (Corbet and Pendlebury 1992, Ackery

56 Australian Entomologist, 2016, 43 (2)

and Vane-Wright 1984, Parsons 1998), particularly on Java and Sumatra and

in New Guinea (Kitching and Scheermeyer 1993, Scheermeyer 1999).

Eight species of Euploea have been recorded from Dauan Island to date, viz.

E. algea amycus Miskin, 1890, E. sylvester sylvester (Fabricius, 1793), E.

alcathoe misenus Miskin, 1890, E. tulliolus dudgeonis (Grose-Smith, 1894),

E. batesii batesii C. & R. Felder, 1865, E. corinna (W.S. Macleay, 1826), E.

netscheri erana (Fruhstorfer, 1910) and E. leucostictos (Gmelin, 1790).

In early January 2010, E. wallacei melia Fruhstorfer, 1904 and E. stephensii

jamesi Butler, 1876, two species previously unrecorded from Australia, were

collected from within mangrove glades adjacent to flowering trees of the

White-flowered Black Mangrove, Lumnitzera racemosa (Combretaceae).

Lumnitzera racemosa (Fig. 3) is a tropical and subtropical species (Lear and

Turner 1977, Williams 1987) that occurs commonly in the low-lying areas on

the eastern and western ends of Dauan Island (Figs 4-5) and typically flowers

within the first month after the onset of the monsoon or wet season. In the

last week of December 2009, through to the second week of January 2010, L.

racemosa flowered quite profusely and attracted large numbers of danaine

butterflies, including Danaus affinis affinis (Fabricius, 1775) (Fig. 6),

Tirumala hamata hamata (Macleay, 1827) and, in particular, several Euploea

species.

Considering the diversity of Euploea in New Guinea, with at least 16 species

known (Yata and Morishita 1985, Parsons 1998, Scheermeyer 1999) and the

proximity of Torres Strait to New Guinea, it is not surprising that 10 species

are now recorded from Dauan Island. This brings the total number of Euploea

species now known from Torres Strait islands to 12 (Braby 2000, Meyer et

al. 2004, Lambkin and Knight 2007).

A critical survival strategy for several Euploea species and other danaine

species in monsoon regions is to aggregate or roost, often in large

communities and typically over the dry season (Kitching and Zalucki 1981,

Scheermeyer 1993, Canzano et al. 2003). Typically, individuals within these

aggregating communities persist in a semi-quiescent state for the duration of

the dry period. These aggregations can be semi-permanent in their roosting

locations and revisited over several dry seasons (Monteith 1982). In contrast

to this dry season behaviour, here I document an opportunistic, wet season

roosting of Euploea and other danaines near nectar sources on Dauan Island

in late December 2009 and early January 2010. The aggregating Euploea

species included the two previously unrecorded from Australia.

The following abbreviations refer to public institutions and private

collections from which material was examined: MDBC - M. De Baar

collection (Euploea spp now in TLIKC); QDAFC - Queensland Department

of Agriculture and Fisheries collection, Brisbane; TLIKC - Joint collection of

T.A. Lambkin and A.I. Knight, Brisbane.

Australian Entomologist, 2016, 43 (2) 57

Figs 1-8. (1) Monsoon cloud band over northern Australia, 9.1.2014. (Australian

Government, Bureau of Meteorology); (2) hill on western end of Dauan composed of

granite boulder stacks; (3) flowering Lumnitzera racemosa on Dauan; (4) northern

aspect of western end of Dauan, panhandle covered mostly with mangroves; (5) aerial

view of southern aspect; (6) Danaus affinis affinis visiting blossom of L. racemosa;

(7) mangrove glade on Dauan after termination of flowering; (8) male Euploea

alcathoe misenus roosting under Rhizophora stylosa in mangrove glade.

58 Australian Entomologist, 2016, 43 (2)

Abbreviations of collectors' names appearing on specimen labels are: GS —

collector unknown; IFTA - Insect Farming and Trading Agency, Bulolo,

Papua New Guinea; JFG — J.F. Grimshaw; JG — J. Guyomar; JH — J. Hancox;

PS - P. Shanahan; RS — R. Straatman; TAL - T.A. Lambkin.

Specimens examined

Euploea wallacei C. & R. Felder, 1860

(Figs 9-14)

QUEENSLAND: 1 (, Dauan Island, Torres Strait, 9°25'S 142?32'E, 9.1.2010, TAL

(TLIKC); 1 9, same data except 10.1.2010 (TLIKC).

PAPUA NEW GUINEA: 1 ĝ, Kiunga, Western Province, -.ii.1991, JH (TLIKC, ex

MDBO); 1 ĝ, Lae, Morobe Province, -.ix.1969, GS (TLIKC, ex MDBC); 7 dd, 1 9,

Sambio Mumeng, Morobe Province, -.xii.1984 (2 4), -.i.1985 (4 43), -.v.1985 (1

d, 1 9), IFTA (TLIKC, ex MDBC); 2 Gd, Wau, Morobe Province, 1200 m,

24.x.1975 (TLIKC, ex MDBC); 1 9, same data except -.iv.1974, PS (TLIKC, ex

MDBC).

INDONESIA: 2 3, Halmahera I., Northern Moluccas, -.vi.1996 (TLIKC, ex

MDBC); 1 3, 1 9, Dobo, Wamar I., Aru Is, 1992 (TLIKC, ex MDBC); 1 d,

Manokwari, West Papua Province, 1989 (TLIKC, ex MDBC).

Euploea stephensii C. & R. Felder, [1865]

(Figs 17-20)

QUEENSLAND: 1 9, Dauan Island, Torres Strait, 9°25'S 142?32'E, 9.1.2010, TAL

(TLIKC).

PAPUA NEW GUINEA: 1 <6, Imonda, West Sepik Province, 15.v.1992, JFG

(QDAFC); 1 2, Nuku Mission, NW Central Highlands Province, -.viii.1977 (TLIKC,

ex MDBC); 1 ĝ, Lae, Morobe Province, 10.xii.1969, GS (TLIKC, ex MDBC); 1 9,

Sambio Mumeng, Morobe Province, -.xii.1984, IFTA (TLIKC, ex MDBC); 1 d,

Janita Village, 10 km E of Popondetta, Oro Province, 22.x.1987, JG (TLIKC, ex

MDBC).

INDONESIA: 1 3, Kofiau I., N of Misool I., 1992 (TLIKC, ex MDBC); 1 c, Waigeo

I., 26.ix.1985, RS bequest (TLIKC, ex MDBC); 1 ĝ, sama data except 2.x.1985, RS

(TLIKC, ex MDBC); 1 ĝ, Yapen I., Irian Bay, Papua Province, -.i.1996 (TLIKC, ex

MDBO); 1 4, Nabire, Papua Province, -.x.1991 (TLIKC, ex MDBC); 1 9, Jaya Pura,

Papua Province, 1989 (TLIKC, ex MDBC).

Figs 9-16. Euploea spp. All figures not to scale: upperside left, underside right

[forewing lengths, in mm, in square brackets]. (9-14) E. wallacei: (9, 10, 13, 14) E. w.

melia; (9-10) Dauan I., Torres Strait, Qld, (9) 4, 9.1.2010 [43 mm], (10) 9, 10.1.2010

[40]; (13) 4, Wamar I., Dobo, Aru, 1992 [38]; (14) £, Wau, Morobe Province, PNG,

24.x.1975 [40]; (11-12) E. w. wallacei: (11) 3, Halmahera I., Northern Moluccas, -

.vi.1996 [42]; (12) 4, Manokwari, West Papua Province, 1989 [38]. (15-16) EF.

leucostictos: Dauan I., Torres Strait, Qld, 9.1.2010, (15) 3 [42], (16) 9 [45].

Australian Entomologist, 2016, 43 (2) 59

60 Australian Entomologist, 2016, 43 (2)

Figs 17-20. Euploea stephensii. All figures not to scale: upperside left, underside right

[forewing lengths, in mm, in square brackets]. (17-19) E. s. jamesi: (17) 9, Dauan I.,

Torres Strait, Qld, 9.1.2010 [35 mm]; (18) 9, Jaya Pura, Papua Province, Indonesia,

1989 [36]; (19) d, Janita Village, 10 km E of Popondetta, Oro Province, PNG,

22.x.1987 [30]. (20) E. s. pumila Butler, 1866: 3, Imonda, West Sepik Province,

PNG, 15.v.1992 [30] (courtesy of QDAFC).

Field observations

Two humid mangrove glades (Fig. 7), with large numbers of L. racemosa

flowers, were preferred loci for several species of Euploea, with several

hundred butterflies observed in each glade at any one time, feeding especially

at flowers of several taller L. racemosa trees in each glade. The Euploea

species were observed mostly between the hours of 0800h and 1000h

Australian Eastern Standard Time (AEST) and again from about 1400h to

1600h AEST. Large numbers of Euploea were also observed communally

roosting on branches and dead sticks within mature L. racemosa trees and on

other large mangrove trees Rhizophora stylosa Griff. (Rhizophoraceae) and

Avicennia marina eucalyptifolia (Valeton) J. Everett (Acanthaceae) (Fig. 8).

In addition, hundreds of Euploea were observed communally roosting in

these loci prior to 0800h AEST and it is likely that they roosted there

overnight, prior to feeding from L. racemosa flowers the following morning.

Australian Entomologist, 2016, 43 (2) 61

By the middle of January 2010, after cessation of the mangrove blossom, the

numbers of Euploea declined markedly within the glades, indicating the

aggregation's opportunistic nature and its direct link to the nectar source. In

previous years on Dauan Island (i.e. Januaries of 2006 and 2008), Euploea

species were observed by the author around L. racemosa, but the abundance

of flowers and butterflies and the Euploea species diversity observed in the

two mangrove loci in December 2009 and January 2010 were unprecedented.

In addition, other regular communal roosting haunts of many Euploea

observed that same year and in previous wet seasons have been in the island

village, around and under flowering trees of Terminalia catappa F. and T.

muelleri Benth. (Combretaceae) and Citharexylum spinosum L.

(Verbenaceae), but again not in the large numbers observed in the mangrove

glades in December 2009 and January 2010.

The majority of Euploea species observed in the summer of 2009-2010 were

those that are often seen on Dauan Island, particularly during the wet season,

viz. E. algea amycus, E. sylvester sylvester, E. alcathoe misenus (taxonomy

as per Lambkin 2005, Braby 2016) (Fig. 8) and E. tulliolus dudgeonis

(taxonomy as per Lambkin and Knight 2007). What was unusual in 2009-

2010 was the occurrence of some Euploea species that are not often observed

on the island, viz. E. batesii batesii (taxonomy as per Lambkin 2013, Braby

2016), E. corinna (taxonomy as per Braby 2010), E. netscheri erana and E.

leucostictos (Lambkin and Knight 2007), with the latter two being relatively

abundant (24 and 30 specimens collected respectively) inside and around the

two mangrove loci, either roosting or feeding on nectar of L. racemosa.

Within these mangrove glades, a male and female of E. wallacei melia (Figs

9-10) were collected feeding at L. racemosa blossom in company with E.

leucostictos (Figs 15-16); one was collected in the morning of 9 January (at

approximately 0900h AEST) and the other in the afternoon of 10 January (at

approximately 1500h AEST).

On 9 January, a single female of E. stephensii jamesi (Fig. 17) was collected

in the afternoon (at approximately 1500h AEST), as it perched on an inner

branch of Rhizophora stylosa just beneath blossoming trees of L. racemosa.

The majority of Euploea specimens collected and observed on Dauan Island

during that particular week were in fresh condition, suggesting they were

newly emerged.

Euploea wallacei melia Fruhstorfer, 1904

Euploea wallacei occurs in the Moluccas from Halmahera to Buru and

eastward on Aru, Waigeo, Mysol, Salawati, Biak, Japen, Roon, mainland

New Guinea and several of its outlying islands, including Daru, to Fergusson

Island (Ackery and Vane-Wright 1984, Parsons 1998). Across its range, the

species is phenotypically variable from west to east, with western populations

of E. w. wallacei from the Moluccas and E. w. confusa Butler, 1866 from

62 Australian Entomologist, 2016, 43 (2)

Indonesian Papua being mostly black or dark brown in colour (Figs 11-12).

Further east in Papua New Guinea, as E. w. melia (Figs 13-14), the species

occurs mostly as a black butterfly with conspicuous ‘orange windows’ on the

forewings (Ackery and Vane-Wright 1984). This black and orange form is

similar to the ‘usipetes’ form of E. leucostictos (Figs 15-16), which is another

polymorphic species with a similar black and orange morph found

predominantly in southern Papua New Guinea (Parsons 1998, Lambkin and

Knight 2007). Euploea leucostictos has a much broader range, from the

Andaman Islands and southern China to the Solomon Islands (Yata and

Morishita 1985).

In Papua New Guinea E. wallacei frequents forests and forest margins where

it is often encountered with the ‘usipetes’ form of E. leucostictos (Ackery and

Vane-Wright 1984, Parsons 1998). Little is known of its life history although

D'Abrera (1978) and Parsons (1998) recorded Ficus sp. (Moraceae) and

Parsonsia sp. (Apocynaceae) respectively as larval host plants.

Due to the similarity of E. w. melia and E. leucostictos (form ‘usipetes’), the

latter is thought to be a Batesian mimic of E. w. melia (Parsons 1998).

Euploea w. melia can be distinguished from females of orange forms of E.

leucostictos by the presence of elongate white stripes in the spaces above

1A+2A and CuA, of the forewing underside, and by the presence of a series

of centrally placed blue spots or streaks (3-5) on the hindwing underside

(Figs 9-14). In addition, females of E. leucostictos always bear at least a

single white spot in the subterminal area of the forewing upperside (Fig. 16)

(Lambkin and Knight 2007). Euploea w. melia is one of the few species of

Euploea where the posterior margin of the male forewing is straight rather

than bowed or convex (Ackery and Vane-Wright 1984). The species was

placed into one of three clades (i.e. Clade 211.3) but, in doing so, Ackery and

Vane-Wright (1984) suggested that this arrangement was at best tentative, as

E. wallacei was the only species in this clade where the posterior margin of

the male forewing was not strongly convex.

Despite the widespread distribution of E. wallacei across New Guinea

(Ackery and Vane-Wright 1984, Parsons 1998) and the proximity of Torres

Strait to southern Papua New Guinea, the species has not been recorded

previously from the northern Torres Strait islands.

Euploea stephensii jamesi (Butler, 1876)

Euploea stephensii primarily occurs in mainland New Guinea but does

extend westwards to the Maluku Islands in the Northern Moluccas of

Indonesia (and including Waigeo and Yapen Islands) and eastwards to New

Britain and the Bismarck Archipelago in Papua New Guinea (Ackery and

Vane-Wright 1984, Parsons 1998). In Papua New Guinea the species is

widespread (Parson 1998) and frequents forest margins, including secondary

forest up to about 1000 m (Ackery and Vane-Wright 1984, Parsons 1998).

Australian Entomologist, 2016, 43 (2) 63

Parsons (1998) briefly studied the life history of E. s. jamesi in Bulolo,

Morobe Province, Papua New Guinea, where he reared larvae on Trophis

scandens (Lour. Hook. & Arn. (Moraceae). Although Parsons (1998)

illustrated a final instar larva, the image was unfortunately referred to twice

in his text as either E. tulliolus dudgeonis or E. s. jamesi and referred to as E.

tulliolus (Fabricius, 1793) in the plate caption. Despite Parsons' (1998)

illustration resembling closely the final instar larva of E. t. tulliolus from

Brisbane, Queensland (Lambkin 2010), it might well have been of E. s.

jamesi, considering that the two species are thought to be closely related

(Ackery and Vane-Wright 1984).

Euploea stephensii is also phenotypically variable across its range and has a

large number of subspecies (Parsons 1998). In the northern half of Papua

New Guinea, individuals of E. stephensii can be remarkably different in

colour (Fig. 20), with some females being a pale blue-silver on the upperside

of the wings (E. s. pumila f. lucinda Grose-Smith, 1894). In the south of

Papua New Guinea, individuals of E. s. jamesi show little polymorphism

(Figs 18-19) and resemble EF. tulliolus, but can be distinguished from the

latter by the shape of their forewings, being more squat and vertically broader

(Parsons 1998). This similarity prompted Ackery and Vane-Wright (1984) to

group these two species, plus E. hewitsoni Felder & Felder, [1865] and E.

darchia (Macleay, 1827), into a ‘tulliolus-complex’, which they tentatively

classed as a ‘clade’ [or more correctly as an ‘informal group’ as per the

International Commission on Zoological Nomenclature ‘Code’ (ICZN

1999)]. Ackery and Vane-Wright (1984) admitted that their assemblage of

the four taxa into this complex was poorly characterised and that the only

significant feature of the group they could determine was its ‘unique

exploitation’ of T. scandens as a larval host plant.

Finally, populations of E. wallacei, E. leucostictos and E. stephensii all show

diverse phenotypic variation across their geographic ranges (Ackery and

Vane-Wright 1984, Parsons 1998, Lambkin and Knight 2007). This,

combined with the difficulty in defining their cladistic placement (Ackery

and Vane-Wright 1984), suggests that these taxa could be further examples of

species complexes within the genus Euploea, thereby highlighting the

difficulties confronted in attempting to unravel these complexes (e.g. Ackery

and Vane-Wright 1984, De Baar 1991, Vane-Wright 1993, Lambkin 2001,

2005, 2013, Lambkin and Knight 2007, Monastyrskii 2011, Tennent 2001,

Treadaway 2012, Yata and Morishita 1985).

Discussion

One of the features of Euploea butterflies in Torres Strait, particularly on

Dauan Island, is their often relative abundance soon after the commencement

of the monsoon rains and their relative scarcity at other times, chiefly during

the dry season. Thus, after the end of the wet season, numbers of Euploea

64 Australian Entomologist, 2016, 43 (2)

noticeably decline on Dauan Island and those surviving enter a reproductive

diapause (Canzano et al. 2003).

Euploea species in the monsoon tropics can typically enter diapause during

the dry season and are known to communally aggregate or roost, quite often

in large numbers, in protected areas such as shaded gullies, caves and under

evergreen thickets of mangroves and bamboo, most often adjacent to water

(Kitching and Zalucki 1981, Monteith 1972, 1982, Scheermeyer 1993,

Canzano et al. 2003).

With the onset of the wet season across Torres Strait and southern Papua

New Guinea come conditions that likely act as cues to end this diapause

(Canzano et al. 2003), resulting in adult roosting butterflies becoming active

and reproductive. Consequently, these individuals that have undergone

diapause give rise to the next generation of Euploea, observed on Dauan

Island as a peak during the first month of the wet season (Lambkin and

Knight 2007). Based on life table data for several other Australian Euploea

species (Meyer 1996, 1997, Lambkin 2001, 2010, Braby 2009) and the fact

that diapausing adults become reproductive again after the start of the wet

season (Canzano et al. 2003), it is estimated that these newly emerged adults

occurring during the early part of the wet season originated from eggs laid by

diapausing mothers approximately 3-4 weeks earlier.

Reproductive diapause in Euploea in Australia is believed to be a significant

survival strategy for the dry season in the monsoon tropics (Monteith 1982,

Canzano et al. 2003), as early instar larvae of many Euploea species feed

solely on fresh growth of their host plants (Clarke and Zalucki 2000, Rahman

and Zalucki 1999), this being unavailable through the drier months (Braby

2000, Canzano et al. 2003). In addition, suitable nectar sources for adult

butterflies are more readily available during the wet season, although it is

thought that some dry season nectar sources can sustain these large

populations during dry periods (Canzano et al. 2003).

On Dauan Island, the majority of specimens of Euploea observed at the very

start of the wet season in November and early December were noted by

Lambkin and Knight (2007) to be in a worn condition. This observation

supports the theory that individuals have persisted by perhaps diapausing in

aggregations on the island since the termination of the previous wet season.

While long-term, dry season aggregations of Euploea are documented in

Australia (Kitching and Zalucki 1981, Monteith 1972, 1982, Valentine 1987,

Scheermeyer 1993, Canzano et al. 2003), there are only brief mentions in the

literature of other forms of communal roosting of Euploea species in

Australia (Meyer 1996, 1997, Braby 2000, Lambkin and Knight 2007) and

overseas, viz. E. mulciber (Cramer, 1777) in Taiwan (Yata and Morishita

1985) and E. midamus (Linnaeus, 1758) and E. core (Cramer, [1780]) in

Hong Kong (Ackery and Vane-Wright 1984, Bascombe et al. 1999). One

Australian Entomologist, 2016, 43 (2) 65

report in Bascombe et al. (1999) similarly described E. midamus roosting

communally at nectar sources of Zanthoxylum avicennae (Lam.) DC

(Rutaceae). Despite the proclivity for Euploea in the monsoon tropics to roost

during the dry season in Australia (Monteith 1972, 1982, Canzano et al.

2003), communal roosting by Euploea at nectar sources is previously

unrecorded in Australia.

In America, nectar source roosting is known during migrations of Danaus

plexippus (Linnaeus, 1758) (Nymphalidae: Danainae), where the butterflies

pause along the way throughout their journey to imbibe on nectar. These

opportunistic stopover sites are believed to be an important resource for

successful migrations of D. plexippus (Brower et al. 2015).

While not strictly analogous to D. plexippus migrations, the phenomenon

observed on Dauan Island in 2009-2010 could indicate that Euploea species

can migrate out of their breeding environments en masse and are attracted

perhaps from remote areas and arrested by discrete nectar-rich loci, such as in

the case recorded here in mangrove glades of profusely flowering L.

racemosa trees. If this is the case, then these temporary aggregations of

Euploea might be migrations possibly originating from the nearby Papua

New Guinea mainland, approximately 10 km to the north of Dauan Island.

Moreover, crow butterflies were observed in January 2014, in numbers,

flying in a northerly direction over the stretch of water that separates Saibai

and Dauan Islands (pers. obs.). This observation does show that Euploea

butterflies are vagile, at least between neighbouring islands.

The synchronization of several events which occurred in the early wet season

of December 2009-January 2010 might have contributed to what appears to

have been an atypical aggregation (when compared with other years), when

Euploea butterflies were overall unusually common, especially as roosting

individuals. This applied to species that are normally infrequently observed

on Dauan Island (e.g. E. leucostictos and E. netscheri erana) and may further

explain the occurrence of the two previously unrecorded species E. w. melia

and E. s. jamesi.

Acknowledgements

I thank the local community councils and island Elders of Dauan Island,

Torres Strait for allowing entry into their community and providing

assistance and lodging during time spent on their island. Appreciation is

given to M.P. Zalucki (University of Queensland) for his critical review of

the manuscript, and to J.S. Bartlett (ODAFC) for allowing access to a

specimen in his care and for the image of this specimen. In addition, the late

M. De Baar (vale 6 January 2011) is thanked for his extensive collection of

Euploea species which was made available to the author on a permanent

basis in 2011. This collection has been of significant assistance to the author

over the period of his research on the genus Euploea in Torres Strait. This

66 Australian Entomologist, 2016, 43 (2)

paper partially fulfils the requirements for a Master of Philosophy degree

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

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Australian Entomologist, 2016, 43 (2): 69-74 69

A NEW SPECIES OF THE TACHYS (S. L.) ECTROMOIDES-GROUP

FROM SOUTHEASTERN QUEENSLAND, AUSTRALIA

(COLEOPTERA: CARABIDAE: BEMBIDIINI)

MARTIN BAEHR

Zoologische Staatssammlung, Münchhausenstr.21, D-81247 Munich, Germany

(Email: martin.baehr@zsm.mwn.de )

Abstract

Tachys federicae, sp. n., in the enigmatic ectromoides species-group (sensu Baehr 1989) of the

genus Tachys Dejean, 1825 (sensu lato), is described from mountains near Brsbane in

southeastern Queensland, Australia. The species is distinguished from all other species of the

ectromoides-group by its striking bicolouration and narrow pronotum. A revised key to all

species of the group is included.

Introduction

In Australia, the carabid genus Tachys Dejean, 1825, in the widest sense,

includes a number of species that are not easily included in one of the

generally accepted genera (or subgenera, according to the opinion of the

respective workers). Thus, for the present, they are treated as species-groups

with doubtful relationships. One of these is the ectromoides-group (sensu

Baehr 1989), which includes about half a dozen species that are characterized

by: presence of two deep foveae in the mentum; a very distinct, elongate,

straight lateral carina at the basis, and a deeply impressed, wide basal part of

the median line on the pronotum; laterad relatively little produced eye; and

very glossy and even slightly sericeous surface of the elytra. The species-

group is distributed mainly in eastern Australia from New South Wales to

Windsor Tableland in northern Queensland, with a single species occurring in

the southwestern part of Western Australia (Sloane 1896, Darlington 1962,

Baehr 1989, 1991, 2003).

Species of the group are rarely collected, perhaps because they appear to live

on or under the bark of trees or in leaf litter. They occur in rain forests as well

as in open forests and woodlands. However, arboricolous habits are not

verified for all species.

Among a number of carabid beetles sent for identification from the

Queensland Museum, Brisbane, was a single specimen of another species of

this enigmatic ectromoides-group that is described in the present paper. The

single female specimen of the new species is outstanding within the

ectromoides-group in its large body size, narrow pronotum and contrasting

colouration, making its description justified.

Materials and methods

Measurements were taken using a stereo microscope with an ocular

micrometer. Body length was measured from apex of labrum to apex of

elytra, length of pronotum along midline, length of elytra from the most

advanced part of the humerus to the very apex.

70 Australian Entomologist, 2016, 43 (2)

For dissection of the female genitalia the specimen was softened for a night

in a jar under moist atmosphere, then the genitalia were removed and

subsequently cleaned for a short while in hot KOH. The habitus photograph

was obtained with a digital camera using AutoMontage and subsequently was

worked with Corel Photo Paint X4.

The holotype of the new species is deposited in the Queensland Museum,

Brisbane (QM).

Tachys federicae sp. n.

(Figs 1-2)

Type material. Holotype 9, QUEENSLAND: 27.295°S 152.749?E, Mt.Tenison

Woods, 631 m, 29 Oct 2009, 19171, G.B. Monteith & F. Turco, sieved litter (QM

Reg. No. QMT224214).

Diagnosis. Characterized and distinguished from all other species of the

group by combination of large body size (in group), narrow prothorax,

distinct, isodiametric microreticulation of pronotum, conspicuously crenulate

elytral striae, and the contrasting colour with red head and pronotum and

almost black elytra.

Figs 1-2. Tachys federicae sp. n.: (1) dorsal view of holotype female, length: 3.65

mm; (2) female gonocoxites (scale bar: 0.1 mm).

Australian Entomologist, 2016, 43 (2) 71

Description. Measurements. Length: 3.65 mm; width: 1.55 mm. Ratios:

width/length of pronotum: 1.36; width base/apex of pronotum: 1.38;

length/width of elytra: 1.49; width elytra/pronotum: 1.63.

Colour (Fig. 1). Head reddish, frons in middle and occiput reddish piceous;

pronotum red; elytra almost black, only the very basal margin slightly lighter,

also the lateral channel inconspicuously paler. Labrum and mandibles dark

red, palpi and antennae dirty yellow, antenna apicad slightly paler. Legs dirty

yellow. Whole lower surface pale red. Surface of elytra slightly iridescent.

Head (Fig. 1). Moderately wide. Eye rather depressed, orbit large, oblique-

convex, length of orbit almost half of length of eye. Temporal sulcus

elongate, deep, straight, slightly oblique, laterally bordered by a conspicuous

ridge, anteriorly prolonged onto clypeus, posteriorly almost attaining level of

posterior border of eye. Anterior margins of clypeus and labrum straight.

Mandible moderately elongate. Dorsal aspect of head markedly trechine-like.

Mentum bifoveate, mental tooth present, though wide and rather obtuse at tip.

Both palpi densely setose. Terminal palpomeres of both palpi elongate,

markedly subulate. Antenna medium-sized, surpassing base of pronotum by

about two antennomeres. Median antennomeres about 1.75 x as long as wide.

Frons with conspicuous isodiametric microreticulation, microsculpture

weaker on clypeus and on neck, meshes there slightly transverse.

Pronotum (Fig. 1). Comparatively narrow in group. Surface in middle

slightly convex. Anterior angle well produced, obtuse at apex, lateral margin

in apical third markedly curved inwards. Apex with moderately deep

excision. Pronotum widest at about middle, behind anterior lateral seta,

moderately and almost straight narrowed towards base, lateral margin slightly

concave only near base. Basal angle right. Base wide, considerably wider

than apex, very slightly produced in middle. Lateral channel anteriorly

narrow, slightly widened towards base, neither apex nor base bordered.

Anterior transverse sulcus shallow, forming a triangle. Median line shallow,

not attaining the apex, suddenly widened and deepened near base to form a

conspicuous longitudinal sulcus. Prebasal transverse sulcus deep, straight,

interrupted in middle. Submarginal carina conspicuous, elongate, very

slightly oblique. Basal grooves large and very deep. Microsculpture on disk

fine though distinct, slightly superficial, consisting of fine, isodiametric to

slightly transverse meshes; in front of anterior transverse sulcus and across

base behind basal transverse sulcus microreticulation strong and regularly

isodiametric. Surface of disk moderately glossy, without any perceptible

punctures.

Elytra (Fig. 1). Comparatively elongate, but wide in comparison to prothorax,

dorsally convex but on disk depressed, with wide base, moderately convex

laterally, widest slightly behind middle. Humerus rounded. Lateral channel

deep and fairly wide. Scutellary striole absent, though scutellary puncture and

72 Australian Entomologist, 2016, 43 (2)

seta present and conspicuous. Striae complete, deeply impressed almost

throughout, lateral striae little shallower, only close to apex all striae are

slightly shallower. Striae at least in anterior half distinctly punctate or even

crenulate. Intervals very slightly raised. 8" stria deeply sulcate in apical half,

very fine in basal half, reaching the humeral group of marginal pores.

Recurrent stria deeply impressed, bearing a strong ridge posterolaterally,

uniting with the 3" stria. No discal punctures present, although with a

setiferous pore within the recurrent stria close to apex. 8 marginal setae

present, arranged in three groups of 4 pores near base, 2 behind middle, and 2

near apex, all punctures large and deep and the setae very elongate.

Extremely fine and superficial transverse lines only visible at very high

magnification (> 150 x), at lower magnification surface appearing glossy and

rather iridescent. Metathoracic wings present.

Lower surface. Metepisternum slightly < 2 x as long as wide. All thoracic

sterna with several erect, elongate hairs. Abdominal sterna with sparse, much

shorter pilosity. Female terminal abdominal sternum quadrisetose.

Legs. Of average size. Tibiae and tarsi with dense and very elongate setosity,

claws edentate. Squamosity of male anterior tarsus unknown.

Male genitalia. Unknown.

Female gonocoxites (Fig. 2). Apex of gonocoxite 1 without setae. Gonocoxite

2 narrow and elongate, slightly curved, with acute apex; with one rather small

dorsomedian ensiform seta located slightly apicad of middle, two narrow but

rather elongate, widely separated ventrolateral ensiform setae situated in

basal half, and one elongate nematiform seta that originates from a circular

pit near apex.

Variation. Unknown.

Etymology. The species name is a patronym in honour of one of the

collectors, Federica Turco of Queensland Museum (now at Australian

National Insect Collection, Canberra).

Distribution. Southeastern Queensland. Known only from the type locality.

Collecting circumstances. The holotype was collected by sieving litter in

subtropical rainforest at medium altitude.

Relationships. This is a unique species within the ectromoides-group. It may

be more closely related to T. fortestriatus Baehr, 2003, which is also from

southeastern Queensland, than to any other species but, without knowledge of

the male genitalia, this is speculative.

Key to species of Tachys ectromoides-group

For better distinction the key from the most recent paper on the group (Baehr

2003) is revised as follows.

Australian Entomologist, 2016, 43 (2) 73

] Elytra with distinct colour pattern; discal elytral punctures present ......... 2

— Elytra unicolourous, or with very indistinct pattern (base ill-delimited

paler); discal elytral punctures absent ........................... esee 3

2 Elytra yellow with wide brown fascia and piceous apex; antenna yellow

throughout; pronotum very wide, base almost as wide as width of

pronotum in middle; surface conspicuously reticulate. Eastern Australia

PIEN ATA EN EAEE URINE T. ectromoides Sloane, 1896

— Elytra piceous with indistinct lighter spots at humeri and in posterior

third; antenna piceous with 1"', 2" and base of 3™ antennomeres yellow;

pronotum evidently narrowed to base; surface almost smooth, nitid.

Southwestern Western Australia ............................... T. marri Baehr, 1989

3 Larger species, length > 3.2 mm; either median antennomeres c. 3 x as

long as wide, or head and pronotum red and elytra contrastingly dark .... 4

— Smaller species, length 2.6-2.9 mm; median antennomeres « 1.5 x as long

as wide; colour not as contrasting — ...............eessseseeeeeeeeeeere nennen 5

4 Colour rather uniformly reddish piceous to piceous; median antennomeres

c. 3 x as long as wide. Northeastern New South Wales .............................

RIDMANS GT n bechs MIMME IEEE LI oA eEE cer eck eres 32.499 900 T. bolus Darlington, 1962

— Colour contrasting: head and pronotum red, elytra dark (Fig. 1); median

antennomeres < 2 x as long as wide. Southeastern Queensland .................

EEEE EE eties eene hue ENS 22722 A Botdcecce Mae he fea caedes T. federicae sp. n.

5 Pronotum narrower, ratio width/length < 1.35; elytra wider in comparison

with pronotum, ratio width elytra/pronotum > 1.5, elytra more oval-

shaped, lateral margins more convex .................sesseseeeeeeeeeeeeee eene 6

— Pronotum wider, ratio width/length c. 1.44; elytra narrower in comparison

with pronotum, ratio width elytra/pronotum « 1.45, elytra less oval-

shaped, lateral margins more parallel-sided. Southeastern Queensland

TTE REIR UP CH RII ASTE TEE Maar T. fortestriatus Baehr, 2003

6 Head, pronotum, and base of elytra reddish, rest light piceous; base of

pronotum wider, ratio base/apex > 1.4; striae less impressed, even basally

barely crenulate, lateral striae feebly indicated. Eastern New South Wales,

southeastern Queensland ............................... T. bolellus Darlington, 1962

— Colour almost black, only pronotum dark piceous; base of pronotum

narrower, ratio base/apex « 1.3; striae well impressed, distinctly crenulate

in basal half, lateral striae well marked. Northeastern Queensland ............

VP ETE EIEEE ET SE PE REATO cas HER A tA T. windsorensis Baehr, 1991

Discussion

Unfortunately, the systematic position of the ectromoides-group within the

‘genus’ Tachys sensu lato is still uncertain, similar to the systematic position

74 Australian Entomologist, 2016, 43 (2)

of a number of other Australian tachyine species and species-groups of

different shape and structure. Although a number of workers in more recent

times have tried to elucidate the generic and subgeneric taxonomy of the

former genus ‘Tachys sensu latissimo’, this is still remarkably controversial.

Even the application of certain well recognizable and generally accepted

genera to the Australian Tachyina leaves a number of Australian species

unplaced, because presently they cannot be included with good reasons in

one of these genera. The species that are combined in the ectromoides-group

form one of these groups of uncertain status.

All species of the ectromoides-group are apparently rare, perhaps because

they have not been searched for in their characteristic habitats. Indeed, almost

all specimens that bear records of their collecting circumstances, apart from

those discovered by the author, have been collected at light or in flight

intercept traps, which implies that these records are from specimens on the

wing. The record of the new species described herein adds another sampling

circumstance, namely sieving from leaf litter. Due to the apparent rarity of all

species, males of some species are so far unknown. Therefore, comparisons

of the male aedeagi of all species are not yet possible, making the

relationships within the ectromoides-group still unclear.

Acknowledgement

My thanks for the kind loan of the new species are due to Dr Geoff Monteith

of Queensland Museum, Brisbane.

References

BAEHR, M. 1989. A new species of the Tachys ectromoides-group from Western Australia

(Coleoptera, Carabidae, Bembidiinae). Spixiana 12: 279-283.

BAEHR, M. 1991. Tachys windsorensis, spec. nov. from North Queensland, a further new

species of the Tachys ectromoides-group (Insecta, Coleoptera, Carabidae, Bembidiinae).

Spixiana 14: 189-192.

BAEHR, M. 2003. A further new species of the Tachys (s.l) ectromoides-group from

Queensland, Australia (Insecta, Coleoptera, Carabidae, Bembidiinae). Spixiana 26: 143-147.

DARLINGTON, P.J. Jr. 1962. Australian carabid beetles XI. Some Tachys. Psyche, Cambridge

69: 117-128.

SLOANE, T.G. 1896. On the Australian Bembidiides, referable to the genus Tachys with the

description of an allied genus Pyrrhotachys. Proceedings of the Linnean Society of New South

Wales 21: 355-377.

Australian Entomologist, 2016, 43 (2): 75-82 75

A REVIEW OF THE SUBGENUS AUSTRODACUS PERKINS OF

BACTROCERA MACQUART (DIPTERA: TEPHRITIDAE: DACINAE)

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

'8/3 McPherson Close, Edge Hill, Cairns, Qld 4870

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

Abstract

The Bactrocera Macquart subgenus Austrodacus Perkins is reviewed and 5 species recorded

from Australia and New Guinea are included: B. abdoaurantiaca Drew, B. alampeta Drew, B.

atrisetosa (Perkins), B. cucumis (French) and B. papuaensis (Malloch), comb. n. (2 unichromata

Drew, syn. n.). Subgenus Hemiparatridacus Drew is placed as a new synonym of subgenus

Austrodacus and its sole species, B. abdoaurantiaca, is transferred, while B. alampeta, B.

atrisetosa and B. papuaensis are transferred to Austrodacus from subgenus Paratridacus Shiraki.

A fourth Papua New Guinean species previously included in Paratridacus, B. mesonotaitha

Drew, is transferred to subgenus Zeugodacus Hendel as a close ally of B. (Z.) sandaricina Drew.

Introduction

This is the third paper in a series reviewing the subgenera of the

economically important fruit fly genus Bactrocera Macquart, made possible

by the revisions of Australasian and Southeast Asian species by Drew (1989)

and Drew and Romig (2013) respectively. Previous papers have reviewed the

Indo-Australian subgenera Calodacus Hancock and Parazeugodacus Shiraki

(Hancock 2015, Hancock and Drew 2015). This paper deals with subgenus

Austrodacus Perkins, which is considered here to contain five described

species known only from Indonesia's Papua Province, mainland Papua New

Guinea and eastern and northern Australia. The recent elevation of the

Zeugodacus group of subgenera to generic status (Virgilio et al. 2015, De

Meyer et al. 2015) is not accepted here and we follow Drew and Romig

(2013) and Hancock and Drew (2015) in retaining all species in genus

Bactrocera. A preliminary statement of our reasons is given in Hancock and

Drew (2015) and a comprehensive case is being prepared.

Genus Bactrocera Macquart

Subgenus Austrodacus Perkins

Austrodacus Perkins, 1937: 56. Type species Dacus cucumis French, 1907, by

original designation.

Bactrocera (Hemiparatridacus) Drew, 1989: 15. Type species Bactrocera

abdoaurantiaca Drew, 1989, by original designation. Syn. n.

Definition. Abdominal sternite V of male with a shallow posterior

emargination; posterior lobe of male surstylus narrow and elongate; pecten of

cilia absent on abdominal tergite III of male; postpronotal setae absent; supra-

alar setae present or absent; prescutellar acrostichal setae present or absent;

two pairs of scutellar setae; scutum with medial postsutural yellow vitta

present and lateral postsutural yellow vittae extending across suture

anteriorly; wing with costal band narrow and not expanding apically; apex of

aculeus very weakly trilobed [subapically ‘keeled’ ].

76 Australian Entomologist, 2016, 43 (2)

Response to male lures. None known for any member of the subgenus. Single

records of B. alampeta and ‘B. unichromata’ from methyl eugenol (Drew

1989) appear to be accidental and have not been repeated.

Included species. B. (A.) abdoaurantiaca Drew, B. (A.) alampeta Drew, B.

(A.) atrisetosa (Perkins), B. (A.) cucumis (French) and B. (A.) papuaensis

(Malloch) (= B. unichromata Drew, syn. n.).

Host plants. Recorded from the fruit of Cucurbitaceae and Solanaceae, with

occasional records from other families (Drew 1989, Hancock et al. 2000).

Comments. The combination of a shallow emargination to sternite V and long

posterior surstylus lobes place this subgenus in the Zeugodacus group of

subgenera as defined by Drew (1989). Although three of the included species

were included previously in subgenus Paratridacus Shiraki (e.g. by Drew

1989), that subgenus has the surstylus lobes shorter and much broader (Hardy

1974 and Fig. 1) than the narrow, elongate lobes seen in Austrodacus.

Paratridacus s.s. also lacks a yellow medial vitta on the scutum, has non-

cucurbitaceous host plants (Garcinia spp: Clusiaceae) and appears to be

better placed in the Melanodacus group of subgenera, as suggested by

Hancock and Drew (2015), with molecular evidence (Krosch et al. 2012)

supporting this association. The weakly trilobed shape of the aculeus apex

appears to be a synapomorphy for subgenus Austrodacus. Although the

aculeus has not been examined in B. abdoaurantiaca or B. alampeta other

characters, including the lack of a pecten on abdominal tergite III, very long

posterior surstylus lobes (Fig. 2) and their overall appearance, suggest that

they also belong in Austrodacus.

1 2

Figs 1-2. Bactrocera spp, male surstylus lobes: (1) B. (Paratridacus) expandens

(Walker); (2) B. (Austrodacus) abdoaurantiaca Drew.

The only other Zeugodacus-group species known from the island of New

Guinea or Australia that lacks a pecten of cilia on abdominal tergite III in

males is B. (Niuginidacus) singularis Drew from Morobe Province, Papua

New Guinea. However, that species has no lateral postsutural yellow vittae,

only one pair of scutellar setae, a fuscous discal marking on the wing, an

elongate-oval abdomen and males that respond to cue-lure. It is also smaller

in size (wing length ca 5 mm) and, in lacking prescutellar acrostichal setae,

Australian Entomologist, 2016, 43 (2) TI

appears to be most closely related to several New Guinean species currently

included in subgenus Sinodacus Zia.

Included species

For detailed morphological descriptions and illustrations see Drew (1989).

B. (Austrodacus) abdoaurantiaca Drew

Bactrocera (Hemiparatridacus) abdoaurantiaca Drew, 1989: 188. Type locality

Aiyura, Papua New Guinea.

Distribution. Eastern Highlands Province, Papua New Guinea.

Host plants. Unknown; swept from Cucurbita sp. (Cucurbitaceae) (1 3, Uni

of Goroka gardens, Goroka, 24.vi.2011, A.D. Rice, in Department of

Agriculture Collection, Cairns) and this is a likely host plant.

Comments. As noted above, the overall morphology of this species suggests a

close relationship with other species of Austrodacus and the monotypic

subgenus Hemiparatridacus is thus regarded as a synonym. It differs from

other Austrodacus species only in having the combination of supra-alar setae

present and prescutellar acrostichal setae absent and appears closest to B.

alampeta. The abdominal setulae are relatively short and white, with a weak

gold reflection on tergites III-V. The above specimen from Goroka has

fuscous abdominal markings as in B. alampeta and faint facial spots.

B. (Austrodacus) alampeta Drew

Bactrocera (Paratridacus) alampeta Drew, 1989: 196. Type locality Mt Hagen,

Papua New Guinea.

Bactrocera (Austrodacus) alampeta Drew: Hancock and Drew 2015: 101.

Distribution. Western Highlands Province, Papua New Guinea.

Host plants. Unknown.

Comments. This species appears closest to B. abdoaurantiaca, sharing with it

a largely black scutum and differing primarily in the presence of both supra-

alar and prescutellar acrostichal setae. The abdominal setulae are relatively

short and white, with a weak gold reflection on tergites III-V.

B. (Austrodacus) atrisetosa (Perkins)

Zeugodacus atrisetosus Perkins, 1939: 29. Type locality Mondo, Papua New Guinea.

Dacus (Zeugodacus) cucumis: Malloch 1939: 412. Misidentification.

Melanodacus rubidus May, 1957: 297. Type locality Goroka, Papua New Guinea.

Syn. May 1962: 64.

Melanodacus atrisetosus (Perkins): May 1962: 64.

Dacus (Paratridacus) atrisetosus (Perkins): Drew 1973: 15.

Bactrocera (Paratridacus) atrisetosa (Perkins): Drew 1989: 197.

Bactrocera (Austrodacus) atrisetosa (Perkins): Hancock and Drew, 2015: 101.

78 Australian Entomologist, 2016, 43 (2)

Types. Although Drew (1989) and Norrbom et al. (1999) referred only to

syntype material for B. atrisetosa, Perkins (1939) stated: ‘Type returned to

British Museum', indicating designation of a single specimen as the holotype,

but he did not state to which of the three listed specimens it referred. Two

females from the type series are present in the Natural History Museum,

London (BMNH: examined), labelled as follows:

HOLOTYPE 9: Papua, Mondo, 5000’, 11.1934, L.E. Cheesman, BM 1934-321 / (red

label) Holotype Dacus atrisetosus Perkins [in BMNH].

PARATYPE 9: Papua, Mafulu, 4000’, 1.1934, L.E. Cheesman, BM 1934-321 / (blue

label) Paratype Dacus atrisetosus Perkins [in BMNH].

The action of Perkins in nominating a single “Type’ fulfils Section 73.1.1. of

the International Code of Zoological Nomenclature (ICZN 1999) for original

designation of a holotype and therefore we accept the above specimen so

labelled as the holotype. Perkins’ third specimen, from Mt Lamington, Oro

Province, although apparently seen by May (1962), has not been found in the

collections of either the Queensland Museum [the current repository of

Perkins' retained specimens] or the Queensland Department of Agriculture

and Fisheries [where May worked] (S. Wright and D. Tree pers. comms) and

is presumed lost. Consequently, its identity has not been confirmed.

Distribution. Recorded from 1200-1650 m in Central, Oro, Morobe and

Eastern Highlands Provinces in Papua New Guinea and Papua Province in

Indonesia. Newly recorded from Vagau, Herzog Mts, ca 4000', Morobe

Province, Papua New Guinea, 9-17.1.1965 (7 SQ, in BMNH) and ‘Tanah

Merah’ [above 1600 m] via Wamena, Papua Province, Indonesia, ‘3°51.95'S,

138°42.44'F’, 6.vii1.1997, G. Bellis (1 9, in Department of Agriculture

Collection, Cairns). A record from Queensland, Australia (Perkins and May

1949) is a misidentification of B. (Hemizeugodacus) aglaiae (Hardy). Dacus

papuaensis Malloch, placed as a synonym of B. atrisetosa by May (1962), is

regarded here as a separate species (see below).

Host plants. Recorded from Citrullus lanatus, Cucumis melo, Cucumis

sativus, Cucurbita pepo, Luffa cylindrica and Momordica charantia (all

Cucurbitaceae) and Lycopersicon esculentum (Solanaceae) (Drew 1989,

Leblanc et al. 2012). The type specimens of ‘M. rubidus’ were collected on

flowers of Euphorbia pulcherrima (Euphorbiaceae) (May 1957) but this is

unlikely to be a host plant. Records from Aglaia sapindina (Meliaceae)

(Drew 1989, Leblanc et al. 2012) are based on a misidentification of B.

aglaiae from Queensland by Perkins and May (1949) before the latter species

was described by Hardy (1951).

Comments. Although B. atrisetosa was referred to subgenus Paratridacus by

Drew (1972, 1989), this was done solely on the presence of supra-alar and

prescutellar acrostichal setae. However, its overall morphology clearly

associates it with B. cucumis rather than with typical members of subgenus

Australian Entomologist, 2016, 43 (2) 79

Paratridacus, particularly the medial yellow vitta on the scutum and the

shape of the male surstylus and female aculeus. The setulose and weakly

trilobed ['keeled'] apex of the aculeus is very similar to that of B. cucumis

(see Drew 1989, figs 335 and 357). The scutum and abdomen frequently have

fuscous markings that vary in extent but unmarked specimens also occur. The

abdominal setulae are long and white, with a gold reflection on tergites III-V.

Malloch's (1939) female B. cucumis from Mondo, Papua New Guinea, is a

misidentification of B. atrisetosa (specimen in BMNH: examined).

B. (Austrodacus) cucumis (French)

Dacus tryoni var. cucumis French, 1907: 307. Type locality Bowen, Queensland.

Dacus cucumis French: Froggatt 1910: 866.

Austrodacus cucumis (French): Perkins 1937: 56.

Dacus (Austrodacus) cucumis French: Hardy 1951: 122.

Bactrocera (Austrodacus) cucumis (French): Drew 1989: 185.

Distribution. Queensland (including Torres Strait islands), Northern Territory

and northern New South Wales, Australia. A record from Mondo in Papua

New Guinea (Malloch 1939) is a misidentification of B. atrisetosa.

Host plants. Recorded primarily from several species of Cucurbitaceae and

Solanaceae (Drew 1989, Hancock et al. 2000). Carica papaya (Caricaceae)

and Passiflora spp (Passifloraceae) are regarded as moderate hosts, while

records from several other families appear to be incidental and in need of

confirmation (Hancock et al. 2000).

Comments. This, the type species of Austrodacus, differs from all other

species in the subgenus in lacking both supra-alar and prescutellar acrostichal

setae. The costal cells are pale fulvous and the abdominal setulae are long

and white with a gold reflection, as in B. atrisetosa; in these two species the

silvery white pruinose areas apically on tergite II and laterally over the

shining spots on tergite V are also more distinct than in all the other species

(C. Lambkin pers. comm.). It is regarded as an economically important pest

of cucurbits and tomatoes.

B. (Austrodacus) papuaensis (Malloch), comb. n.

Dacus (Zeugodacus) papuaensis Malloch, 1939: 412. Type locality Bulolo, Papua

New Guinea.

Bactrocera (Paratridacus) unichromata Drew, 1989: 200. Type locality 20 km SE of

Port Moresby, Papua New Guinea; syn. n.

Bactrocera (Austrodacus) unichromata Drew: Hancock and Drew 2015: 101.

Distribution. Below 1250 m in East Sepik, Morobe and Central Provinces,

Papua New Guinea (Malloch 1939, Drew 1989); also in northeastern Papua

Province, Indonesia (White and Evenhuis 1999). Additionally recorded from

Moale Plantation, Wau, Morobe Province, 3800', xii.68-v.69, Mrs J.E.

80 Australian Entomologist, 2016, 43 (2)

Benson (1 9, in BMNH); Kapakapa [Gabagaba, coast ca 50 km SE of Port

Moresby], Central Province, 1891, L. Loria (1 £, 1 9, in BMNH); and Laloki

Research Station [ca 20 km E of Port Moresby], Central Province,

2.viii.2000, per D. Tenakanai, [reared] ex Luffa cylindrica (2 63,9 99, in

Department of Agriculture Collection, Cairns).

Host plants. Reared from Luffa cylindrica (Cucurbitaceae) [new record].

Comments. As with B. atrisetosa, the morphology of this species clearly

associates it with B. cucumis rather than with typical members of subgenus

Paratridacus, particularly the medial yellow vitta on the scutum, the weakly

trilobed aculeus and the long posterior surstylus lobe (ca 1.5 times as long as

width of surstylus; cf. B. cucumis: Hardy 1951, fig. 3a). Unlike B. cucumis,

however, both supra-alar and prescutellar acrostichal setae are present. It

most resembles pale forms of B. atrisetosa, differing in the denser, long

yellow setulae on abdominal tergites II-V, pale third antennal segment and

colourless costal cell c; it also occurs primarily at lower altitudes.

Although B. papuaensis was previously treated as a synonym of B. atrisetosa

(e.g. May 1962, Drew 1989, Norrbom ef al. 1999), this was based on

presumed tenerality (May 1962). However, examined photographs of the

[non-teneral] holotype male (in Australian Museum, Sydney) indicate a pale

thorax without fuscous markings, a pale third antennal segment, colourless

costal cells, dense and yellow abdominal setulae and a long, finger-like

posterior surstylus lobe; the type specimens were also collected well below

1200 m. Hence, B. papuaensis is removed from the synonymy of B.

atrisetosa and treated here as a senior synonym of B. unichromata, which

agrees with it in all respects.

Excluded species

A further species referred to Paratridacus by Drew (1989), B. mesonotaitha

Drew, is known only from the type female; hence the presence or absence of

the pecten cannot be determined. However, the broader costal band

(confluent with vein R4,5) and more distinctly trilobed aculeus (see Drew

1989, figs 363 and 363A) closely resemble those of B. (Zeugodacus)

sandaricina Drew, its potential sister-species; hence B. mesonotaitha is

placed here in subgenus Zeugodacus, as indicated by Hancock and Drew

(2015). The two species differ primarily in the width of the anepisternal stripe

and both are known only from East Sepik Province, Papua New Guinea.

Key to Austrodacus species

] Scutum black (with yellow lobes and vittae); anepisternal yellow stripe

reaching line of anterior notopleural seta; facial spots faint or absent ...... 2

— Scutum mostly or almost entirely red-brown or orange-brown, rarely

extensively black; anepisternal yellow stripe not reaching line of anterior

notopleural seta; facial spots present ......................eeseseeeeeeeeeeneeeneenn 3

Australian Entomologist, 2016, 43 (2) 81

2 Prescutellar acrostichal setae present; costal cells colourless .....................

DL NN EEI TERASS A TA A-E ET A TETERE OE bento B. alampeta Drew

— Prescutellar acrostichal setae absent; costal cells with a pale fuscous tint

Meu IU 2e er ERASER EDT COCA EERE FLA DOD Mene lr B. abdoaurantiaca Drew

3 Prescutellar acrostichal and supra-alar setae absent; scutum orange-brown

without fuscous markings; anepisternal yellow stripe reaching almost to

anterior notopleural seta ....................... eee B. cucumis (French)

— Prescutellar acrostichal and supra-alar setae present; scutum red-brown

with or without fuscous markings; anepisternal yellow stripe reaching

midway between anterior margin of notopleural callus and anterior

notopleural Setaes csse ie ER rr ER ERE ARR RELIER QUSS SE CeCe rRnntn rn tryst E AES nnn ans 4

4 Abdominal tergites III-V with setulae distinctly yellow; third antennal

segment entirely fulvous (extreme apex sometimes darker); scutum

without black markings; costal cell c colourless ....................................sse.

— Abdominal tergites III-V with setulae white with a gold reflection; third

antennal segment fuscous apically and on outer surface; scutum often

with a pattern of black markings; costal cell c pale fulvous .......................

Nhaxayessegertieeees eec nid partite seni e Horror pee c irent B. atrisetosa (Perkins)

Acknowledgements

We thank Sally Cowan (Department of Agriculture, Cairns) and Daniel

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

Bickel, Russell Cox and John Martin (Australian Museum, Sydney) for

photographs and information on the holotype of B. papuaensis, Christine

Lambkin and Susan Wright (Queensland Museum, Brisbane) and Desley

Tree (Queensland Department of Agriculture and Fisheries, Brisbane) for

checking types and information on their collection holdings.

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and VIRGILIO, M. 2015. A review of the current knowledge on Zeugodacus cucurbitae

(Coquillett) (Diptera, Tephritidae) in Africa, with a list of species included in Zeugodacus.

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and CLARKE, A.C. 2012. A molecular phylogeny for the tribe Dacini (Diptera: Tephritidae):

systematic and biogeographical implications. Molecular Phylogeny and Evolution 64: 513-523.

LEBLANC, L., TORA VUETI, E., DREW, R.A.I. and ALWOOD, A.J. 2012. Host plant records

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Australian Entomologist, 2016, 43 (2): 83-87 83

A NEW ORGAN IN MALE ZYGAENID MOTHS

(LEPIDOPTERA: ZYGAENIDAE: PROCRIDINAE)

BERNARD MOLLET! and GERHARD M. TARMANN?

'16, Parc Vatonne, 91190, Gif-sur-Yvette, France

?Tiroler Landesmuseen, Ferdinandeum, Naturwissenschaftliche Abteilung, Feldstrasse 11a, A-

6020 Innsbruck, Austria

Abstract

A new structure is described comprising a brush of expansible hairs on each side of the thorax in

males of some Australian zygaenid moths in the genera Pollanisus Walker, Hestiochora Meyrick

and Onceropyga Turner. The structure is described and illustrated and its taxonomic implications

are discussed.

Introduction

Recent investigations into the morphology of Australian zygaenids have

revealed a discrete organ that is difficult to discern. This structure is rarely

visible on dried specimens and was first observed on Pollanisus

viridipulverulenta (Guérin-Méneville, 1839) and P. apicalis (Walker, 1854).

It consists of a bunch of hairs of clear colour arranged as a hair-pencil,

inserted laterally on each side of the thorax between the prothorax and

mesothorax, the individual hairs reminiscent of androconial scales (Fig. 1).

Fig. 1. Everted hair-pencil on the thorax of Pollanisus viridipulverulenta male.

84 Australian Entomologist, 2016, 43 (2)

mesot

View A

Left side Right side

hair pencil at rest hair pencil everted

| View B

posterior lateral fold memb 1

\ x

SSS closed position

/ anterior lateral fold \

- memb 2 hair pencil

opened position

3 fold

co | co | N sps

0.1 mm

Figs 2-3. (2) lateral view of the thoracic segments of Pollanisus commoni, imago. ane:

anepisternum; co: coxa; epist: episternum; mesot: mesothorax; par: parapatagium; pat:

patagium; prot: prothorax; teg: tegula; p.f.: protection fold; h.p: hair pencil. (3)

prothorax of Pollanisus trimacula, view A from mesothorax. Left side with tightly

packed bristle stowed in the protecting fold and right side with bristle expanded. ane:

anepisternum; furc: furcopleural bridge; memb 1: membrane connected to

mesothorax; memb 2: membrane connected to coxal; sps: spinasternum sectioned.

Australian Entomologist, 2016, 43 (2) 85

These hairs are confined within a weakly sclerotized translucent fold,

connected to the posterior part of coxa 1 and to the inner skeleton by a fine

membranous diaphragm (Fig. 2). This fold covering the densely grouped

hairs (20-30) is set distally to allow the ‘blooming’ of these hairs, in the form

of a brush, laterally on the outer side of the thorax (Fig. 3).

Discussion

This organ is present on all the species of Pollanisus Walker, 1854 that we

were able to dissect, e.g. P. viridipulverulenta, P. apicalis, P. empyrea

(Meyrick, 1888), P. cupreus (Walker, 1854), P. lithopastus Turner, 1926, P.

commoni Tarmann, 2004, P. contrastus Tarmann, 2004, P. eumetopus

Turner, 1926, P. angustifrons Tarmann, 2004, P. trimacula (Walker, 1854),

P. marriotti Kallies & Mollet, 2011, P. incertus Tarmann, 2004, P. cyanota

(Meyrick, 1886) and P. calliceros Turner, 1926. It is also present on those

species of Hestiochora Meyrick, 1886 that we were able to dissect, i.e. H.

xanthocoma Meyrick, 1886 and H. furcata Tarmann, 2004, and on

Onceropyga anelia Turner, 1906 (Fig. 5).

A modified organ (Fig. 4) without bristles and lateral fold is also present in P.

calliceros. In the genus Hestiochora the organ looks like a bulbous pocket

with soft membranous fold in H. xanthocoma (Fig. 7), while it is well

developed with a large posterior lateral fold in H. furcata (Fig. 6). No bristles

were visible on these two species but further specimens must be examined to

ascertain whether these bristles were merely lost during the lifetime of the

examined specimens or whether the organ is genuinely atrophied. However,

the presence of free arms on the furcopleural bridge is another good generic

character for Hestiochora.

This structure is absent in other Australian Artonini genera, such as

Myrtartona Tarmann, 2004, and Australartona Tarmann, 2004, as well as in

other Procridinae, viz. species of Artona Walker, 1854, Balatea Walker,1865,

Lophosoma cuprea (Walker, 1856), Ephemeroidea ariel Hampson, 1893, E.

virescens Snellen, 1903, species of Clelea Walker, 1854, and Chrysartona

Swinhoe, 1892, Thibetana sieversi (Alphéraky, 1892), T. delavayi (Oberthür,

1894) and in species of European, Asian and American genera of Prodicrini

and in the genus Zygaena Fabricius, 1775.

As far as we know, no similar characters that are localized on the thorax in a

lateral fold located between the pro- and mesothorax have been found in

other Lepidoptera. If the situation in Pollanisus, Hestiochora and

Onceropyga is unique, we have another very good autapomorphy for these

three Australian genera.

Some Amuria Staudinger, 1887 species have also been examined but no setae

in the lateral position described above have been observed so far. However,

similar hairs were found between the forelegs on the prothorax on Amuria

cyclops Staudinger, 1887 and one other Amuria sp. Also, on Pseudoamuria

86 Australian Entomologist, 2016, 43 (2)

melaleuca (Jordan, 1908) from New Guinea, two brushes are clearly

developed but they appear to be inserted ventrally between the prothoracic

coxae rather than laterally, on the outside, as in Pollanisus, Hestiochora and

Onceropyga.

Moreover, if this character is typical for Amuria and Pseudoamuria Tarmann,

2004, it is another argument not to treat the brown 'Artona' species as Artona

or Homophylotis 'Turner, 1904, as both genera lack these brushes.

0.1 mm NY.]

^w anterior lateral

^. posterior lateral fold

Figs 4-7. Posterior view of prothorax relative to mesothorax: (4) P. calliceros; (5) O.

anelia; (6) H. furcata. br. f.: free arms on furcopleural bridge; (7) H. xanthocoma.

Australian Entomologist, 2016, 43 (2) 87

Due to the paucity of material in collections, no specimens belonging to the

genera Amuria, Pseudoamuria and Homophylotis were completely dissected

to check whether any protecting fold, pocket, gland or system connected with

such bristles were present.

The function of this organ is unknown.

The taxonomic value of this character at genus level needs to be studied in

greater depth, especially for Pollanisus and Onceropyga and also further

specimens of Hestiochora need to be examined. On the other hand, the

presence of free arms on the furcopleural bridge is another good generic

character for Hestiochora. The specific differences recognized so far in

Hestiochora are remarkable (between H. furcata and H. xanthocoma).

Acknowledgements

We are indebted to the late N.P. Kristensen (Copenhagen, Denmark) for

fruitful discussion and to Mr A. Kallies (Melbourne, Australia) for providing

zygaenid specimens.

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Palaearctic Zygaenidae. Stenstrup; 304 pp.

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2000. Simferopol; 360 pp.

88 Australian Entomologist, 2016, 43 (2)

RECENT LITERATURE

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2016. Ant megadiversity and its origins in arid Australia. Austral Entomology 55(2): 132-137.

DOWNES, M.F.

2015. Annual cycle of nest composition in the queen-dimorphic weaver ant Polyrhachis

australis Mayr, 1870 (Hymenoptera: Formicidae) in northern Queensland. Austral

Entomology 54: 87-95.

2016. The dimorphic queens of Polyrhachis australis (Hymenoptera: Formicidae). Austral

Entomology 55(2): 163-169.

FLETCHER, M.J. and MONTEITH, G.B.

2016. History of the Australian Entomological Society. Austral Entomology 55(2): 121-131.

GIFFNEY, R.A. and KEMP, D.J.

2016. Maternal care behaviour and kin discrimination in the subsocial bug Tectocoris

diophthalmus (Hemiptera: Scutelleridae). Austral Entomology 55(2): 170-176.

GULLAN, P.J. and WILLIAMS, D.J.

2016. Anew pupillarial scale insect (Hemiptera: Coccoidea: Eriococcidae) from Angophora in

coastal New South Wales, Australia. Zootaxa 4117(1): 85-100.

JAY, K.R., POPKIN-HALL, Z.R., COBLENS, M.J., OBERSKI, J.T., PRASHANT, P.,

SHARMA, P.P. and BOYER, S.L.

2016. New species of Austropurcellia, cryptic short-range endemic mite harvestmen

(Arachnida, Opiliones, Cyphophthalmi) from Australia’s Wet Tropics biodiversity

hotspot. ZooKeys 586: 37-93.

KLOPFSTEIN, S.

2016. Nine new species of Dimophora from Australia (Hymenoptera: Ichneumonidae): new

insights on the distribution of a poorly know genus of parasitic wasps. Austral

Entomology 55(2): 185-207.

KOZLOV, M.V.

2016. Taxonomic revision of Australian long-horn moths of the genus Nemophora

(Lepidoptera: Adelidae). Zootaxa 4097(1): 84-100.

LAMBKIN, K.J.

2016. Revision of the Scytinopteridae (Hemiptera: Cicadomorpha: Scytinopteroidea) of the

Queensland Triassic. Zootaxa 4117(4): 580-590.

MA, Y., ZHAO, C. and GREENSLADE, P.

2016. A new species of Metacoelura (Collembola: Paronellidae) from Australia, and

redescription of Metacoelura articulata. Zootaxa 4105(4): 381-388.

MALIPATIL, M.B., KWAK, M.L. and GUNAWARDENE, N.

2016. A revision of the Australian genus Trachylestes with the description of two new species

(Hemiptera: Heteroptera: Reduviidae: Harpactorinae). Zootaxa 4105(4): 88-100.

MOUND, L.A. and TREE, D.

2016. Genera of the leaf-feeding Dendrothripinae of the world (Thysanoptera, Thripidae),

with new species from Australia and Sulawesi, Indonesia. Zootaxa 4109(5): 569-582.

RICHARDSON, B. J.

2016. New genera, new species and redescriptions of Australian jumping spiders (Araneae:

Salticidae). Zootaxa 4114(5): 501-560.

TAYLOR, R.W. and ALPERT, G.

2016. The myrmicine ant genus Metapone Forel (Hymenoptera: Formicidae): a global

taxonomic review with descriptions of twelve new species. Zootaxa 4105(6): 501-545.

TRIBULL, C.M.

2016. Five new species and key for Australian Epyris Westwood (Hymenoptera, Bethylidae).

Zootaxa 4105(4): 353-367.

Australian Entomologist, 2016, 43 (2): 89-100 89

THE NEST OF POLYRHACHIS AUSTRALIS MAYR

(HYMENOPTERA: FORMICIDAE)

MICHAEL F. DOWNES

26 Canara Street, Cranbrook, Townsville, Qld 4814 (Email: mikedownes @ bigpond.com)

Abstract

Many species of the large ant genus Polyrhachis F.R. Smith make nests using silk from their

own larvae or from spiders. This paper reports on 400 nests of Polyrhachis australis Mayr

dissected at Townsville, Queensland, between 2009 and 2015. Details are given of host plants,

nest structure, materials used (carton and silk), placement of brood within nests, resistance to

disturbances such as rain and nest longevity. Carton nests are typically lined with flat-sheet silk.

Complex internal structure gave more internal surface area, so absolute nest size is not a reliable

indicator of ant numbers. Occasional use of fluffy spider-silk in outer walls led to more flaccid

nest structure. Use of a sticky, ductile form of silk, probably derived from moths, was also

identified. Dedicated brood chambers were not noted, but brood clumping was usual, possibly

representing offspring of different queens. Brood was attached to the nest substrate by diffuse

silk strands. Individual nests could persist for 15 months and grew little after an initial period of

expansion.

Introduction

The exceptionally wide variety of nesting habits in the species-rich ant genus

Polyrhachis Fr. Smith was first reported by Jacobson and Wasmann (1905)

and has since been the subject of many investigations and reports (e.g. Ofer

1970, Hólldobler and Wilson 1983, Dorow and Maschwitz 1990, Robson and

Kohout 2005, 2007, Robson et al. 2015, Tranter and Hughes 2015), while the

use of silk in nest construction has been documented in at least 264

Polyrhachis species (Liefke et al. 1998). There are two confirmed sources of

the silk: from spiders and from larvae of the ants themselves (Robson and

Kohout 2007). Spider silk in nests was reported by Collart (1932) for P.

laboriosa F. Smith, Dwyer and Ebert (1994) for P. australis (as P. doddi

Donisthorpe) and P. pilosa Donisthorpe, and Robson (2004) for P. turneri

Forel. Dwyer and Ebert (1994) reasoned that using spider silk as a first option

in the construction of nests may be the norm for arboreal Polyrhachis weaver

ants, since queens establishing nests independently (haplometrotically) have

no larvae with them, while those establishing nests dependently (i.e. with

worker assistance) often rely on the workers setting up the nests before any

queens arrive; those workers may not import larvae until shelter is sufficient.

Nests can be initiated by fewer than 20 workers in these cases and may be

located preferentially where spiders are sheltering (Dwyer and Ebert 1994).

Polyrhachis australis, whose original habitat is rainforest and woodland

margins (CSIRO 2010), has adapted to the semi-natural suburban

environment of humans, nesting in gardens that are dense, moist and shady.

Methods

Between September 2009 and December 2012, 220 P. australis nests were

collected in Townsville, northern Queensland (Australian dry tropics,

19°18'S, 146?45'E) and their contents recorded. That study (Downes 2015)

90 Australian Entomologist, 2016, 43 (2)

documented seasonality in numbers and life history stages of the weaver ants.

From 2013 to 2015, another 180 nests were examined. Descriptions of the

structure and dynamics of all 400 nests are presented here.

External nest dimensions (missing data 45, n = 355) were taken to estimate

nest size (volume), using the protocol of Downes (2015). Details of the

internal structure were noted during nest dissections. Carton and silk, the

major external and internal constituents other than the supporting architecture

(leaves of the host plants, for the most part), drew particular attention,

especially in the later years of the study. Three-dimensional flocculent

masses of fibres, distinct from the flat sheets of silk lining the interior, were

assumed to be spider silk, following Dwyer and Ebert (1994), Robson (2004)

and Robson and Kohout (2005, 2007, 2008).

Host plants were recorded from May 2013 onwards, but these records must

remain anecdotal: corresponding data on host plant frequency in the habitat

would be needed to quantify preferences.

Between August 2013 and March 2015, 63 nests were tagged in situ and their

growth, decline and fates monitored regularly (weekly or more often).

Results

Host plants

Nests were situated not only on or between leaves (Figs 1-2), but also under

bark or within the hollows of stems. Banana plants (Musa sp., Musaceae)

afforded retreats of the latter kind; likewise the tough, tubular woody fronds

of the Cocos or Queen palm, Syagrus romanzoffiana (Arecaceae). Any

detached piece of curled bark was a potential P. australis nest site.

Other host plants, additional to those in Downes (2015), included the flame

tree (Brachychiton acerifolius, Malvaceae), frangipani (Plumeria rubra,

Apocynaceae) mock orange (Murraya paniculata, Rutaceae), native

mulberry (Morinda citrifolia, Rubiaceae), weeping paperbark, (Melaleuca

leucadendra, Myrtaceae) and powder-puff (Calliandra haematocephala,

Fabaceae). A notable exclusion was yellow oleander (Cascabela thevetia [=

Thevetia peruviana], Apocynaceae), which grew beside and between plants

used for nesting by P. australis. Among the unexpected locations of P.

australis nests were abandoned nests of the green weaver ant, Oecophylla

smaragdina Fabricius.

External structure

Nests were typically built between living leaves, but could also be

constructed on a single flat, folded or curled leaf, living or dead, with lengths

ranging from 2-40 cm. Large leaves, e.g. those of the umbrella tree Schefflera

actinophylla (Araliaceae), provided scope for larger than average nests, but

did not lend themselves to structured interiors. Complex interior structure

was more a feature of nests built on trees with small leaves, e.g. Calliandra,

Australian Entomologist, 2016, 43 (2) 91

because leaves incorporated into the nest interior necessarily produced

complete or incomplete partitions.

Nests could be clustered, e.g. six nests in adjacent stems of one banana plant.

Sometimes, nests were close (2-3 cm apart) or contiguous, so that the only

criterion for their being separate nests was the lack of any internal

connection. In one case there was a nest within a nest: an inner partition of

(outer-wall) carton separated the interior into two apparently independent

areas.

Figs 1-2. Nests of Polyrhachis australis Mayr: (1) workers at entrance of nest

constructed between two leaves. Scale bar 10 mm; (2) nest incorporating several host

plant leaves. Scale bar 20 mm. Photos by Malcolm Tattersall.

Notwithstanding the variety of form, nests could be assigned to one of two

broad types on the basis of construction materials. The commonest were nests

with dark-coloured exteriors consisting of densely-packed particulate matter

(carton) lined wholly or partly by larval silk. The other kind (4.596) had fine,

sparse, particulate plant (and perhaps insect) fragments embedded in cream-

coloured, fluffy, three-dimensional masses of convoluted fibres assumed to

be spider silk (see Methods), making the nests conspicuously pale and

flaccid. Despite their structural weakness, spider-silk nests could be large

when hosted by trees with large leaves.

The mean nest size was 32.4 + 69.9 cm’, range 1.5-1008 cm’, n = 355; the

median nest size was 14 cm’. Hence the distribution was positively skewed

due to a small number of relatively large nests. The number of occupants,

however, correlated only moderately with nest size, R? = 0.47, n = 302, and

the density of ants in nests was similarly affected. See Downes (2015) for

more quantified details.

Entrance/exit holes of carton-based nests were ca 4 mm in diameter and

numbered up to three in large nests. Some had a shallow, raised lip that could

be exaggerated to form a squat turret-like structure. Sometimes an interior

portal in a partition (see below: internal structure) took the form of an

entrance hole, i.e. having the same width and with (or without) a raised lip.

92 Australian Entomologist, 2016, 43 (2)

m 1373 = l 3 S :

Figs 3-6. Nest structure of Polyrhachis australis Mayr: (3-4) perforated partitions of

reinforced silk inside nest. Scale bars 5 mm; (5) nest carton in form of fine and coarse

particulate matter adpressed against silk lining. Scale bar 5 mm; (6) nest carton of

admixed particles. Scale bar 1 mm.

Internal structure

Internal nest structure ranged from none to very complex with multiple silk-

lined galleries, channels and partitions. Rarely, the silk lining of the exterior

walls was reduced or absent. Internal partitions could have one or more

perforations that were usually round or oval (Figs 3-4). The form and extent

of the partitions could follow the arrangements of the constituent leaves, in

which case they were almost invariably silk-lined; but often the partitions

were independently formed from thicker, reinforced (i.e. multi-layered) silk

which could support its own weight independently. Partition perforations

could be small or large relative to the size of the partition, so that in some

cases the ‘partition’ was no more than a strut or pillar crossing a wide gap.

Partitions were almost always transverse relative to any long nest axis, but

could be longitudinal, in one case almost dividing the entire nest internally

into an upper and lower tier with connecting holes before breaking down into

ragged gaps supported by reinforced silk pillars; in another extending

unperforated to the inner end of the nest, forming two blindly-ending

chambers only connected at their forward ends. Partition directions were

indeterminate in nests without a long axis. Internal partitions occurred in

Australian Entomologist, 2016, 43 (2) 93

spider-silk nests as well, but were usually of pale sheet silk (like unbleached

paper), not thickly reinforced; they could also be of the same fluffy spider-

silk as the nest walls, but if so were toughened, presumably with the same

(larval) sheet silk used for ‘standard’ partitions. The chambers demarcated by

the partitions varied widely in size. The antechamber accessed directly from

the entrance hole could be large or small relative to nest size. Rarely, the

antechamber was a 'false', unoccupied one, lined at its inner end with

‘exterior’ carton with another entrance hole — i.e. the nest had a ‘porch’. Only

once did exterior carton form a partition deeper in the nest.

The most consistent and consequential feature of the internal structure was

the one most logically expected, that host plant leaf size and shape influenced

and largely governed the interior nest structure. In general, the smaller the

leaf size, the more leaves were incorporated into the interior structure, the

more complex that structure became, the greater was the internal surface area

and the larger the number of ants able to occupy the nest, relative to the size

of the nest (Fig. 2). Up to 10 or 12 Calliandra leaflets could be bound within

a single nest. In nests of curled-up bark fragments, the curls of bark typically

extended into the interior, providing chambers and increased surface area in

the form of helical tunnels. Twigs and stems of the host plant could also

extend through the nest interior.

Carton and silk

Carton was composed of grasses, bark and other plant material, admixed with

soil and mineral debris and adpressed against the silk lining (Figs 5-7). It

could be soft or hard, its fragmented matter fine or coarse (Fig. 5). While

typically dense, firm, gritty, compact and more or less rigid, it could also be

limp, sparsely endowed with grassy fibres and other imbedded or extruding

matter (Fig. 7), often sparse enough to be effectively transparent, with ants

visible inside. It could be as soft as moist paper, or crisp and dust-dry, and

still contain thriving ants and brood.

Flat-sheet silk lining (Fig. 8) is produced by larvae held and manipulated by

workers. Especially when against wood, as with the Cocos palm fronds and

other curled bark, this silk could be fine and difficult to see but could also

truly be absent, leaving the brood against the bare wood. When deficient in a

nest of dead and living leaves, it was invariably the dead leaves that were

unlined. Spider silk (Fig. 9) typically formed a limp, flocculent mass. It had

very little supporting power, gave easily under its own weight and served

only to connect leaves, the latter supporting the nest.

A third kind of silk, tough, dry, ductile and sticky, was also occasionally

present (Fig. 10). Probably derived from the pupal cocoons of moths, as

discussed below, it could be cut easily with scissors but not torn easily by

pulling. It occurred as a semi-transparent wall taking up more than half the

length of one nest and was encountered in small amounts in several others.

94 Australian Entomologist, 2016, 43 (2)

Figs 7-10. Nest structure of Polyrhachis australis Mayr: (7) nest carton relatively

sparsely packed against inner silk lining visible as a pale swathe from upper left to

lower right. Scale bar 5 mm; (8) flat-sheet silk from larvae, lining the carton of nest

exterior which is visible along lower edge. Scale bar 5 mm; (9) fluffy (spider) silk

inside nest. Scale bar 0.5 mm; (10) ductile moth silk teased out from moth pupal

cocoon inside nest. Scale bar 5 mm.

Brood

Dedicated brood chambers were never identified, but clumping of brood was

the norm (Fig. 11). These brood clumps could be anywhere except close to a

nest entrance. Brood was rarely found in contact with carton, almost always

lying on silk lining living leaves or, less commonly, dead leaves and never

against unsilked leaves. Most brood, especially larvae, was held against the

substrate by loosely applied silk threads of unknown source. These could be

effectively invisible, revealed only against a dark background (Figs 12-13).

Longevity and fate

Carton nests were on the whole resistant to rain but were always at risk of

damage by rain or wind and, even in dry, windless conditions, they could be

breached by falling vegetation. Such damage, when it did occur, inevitably

made the ants and their brood vulnerable to invaders. An infestation of the

big-headed ant, Pheidole megacephala Fabricius, in a breached nest in

February 2014 is an example. The fate of a nest was tied to the fate of the

foliage hosting it, the oldest observed (15 months) being enclosed by a strong

Australian Entomologist, 2016, 43 (2) 95

leaf curled firmly around a stout twig. Of 63 nests tagged and monitored, 31

ended their time naturally with mean longevity 3.4 months. The rest were

taken for censusing at known ages. Only five lasted more than a year.

7 E

Figs 11-13. Early stages of Polyrhachis australis Mayr: (11) brood cluster, including

eggs (e), early (el) and late (ll) instar larvae, a worker pupa (wp) and a male pupa

(mp). Scale bar 5 mm. (12-13) larvae inside nest: (12) larvae seemingly unattached to

the leaf surface. A tear in the silk below the mid-vein reveals attachment between

larvae and silk; (13) larvae held to each other and the surface by a loosely-spun web

of silk. Scale bars 2 mm.

Most tagged nests remained discrete and autonomous, but some developed a

contiguous offshoot or incompletely detached side branch — it was not always

96 Australian Entomologist, 2016, 43 (2)

possible to decide which; or a nest would relocate to a new position up to 10

or 15 cm away for no evident reason (this was judged to be a relocation, not a

case of budding, if no ants remained in the ‘mother’ nest). One nest relocated

to a loop of the plastic flagging tape marking its location.

Nests did not grow slowly and steadily to a maximum size, as expected.

Instead, an initial growth phase of relatively short duration (as little as a week

to ten days in some known cases), was followed by a phase typically lasting

several months during which the nests decreased in size: of 27 nests for

which initial (end of first growth phase, as well as could be judged) and final

nest sizes were recorded, mean nest size decreased by 23% (from 13.9 cm? to

10.7 cm’). Once established on certain leaves of the host plant, there was

almost no tendency to expand the nest by including other leaves, even when

adjacent leaves were close enough. So nests established on trees with large

leaves (e.g. umbrella trees) reached a large size rapidly and remained large,

while those on small-leafed plants did not grow much more, if at all, after

enclosing the initial leaves, and hence remained small. There was often no

evident reason for the demise of failed nests and perfect, undamaged nests on

thriving host leaves were sometimes empty.

Discussion

Host plants and structure

Without data on the relative availability (habitat coverage) of host plants,

their relative frequencies, even if available, would be of no value; so a

checklist must suffice.

Also, because the nests of many Polyrhachis weaver ants, unlike Oecophylla

F. Smith, show high variability in location and host plants (Dwyer and Ebert

1994, Liefke et al. 1998, Robson and Kohout 2007), any list of host plants for

P. australis becomes less intriguing than plants that are never used despite

being available. One such was Cascabela thevetia, which never hosted a P.

australis nest over the 5-year period, showing that host plant selection does

not simply reflect relative abundance. C. thevetia’s intrinsic toxins may be

repellent, or its leaves too narrow (0.5-1.5 cm) for adequate support and

concealment. The latter problem has been solved in a unique way by P.

muelleri Forel: its silk nests are translucent and its larvae are green,

camouflaging them against the leaf surface (Dorow ef al. 1990). A general

understanding of nest site selection by ants will require more studies of the

sort done by Campbell et al. (2013) on arboreal thorn-dwelling ants.

Although arboreal Polyrhachis ants may pull and slightly bend a leaf at its

edge (Hólldobler and Wilson 1983), they cannot curl leaves into desired

positions, because they lack the uniquely cooperative chain-forming

behaviours characteristic of Oecophylla. Hence we find P. australis (and P.

pilosa) preferentially nesting in the folds of leaves curled either naturally or

by agents, especially jumping spiders (Dwyer and Ebert 1994).

Australian Entomologist, 2016, 43 (2) 97

The same authors found that by teasing apart strands to provide ‘a flocculent

mass of fibres', P. australis and P. pilosa constructed nests entirely of spider

silk in the laboratory, when deprived of any other material; while in the field

(SE Queensland), the use of spider silk was found to be common and could

be extensive. In northern Queensland, nests composed primarily of silk from

spider webs or shelters are relatively uncommon but do occur. The sparing

use of particulate matter in a nest otherwise made solely of spider silk has

also been reported for a rock-dwelling spiny ant, P. turneri (Robson 2004).

The number and placement of nest entrances varies widely in P. australis, but

can be fixed in other species, e.g. P muelleri (Dorow et al. 1990).

Nest size (volume) was a poor indicator not only of number and hence

density of occupants, but also of nest age. Chambers large or small could be

occupied or unoccupied and, on this basis alone, volume would not be an

expected index of other parameters. Even a moderate correlation of nest size

with number of occupants was surprising given the extent to which internal

structure governs available surface area.

The nests of some arboreal Polyrhachis species contain several chambers,

others only a single chamber (Liefke et al. 1998). An approximate

consistency in the number (6-7) and arrangement of internal chambers may

be a feature of nest design in P. delecta Kohout (Tranter and Hughes 2015),

but larger sample sizes would be needed to confirm this. P. australis nests

showed no such consistency, responding rather to the constraints and

opportunities afforded by the foliage of the host plant. The 'porch' feature in

some nests may be a device to buffer the effects of rain and/or invaders.

Because queens and other castes relocated during nest dissection, no evidence

could be found for the possibility of intranidal oligogyny, that is, the

demarcation of separate zones controlled by individual queens. Nothing in

the queens' behaviour, however, lent support to the notion that they were

territorial.

Carton and silk

The term 'carton', loosely used in the past, is nowadays taken to mean the

nest's exterior mass of particulate material, excluding the inner silk lining

(Robson and Kohout 2007). However, while the two layers are discrete, silk

may be incorporated into the carton layer as well, as reported by Hólldobler

and Wilson (1983) for P. robsoni Kohout.

Not all arboreal weaver ants of the genus Polyrhachis make nests of carton.

The nests of P. bicolor F. Smith and P. muelleri, for example, are made

largely or exclusively of loosely woven, pure spun silk, leaving them

translucent (Jacobson 1908, Dorow et al. 1990, Liefke et al. 1998). Some P.

australis nests are partly translucent when the carton material is minimal.

The production and use of silk for nest building in Polyrhachis has a

surprisingly labile evolutionary history (Robson et al. 2015). It is almost

98 Australian Entomologist, 2016, 43 (2)

exclusively a feature of arboreal species and the flat-sheet silk used for

interior lining appears to be universally of larval origin in arboreal ants of this

genus (Robson and Kohout 2007). Polyrhachis australis larvae do not spin

pupal cocoons, so the silk they produce is presumably dedicated wholly to

nest building and repair. The images of apparently fluffy silk being extracted

from an ant larva by two workers (Brisbane Insects 2013) must remain a

curiosity pending clearer evidence.

Harvesting and recycling of spider silk by P. australis and P. pilosa has been

directly observed by Dwyer and Ebert (1994), who proposed that the use of

spider silk promotes polydomy and hence control of territory and food

resources, while obviating the need to expose larvae to desiccation and other

hazards when new nests are being initiated. The frequency (1296) with which

jumping spiders (family Salticidae) were found with the weaver ants in the

present study suggests that a mutualism might exist between them, with P.

australis preferentially locating in places where these spiders are plentiful.

Several lepidopteran nest associates, in particular a stathmopodine moth

symbiotic with P. australis, produced silk apparently of material benefit for

the ants, perhaps in part offsetting the depredations of the moth larvae.

Brood

Workers were slow to relocate brood during nest dissection, probably

because the silk strands anchoring the brood to the substrate had to be cut

first. Hence the original clumping of brood was evident. The anchoring

would have minimised dislodgment when the nest was buffeted by wind or

jarred by falling fronds. Brood anchored by silk strands was also noted by

Dorow et al. (1990) for P. muelleri and by Liefke et al. (1998) for several

other Polyrhachis species. Whether the brood clumps of P. australis

represent the output of different queens is unknown.

Ants, especially the brood, are particularly vulnerable to infection on account

of their social habits and low intracolonial genetic diversity (Graystock and

Hughes 2011, Tranter et al. 2014). Hence, these social insects keep their

nests exceptionally clean (Hólldobler and Wilson 1990). Their larval silk may

aid in warding off disease-carrying agents (Fountain and Hughes 2011) and

grooming, as well as nest hygiene, plays a part in disease resistance

(Fefferman et al. 2007). Additionally, segregation of brood clumps into

different chambers, as seems to occur in P. delecta, could play a part in

minimising the spread of harmful agents (Tranter and Hughes 2015). Such

segregation was not evident in P. australis nests, however.

Longevity and fate

Polyrhachis australis nests are necessarily well adapted to a monsoonal

climate, but excessive use of spider silk in their construction increases their

vulnerability to rain (Dwyer and Ebert 1994). The common carton form of

the nest showed no evidence of being thicker or denser on its uppermost part,

Australian Entomologist, 2016, 43 (2) 99

as occurs in the nests of Camponotus senex F. Smith (Hólldobler and Wilson

1983). The social structure of P. australis populations favours polygyny

(Downes 2015), consistent with the suggestion of Oliveira et al. (2011) that

polygyny in the arboreal ant Odontomachus hastatus Fabricius is promoted

when nests are liable to destruction by rain.

An understanding (at least my understanding) of the apparently patternless

set of nest relocations, size fluctuations, hasty desertions of seemingly perfect

nests together with reluctance to abandon other seriously defective ones, to

say nothing of how budding as a reproductive strategy operates within these

constraints, is a distant prospect. Nest longevity is inseparable from the

longevity and changing disposition of the host vegetation and it would be

surprising if polydomy was not in some measure driven by these dynamics.

Since nest size (volume) bore no reliable relation to total ant numbers and

hence to colony productivity, the lack of nest growth (or even the typical nest

shrinkage) in nests monitored for size cannot be taken as indicating any

decline in viability.

Acknowledgements

I thank Malcolm Tattersall for the photographs of nests in situ on host plants

and for access to P. australis nests on his property. Liz Downes, Leigh

Winsor and the carers of the Ross River Bush Garden helped with plant

identification. Ted Edwards kindly lent his time and expertise in efforts to

identify the moth mentioned in this article.

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Australian Entomologist, 2016, 43 (2): 101-107 101

A NEW SPECIES OF MACROTRISTRIA STAL FROM THE

SHOALWATER BAY REGION OF CENTRAL QUEENSLAND

(HEMIPTERA: CICADIDAE: CICADINA E)

L.W. POPPLE

Entomology Section, Queensland Museum, South Brisbane, Qld 4101

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

Abstract

Macrotristria stevewilsoni sp. n. is described from a remote coastal locality in the Shoalwater

Bay region, central Queensland. The species exhibits a distinctive appearance when compared

with its congeners and may be quite restricted in distribution.

Introduction

The genus Macrotristria Stal contains a group of large cicadas distributed

across eastern, northern and western Australia, being most diverse in the

north (Moulds 1990). It was revised substantially by Burns (1964), with

additional species described subsequently by Moulds (1992). The definition

of the genus was recently redefined by Moulds (2012), including a useful set

of distinguishing features. It currently includes 18 species endemic to

Australia and one species from Madagascar (Moulds 2012). The new species

described here is known only from a single locality in the Shoalwater Bay

region of central Queensland. It was discovered through the donation of a

pair of specimens collected from this region during an expedition that took

place in 1992. Terminology for morphological features and higher

classification follows Moulds (2005).

Genus Macrotristria Stal

Macrotristria stevewilsoni sp. n.

(Figs. 1 A-C, 2A-B, 3)

Type material. Holotype 6, QUEENSLAND: [all hand written] 4.5 km N.W. Cliff Pt,

22?3T'lT"S, 150°46’22”E, Shoalwater Bay, 6-2-92 [6.11.1992], S. Wilson. Paratype

Q, [all hand written] 4.5 km N.W. Cliff Pt, Shoalwater Bay, 8 Feb 1992, S. Wilson,

Macrotristria sp. Both in Queensland Museum, Brisbane (Reg. Nos QMT23470

(Holotype) and QMT23471 (Paratype)).

Description. Male (Figs 1A-B, 2A-B). Head. Supra-antennal plate dark

reddish brown to black with brown to orange brown along posterior margins;

postclypeus reddish brown, dark brown to black centrally, with a yellow-

brown spot on anterior; dorsal surface medially yellow brown, streaked dark

brown and reddish brown laterally; gena pale brown, with long silver

pubescence; mandibular plate dark brown to black, paler along margins,

covered by long silver pubescence; frons black, yellow brown anteriorly;

vertex yellow brown with a black ring on each lateral side, extending to

supra-antennal plate and area surrounding ocular tubercles, with sparse silver

pubescence; ocelli pale red; compound eyes brown; anteclypeus reddish

brown with long silver pubescence, rostrum black, dark brown at base and

102 Australian Entomologist, 2016, 43 (2)

along lateral edges, extending to posterior margins of hind coxae; antennae

deep brown to black, pedicels brown proximally, dark brown distally.

Thorax. Pronotum mainly reddish brown, pale brown on medial anterior

margin, with a yellow brown longitudinal fascia medially, broader proximally

and bordered with dark reddish brown to black, extensively so around

posterior edge of fascia; pronotal collar pale brown to dull brown, dark brown

along posterior margin; narrow dark brown band commonly also present

adjacent to pronotal collar. Mesonotum mainly yellow brown with a black

fascia extending from area surrounding scutal depressions, narrowing

proximally and terminating acutely without reaching anterior margin;

submedial sigilla dark reddish brown to black, discrete; lateral sigilla diffuse

dark reddish brown, somewhat irregular in outline, posteriorly rounded, just

reaching anterior arms of cruciform elevation; cruciform elevation yellow

brown to reddish brown, with a conspicuous darker brown or black area

between anterior arms, surrounded with long silver pubescence; scutal

depressions black. Metanotum dark brown, pale brown on posterior margin.

Wings. Forewings: pterostigma pale brown, semi-opaque; basal membrane

red; distinct infuscations along crossveins r, r-m, m and m-cu, and at the

distal ends of veins RA», RP, Mj, M5, M3, M, and CuA;; colour of costal

veins pale brown, CuP+1A and CuA, yellow brown, remaining venation

brown to orange-brown. Hindwings without infuscations; plagas and areas

surrounding veins 3A and 2A mainly white with diffuse red patches; colour

of veins CuA, M, RA and av tending dark brown, other veins grading from

pale brown to brown.

Legs. Fore coxae pale brown with brown streaks, brown on anterior faces;

fore femora brown to orange with dark brown spines; mid and hind coxae

dominantly brown to pale brown, darker medially on anterior faces;

meracantha spikes small, pale brown, barely overlapping opercula; mid and

hind femora brown tending reddish brown anteriorly; fore and mid tibiae

mostly reddish brown, dark brown to black basally and, to a lesser extent,

apically; hind tibiae dominantly reddish brown, darker apically and on spines;

tarsi dark brown on outer sides, pale brown on inner sides; pretarsi dark

reddish brown.

Opercula. Outline broadly rounded, not overlapping; plates gently domed in

disto-medial area, mainly pale brown, tending dark brown to black towards

inner margins and reddish brown on lateral edges.

Abdomen. Tergite 2 with well-developed flexing on paramedial anterior

segment, appearing as a weak flange when viewed ventrally; tergite 1 dark

reddish brown; tergites 2 to 7 each conspicuously reddish brown medially,

dark brown to black anteriorly and laterally, with conspicuous silver

pubescence; tergite 8 reddish brown to brown, dark brown to black towards

posterior margin; sternite I pale brown; sternite II dark brown; sternites III-

VIII reddish brown.

Australian Entomologist, 2016, 43 (2) 103

Fig. 1. (A-C) Macrotristria stevewilsoni sp. n., 4.5 km north-west of Cliff Pt,

Shoalwater, central Queensland (22?37' 17"S 150°46’22”E): (A) male holotype dorsal

view; (B) male holotype ventral view; (C) female paratype dorsal view.

Genitalia. Pygofer brown, tending dark brown to black laterally posteriorly,

including dorsal beak; upper lobes rounded in both lateral and dorsal views;

basal lobes visible in lateral and dorsal views, with expanded rounded apices,

covered in conspicuous hairs; uncus undivided, long, as wide as distance

between upper pygofer lobes, with rounded termination (Fig. 2). Internal

features not examined.

104 Australian Entomologist, 2016, 43 (2)

Timbals. Almost completely enclosed by timbal covers; appearing to have a

form typical of the genus, with long ribs visible. [The timbal covers of the

holotype male were not removed for further examination].

Female (Fig. 1C). Colouration and markings similar to male. Abdominal

segment 8 reddish brown dorsally and ventrally, pale brown to brown

laterally, with distinct, black, longitudinal markings along dorsolateral sides

and surrounding dorsal beak. Ovipositor brown, dark apically; ovipositor

sheath reaching «1 mm beyond anal styles and dorsal beak.

Measurements. N = 1 3, 1 9, mm; body length: § 39.4; 9 37.5; forewing

length: ¢ 51.4; 9 46.4; hindwing length: 3 16.7; 9 15.7; head width: 3 15.7;

O 3.09.

Distinguishing characters. Macrotristria stevewilsoni exhibits a unique

appearance that easily distinguishes it from all other Australian species in the

genus. It differs from M. bindalia Burns, M. dorsalis Ashton, M. douglasi

Burns, M. extrema (Distant), M. frenchi (Distant), M. intersecta (Walker), M.

kulungura Burns, M. lachlani Moulds, M. vittata Moulds and M. worora

Burns by the presence of distinct infuscations along crossveins r, r-m, m and

m-cu, and at the distal stems of veins RA;, RP, Mj, M5, M3, M, and CuA,. It

can be distinguished from M. angularis (Germar), M. kabikabia Burns and M.

maculicollis Ashton by having a predominantly dark brown to reddish brown,

rather than black, abdomen and by possessing a white, as opposed to

extensively orange, hindwing plaga. It differs from M. godingi Distant and M.

sylvara (Distant) by the interior colouration of the pronotum, which is

reddish brown rather than predominantly yellow-brown or green. It differs

from M. hieroglyphicalis (Kirkaldy) and M. thophoides Ashton by having

mainly hyaline, rather than dark brown, basal cells on the forewings and from

M. doddi Ashton by having a conspicuous black border encompassing the

central fascia of the pronotum.

Distribution, habitat and behaviour. Macrotristria stevewilsoni sp. n. is

known from a single location on military land to the northwest of Cliff Point

in the Shoalwater Bay region of central Queensland (Fig. 3). The site is

mapped as regional ecosystem 8.2.14, which is characterised by coastal dune

vegetation dominated by Banksia integrifolia, Corymbia tessellaris and/or

Acacia disparrima, with additional species typically associated with

rainforest communities (Queensland Herbarium 2015). This habitat type is

also present in the Byfield area further south; however this species has not

been encountered there, despite multiple cicada collecting expeditions to this

area (M.S. Moulds pers. comm.). This cicada also has the potential to occur

along the coast further north in central Queensland (e.g. Whitsunday Coast);

however, this would be dependent on the distribution of the species crossing

the St Lawrence Gap, a conspicuous >100 km wide dry corridor that includes

the coastline (Webb and Tracey 1981, Chapple et al. 2011, Fig. 3).

Australian Entomologist, 2016, 43 (2) 105

Fig. 2. Macrotristria stevewilsoni sp. n., holotype male: (A) pygofer viewed laterally

from left; (B) pygofer viewed from a posterior ventral angle to reveal the presentation

of the uncus.

106 Australian Entomologist, 2016, 43 (2)

Other reasons for the lack of detection of this species outside the Shoalwater

Bay area could include a multiple-year life cycle, such that populations

appear as adults infrequently. When present, as for other species in the genus,

adults most likely position themselves on the upper branches of trees,

although observations on this particular species are currently lacking.

Calling song. The males possess well-developed timbals for song production.

Other species in the genus typically produce a loud whine; however the call

of this new species currently remains unknown.

Etymology. This new Mactrotristria species is named in honour of Mr Steve

Wilson, a respected herpetologist, who has a long-standing interest in cicadas

and collected the only available specimens.

Q E ind

1 AG MOD.

3 be, CO BE ) 2j.

i — A — Great Barfien V —

P k ) ais C Reef È x

ONCE à XE ae :

* Mackay v WR OMS JE y

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- St Lawrence collection site

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Byfield

100 km e Yeppoon

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* Mount Morgan NN

Fig. 3. Map of coastal central Queensland showing the single location where the two

available specimens of Macrotristria stevewilsoni sp. n. were collected.

Australian Entomologist, 2016, 43 (2) 107

Acknowledgements

I am grateful to the Queensland Museum for staff and resources to enable the

rapid completion of this study. In particular, I thank Geoff Thompson and

Andy Wang for images of the specimens. Max Moulds kindly checked

through his extensive collection of cicadas for additional material. In

addition, Tony Ewart and Max Moulds provided helpful comments on the

distribution of this species.

References

BURNS, A.N. 1964. Revision of the genus Macrotristria Stal (Cicadidae-Homoptera-

Hemiptera) with descriptions of new species. Memoirs of the National Museum of Victoria 26:

71-123.

CHAPPLE, D.G., HOSKIN, CJ. CHAPPLE, S.N.J. and THOMPSON, M.B. 2011.

Phylogeographic divergence in the widespread delicate skink (Lampropholis delicata)

corresponds to dry habitat barriers in eastern Australia. BMC Evolutionary Biology 11: 191.

MOULDS, M.S. 1990. Australian cicadas. New South Wales University Press, Kensington; 217

pp. 24 pls.

MOULDS, M.S. 1992. Two new species of Macrotristria Stal (Hemiptera: Cicadidae) from

Queensland. Australian Entomological Magazine 19(4): 133-138.

MOULDS, M.S. 2005. An appraisal of the higher classification of cicadas (Hemiptera:

Cicadoidea) with special reference to the Australian fauna. Records of the Australian Museum

57: 375-446.

MOULDS, M.S. 2012. A review of the genera of Australian cicadas (Hemiptera: Cicadoidea).

Zootaxa 3287: 1-262.

QUEENSLAND HERBARIUM. 2015. Regional Ecosystem Description Database (REDD).

Version 9.0 (April 2015).

WEBB, L.J. and TRACEY, J.G. 1981. Australian rainforests: patterns and change. Pp 607-694,

in: Keast, A. (ed.), Ecological biogeography of Australia. W. Junk, The Hague.

108 Australian Entomologist, 2016, 43 (2)

CORRIGENDA

The following corrections should be made to Sands (2015), which appeared

in Volume 42, Part 4 of the Journal:

Page 231, Legend: Figure numbers 35, 36, 37 and 38 should read 27, 28, 29

and 30, respectively.

Page 243: The heading Philiris lucina Waterhouse & Lyell, 1913 should read

Philiris lucina Waterhouse & Lyell, 1914.

Page 248, paragraph 1: The last two sentences, from “The illustrated female

' to '... both wings.’ should be deleted. The female mentioned in the

penultimate sentence is of P. azula azula Wind & Clench.

SANDS, D.P.A. 2015. Review of Australian Philiris Röber (Lepidoptera: Lycaenidae), with

notes on variation and descriptions of two new subspecies from Cape York Peninsula. Australian

Entomologist A2(4): 219-252.

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THE AUSTRALIAN

Entomologist

Volume 43, Part 2, 24 June 2016

BAEHR. M.

A new species of the Tachys (s.1.) ectromoides-group from southeastern

Queensland, Australia (Coleoptera: Carabidae: Bembidini)

DOWNES, M.F.

The nest of Polyrhachis australis Mayr (Hymenoptera: Formicidae)

HANCOCK, D.L. and DREW, R.A.I.

A review of the subgenus Austrodacus Perkins of Bactrocera Macquart

(Diptera: Tephritidae: Dacinae). ...................sssssssssseesereneneneerenne 75

LAMBKIN, T.A.

First Australian records of Euploea wallacei melia Fruhstorfer, 1904

and Euploea stephensii jamesi Butler, 1876 (Lepidoptera: Nymphalidae:

Danainae) from Dauan Island, Torres Strait, Queensland, with notes on

the aggregation habits of Euploea Fabricius species near flowering

MOLLET, B. and TARMANN, G.M.

A new organ in male zygaenid moths (Lepidoptera: Zygaenidae:

Procridinae)

POPPLE, L.W.

A new species of Macrotristria Stal from the Shoalwater Bay Region of

central Queensland (Hemiptera: Cicadidae: Cicadinae)

WELLS, A. and KJER, K.

Norfolk Island's caddisfly is a New Zealander (Trichoptera:

kiydrophilida6y, 2:5. a Uus ue Hr eee gere EET Hn entren HEAR tates oad 49

RECENT LITERATURE

CORRIGENDA

ISSN 1320 6133

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