THE AUSTRALIAN ENTOMOLOGIST

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Queensland Museum

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University of Queensland

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Queensland Museum

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Queensland Museum

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Membership is open to anyone interested in Entomology. Meetings are normally held

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Fly Group

Cover: The very large and strong-flying robberfly Blepharotes coriarius

Wiedemann is widespread across eastern Australia. Blepharotes contains six

described and a similar number of undescribed species, restricted to Australia and

New Guinea. They are easily recognised by their flat, usually yellow or orange

abdomens, that bear dense, lateral tufts of hairs. From an original drawing by Geoff

Thompson.

Australian Entomologist, 2001, 28 (3): 65-68 65

A NOTE ON THE LARVAL FOOD PLANTS OF GRAPHIUM

WEISKEI (RIBBE) (LEPIDOPTERA: PAPILIONIDAE) IN PAPUA

NEW GUINEA

M.F. BRABY' and J. ARMSTRONG?

! Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA

02138-2902, USA

2110 Fernloff Road, Wamboin, NSW 2620

Abstract

Two species of plants, Dryododaphne crassa Schodde (Monimiaceae) and Cryptocarya sp.

(Lauraceae), are recorded as potential larval food plants for Graphium weiskei (Ribbe), based on

ovipositing females in the Kegsugl area, Simbu Province, Papua New Guinea, in upper montane

moss forest during June 1999.

Introduction

Graphium weiskei (Ribbe, 1900) (Figs 1-2) is distributed throughout

mainland New Guinea, occurring as far west as the Arfak Mountains in Irian

Jaya (Indonesia), to Goodenough I. in the D’Entrecasteaux Islands of Papua

New Guinea (Parsons 1998). It most commonly occurs in mid to upper

montane primary forest at altitudes between 1,200 and 2,000 m, although it

has been recorded at altitudes well below and above this range. The species is

well known for its unique and extraordinary colours of pink, mauve and

turquoise. For this reason, high quality specimens are in strong demand for

the overseas butterfly trade and currently sell at 1.5 Kina per specimen (Insect

Farming and Trading Agency [IFTA]).

Surprisingly little is known of the larval food plants and the early stages have

not been described formally. Indeed, all specimens exported by IFTA are

obtained as wild-caught adults (through a network of local collectors) and not

through captive breeding and harvesting of the immature stages. Haugum and

Samson (1980) noted that a female had been observed to oviposit on a

‘species of Sassafras (Lauraceae)’ at Wau and that oviposition also was

observed on a small, unidentified tree at Erume. Parsons (1998) considered

that the ‘small, unidentified tree’ represented a species of Cinnamomum

(Lauraceae) but did not provide evidence for this conclusion. The genus

‘Sassafras’ does not exist; however, the name ‘sassafras’ is used frequently as

the common name for Doryphora sassafras Endl., a species which belongs in

the Monimiaceae, not Lauraceae. Hence, there is considerable doubt over the

identity of the larval food plants in Papua New Guinea.

The following observations, although based on oviposition records only, are

documented here because of the general paucity of reliable information. The

species listed are considered to represent likely food plants and, hopefully,

will stimulate further searching and rearing of the immature stages (and

eventual documentation of the life history), leading to the sustainable farming

of this exquisite butterfly.

66 Australian Entomologist, 2001, 28 (3)

Observations

During June 1999, while stationed for eight days collecting in the Kegsugl

area of Simbu Province, Papua New Guinea, two separate observations were

made of ovipositing females of G. weiskei weiskei in upper montane moss

forest.

The first observation was made at 1350 h on 4 June, near the Lake Pindi

Yaundo Lodge, about 3 km north of Kegsugl, at an altitude of approximately

2,800 m. A female was observed at about eye level (using binoculars, 10x25

magnification) to spend several minutes circling around and settling in the

canopy of a large (ca 30 m in height) Dryododaphne crassa Schodde

(Monimiaceae) (Fig. 3), growing on a steep slope above a watercourse. The

female was seen to eventually settle on the underside of a branch about 2-3 m

from the top of the tree. Once settled she then curled and extended her

abdomen in a characteristic ovipositing manner and laid an egg on the bark,

before flying off to ‘inspect’ other areas of the tree. Further observations of

oviposition and the egg were not possible due to the density of the vegetation

and the height of the tree. However, it is probable that other eggs were being

laid, given the intensity with which the female was searching the tree and

frequently settling.

The second observation occurred at 1225 h on 7 June, in a gully about 3-4 km

south-east of Kegsugl, at an altitude of approximately 2,700 m. A female was

observed at close range to spend several minutes intensely searching, whilst

in flight, the foliage of a shrub (ca 2 m in height) of Cryptocarya sp.

(Lauraceae), growing on the edge of a watercourse. During this period, the

female frequently settled and laid several eggs on the underside of older

leaves in the lower half of the plant. All eggs were deposited singly and were

typical of the genus Graphium Scopoli, being spherical, smooth and

yellowish-green in colour. The female (Figs 1-2) was eventually netted and

retained as a voucher specimen.

Voucher specimens of the two plants mentioned above are deposited in the

Herbarium of the Papua New Guinea Forest Research Institute, Lae.

Discussion

Graphium weiskei belongs to a distinct taxonomic clade (the weiskei group),

within the sarpedon group of subgenus Graphium. The weiskei group

includes five other closely related species (Okano 1984, Hancock 1985,

Parsons 1998, Miiller and Tennent 1999). The life histories and larval food

plants are unknown for four of these species: G. batjanensis Okano from

Batjan [= Bacan], northern Maluku (Indonesia), G. stresemanni (Rothschild)

from Ceram [= Seram], southern Maluku (Indonesia), G. kosii Miiller &

Tennent from New Ireland (Papua New Guinea) and G. gelon (Boisduval)

from New Caledonia and the Loyalty Is.

Australian Entomologist, 2001, 28 (3)

Figs 1-3. (1-2) Graphium weiskei, adult female, upper and undersides; (3)

Dryododaphne crassa (Monimiaceae), putative larval food plant of G. weiskei at

Kegsugl (2,800 m), Simbu Province, Papua New Guinea.

68 Australian Entomologist, 2001, 28 (3)

The fifth species in the group is G. macleayanum (Leach), whose larvae, in

Australia, are recorded feeding on a large number of plants belonging

primarily to the Monimiaceae and Lauraceae, with a few species of Rutaceae

and Winteraceae also being used (Common and Waterhouse 1981).

Doryphora sassafras is commonly used in New South Wales and at least

three species of Cryptocarya are recorded as host plants. It is therefore

interesting to note that the putative food plants recorded for G. weiskei in

eastern mainland Papua New Guinea also belong to the Monimiaceae and

Lauraceae, two primitive and closely related families of plants. Since closely

related species of butterflies frequently share similar larval food plants at the

higher (and sometimes lower) taxonomic levels as a result of coevolution

(Ehrlich and Raven 1965), it is considered very likely that G. weiskei larvae

would feed on plants within the Monimiaceae and Lauraceae and that

Dryododaphne crassa and Cryptocarya sp. almost certainly represent larval

food plants.

Acknowledgments

We are greatly indebted to Joe Wiakabu of the Papua New Guinea Forest

Research Institute (Lae) for identifying the plants. William Wanuma and

Peter Parr of Denglagu Mission and John Dobunaba of the Papua New

Guinea Forest Research Institute kindly assisted in other ways.

References

COMMON, I.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. 2nd, revised

edition. Angus and Robertson, Sydney; xiv+682 pp.

EHRLICH, P.R. and RAVEN, P.H. 1965. Butterflies and plants: a study in coevolution.

Evolution 18: 586-608.

HANCOCK, D.L. 1985. Notes on the taxonomy and distribution of Indo-Australian Papilionidae

(Lepidoptera). Australian Entomological Magazine 12: 29-34.

HAUGUM, J. and SAMSON, C.J. 1980. Notes on Graphium weiskei. Lepidoptera Group of

1968 Newsletter, Supplement 8: 1-12.

MULLER, C.J. and TENNENT, W.J. 1999. A new species of Graphium Scopoli (Lepidoptera:

Papilionidae) from the Bismarck Archipelago, Papua New Guinea. Records of the Australian

Museum 51: 161-168.

OKANO, K. 1984. On the butterflies of ‘Graphium weiskei' group (Papilionidae) with

description of a new species. Tokurana 8(1): 1-20.

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

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

Australian Entomologist, 2001, 28 (3): 69-84 69

EMERGENCE PATTERNS AND DENSITIES OF CICADAS

(HEMIPTERA: CICADIDAE) NEAR CALOUNDRA, SOUTH-EAST

QUEENSLAND

A. EWART

Entomology Section, Queensland Museum, PO Box 3300, South Brisbane, Qld 4101

Abstract

A census of cicada exuviae, collected regularly over 5 months, is reported from a 5.0 ha site of

coastal parkland with fringing mangroves, at Golden Beach, 3-4 km south of Caloundra, S.E.

Queensland. Two emergence patterns occurred, both extending over about 4 months. One type

characterized the higher density species Psaltoda plaga (Walker), P. claripennis Ashton, P.

harrisii (Leach) and Arunta perulata (Guérin-Méneville), in which “explosive” emergence

reached a peak in early-mid December, then slowly decreased over 75-80 days. Male/female

ratios were initially high, but progressively decreased to become female dominated. The second

pattern, shown by the low density species Cicadetta hackeri (Distant) and Abricta curvicosta

(Germar), showed “diffuse” emergence with no systematic change of sex ratios. Exuvial

densities for the six species were, respectively, 3,928, 3,582, 424, 292, 93 and 50 per ha, with a

total density of 8,369 per ha (49 per tree), consistent with overseas data. Emergence of the six

cicada species was synchronous, but their songs are species specific.

Introduction

Cicadas are a characteristic, conspicuous and often noisy component of the

south-east Queensland (SEQ) summer insect fauna. Along coastal SEQ,

cicada numbers are usually high and extend into adjacent mangrove zones.

Common species include Psaltoda plaga (Walker), Psaltoda claripennis

Ashton, Psaltoda harrisii (Leach), Arunta perulata (Guérin-Méneville),

Arunta interclusa (Walker), Pauropsalta rubea (Goding & Froggatt),

Pauropsalta aktites Ewart, Cicadetta hackeri (Distant), Cicadetta oldfieldi

(Distant), Cicadetta stradbrokensis (Distant), Abricta curvicosta (Germar)

and Birrima varians (Germar). Surprisingly, only rather qualitative and

anecdotal published data exist as to relative abundance and emergence

patterns of Australian cicada species (e.g. summaries in Moulds 1990),

although Coombs (1996) has reported on a four year survey of seasonal

cicada occurrences for the New England Tablelands.

Cicada numbers are difficult to estimate quantitatively. The adults are cryptic,

wary, mobile and often occur high in tree foliage, while the nymphal stage is

passed underground within 'root-crown' systems. Previously published

estimates of densities, from USA, Italy, South Africa and New Zealand, are

based on counts of emergence holes, counts of nymphal skins (exuviae) and

emergence traps (Dybas and Davis 1962, White et al. 1979, Karban 1984,

Dean and Milton 1991, Milton and Dean 1992, White and Sedcole 1993,

Williams et al. 1993, Anderson 1994), together with sound level

measurements (Patterson et al. 1997).

The aims of the present survey were: (i) Estimate population densities of six

cicada species within a selected coastal section of SEQ, by means of exuvial

counts, during a complete emergence season; (ii) Document emergence

70 Australian Entomologist, 2001, 28 (3)

patterns and synchrony, including sex ratios; (iii) Estimate longevity of adult

populations; (iv) Compare differences of song characteristics, considered

critical to mate selection, between the temporally and spatially overlapping

species.

Study area and methods

The census area comprised a narrow strip of coastal parkland, adjacent to

high water mark, at Golden Beach, between 3.2 to 4.2 km south of Caloundra,

SEQ. The area extended south from the ‘Military Jetty’ (26°50.06'S,

153°07.12'E) adjacent to the Bribie Channel, to Bells Creek, then to the

western end of a local park (Jensen Park) adjacent to Bells Creek (26°50.41'S,

153°06.62'E). The site area can be referenced on Queensland Topographic

Map Series R834, sheet 9544-III-NW, Caloundra (special), 1:25,000. The

traverse length was 1.38 km, area 5.0 ha (excluding mangroves). The site

consists of Holocene beach sands overlying estuarine muds and silts.

Dominant trees on the site are Coast She-Oak (Casuarina equisetifolia),

Swamp Oak (Casuarina glauca), Blue Gum (Eucalyptus tereticornis),

Common Paperbark (Melaleuca quinquenervia), Cotton Tree (Hibiscus

tilaceous) and exotic Norfolk Pine (Araucaria heterophylla). Less common

trees include Tuckeroo (Cupaniopsis anacardiodes), Grey Ironbark

(Eucalyptus drepanophylla), Swamp Oak (Banksia integrefolia), Hickory

Wattle (Acacia aulacocarpa), Cabbage Tree Palm (Livistonia australis),

Screw Pine (Pandanus tectorius), Red Ash (Alphitonia excelsa) and Bribie

Island Pine (Callitris columellaris). Mangroves are dominated by Avicennia

marina with small numbers of Bruguiera gymnorrhiza.

Previous observations indicated that the site has an abundant cicada fauna

between December-February. Exuviae (Figs 1-6) of the following six species

were found in sufficient numbers for census purposes, listed in decreasing

order of abundance based on qualitative estimates of intensities of singing: P.

plaga (dominant), P. claripennis, A. perulata, P. harrisii, C. hackeri and A.

curvicosta. Systematic searches were normally undertaken daily (less often

each second day), from 10 October 1997 to 27 March 1998. The last day that

newly emerged nymphs were located was 11 March. Results are presented

(Figs 7-12) as successive two-day averages. Where a day was missed, the

exuviae collected on the following day were averaged over the appropriate

two-day period. The only major sampling gap occurred between 29 January

and 16 February; exuvial numbers collected on 17 February were linearly

extrapolated through the sampling gap, assuming a male/female ratio of unity

(close to observed ratios either side of gap). Exactly the same area and trees

were searched each day and all exuviae found were collected, categorised and

counted. Exuviae were conspicuous, normally occurring on tree trunks to

heights of about 3.5 m (rarely higher), with small numbers occurring on long

grass.

Australian Entomologist, 2001, 28 (3) 71

Figs 1-6. Exuviae of the six species of cicada surveyed from southern Golden Beach,

Caloundra. (1) P. plaga; (2) P. claripennis; (3) P. harrisii; (4) A. curvicosta; (5) C.

hackeri; (6) A. perulata. 1-4 and 6 are males; 5 female. Scale lines = 1 cm. 1A, 2A,

3A, are fore leg femora of the exuviae of, respectively, P. plaga, P. claripennis and P.

harrisii. Scale bars = 5 mm.

72 Australian Entomologist, 2001, 28 (3)

Eclosion occurred mostly at night but, at the height of the season, ‘stragglers’

emerged during morning, even to midday. Changing cicada densities were

therefore equated with the number of exuviae progressively collected during

the entire emergence period. 41,842 exuviae were recovered in total. Of the

852 trees occurring within the census site, 90% were ‘productive’, with

exuviae found on them at some stage during the emergence season.

Specific identification of exuviae was based on a sample subset from which

adults were observed emerging. The separation of exuviae of A. perulata, A.

curvicosta and C. hackeri, from each other and from the three Psaltoda

species, was achieved using size, markings and abdominal morphology (Figs

1-6). The overall morphologies of the three Psaltoda species, however,

extensively overlap, including the fore leg femora. In view of the large

numbers involved, identification was based on total body length, as listed in

Table 1. The following size ranges were used: P. plaga 222.5 mm; P.

claripennis 219.9, «22.5 mm; P. harrisii £19.9 mm.

Notwithstanding the size discontinuities within the subset of reference

samples, it was evident during counting that size continuities do exist between

the three Psaltoda species, with females showing an overall bias towards

slightly smaller sizes within each range. It is therefore possible that some

‘leakage’ of female exuviae into the adjacent smaller range division has

occurred. A further potential difficulty is the curvature developed in exuviae

during drying; when excessively pronounced, allowance for curvature was

made. The reference specimens, however, also exhibited varying degrees of

curvature (Figs 1-6), ensuring that the effect is minimised. All exuviae were

sexed according to shape of the developing genitalia at the ventral tips of the

nymphal abdomens.

Table 1. Body length statistics of identified Psaltoda exuviae. (Measurements are in

mm).

Males Females Total

Mean G n Mean (o) n Mean [9] Range n

P. plaga — 23.92 1.26 12. 24.43 143 4 23.97 1.26 22.71- 16

25.23

P. clari- 21.37 0.94 10 20.66 1.19 Mil AL) aa 19.89- 21

pennis 22.11

P. 18.75 1.28 10 18.42 1.00 5 18.66 1.17 17.47- 15

harrisii* 19.81

* Supplemented with material collected from outside study area.

Results and discussion

The results of the census are summarised in Table 2.

Australian Entomologist, 2001, 28 (3) 73

Table 2. Summary of data and results of exuvial census, October 1997 to March

1998, at South Golden Beach, SEQ.

Traverse length: 1.38 km Traverse area: 5.0 ha

Total number of trees: 852 Number of ‘productive’ trees: 763

Total exuviae collected: 41,842 Mean exuviae per tree: 49 (55 per

‘productive’ tree)

Species Peak Date of Date of Mean Sex Number of

eclosion first last density ratio days of

dates emergence emergence (perha) (M/F) nymphal

emergence

period

P. plaga 4-13 Dec 20 Nov 10 March 3928 1.05 112

P. claripennis 4-11 Dec 20 Nov 10 March 3582 0.97 112

P. harrisii 8-11 Dec 28 Nov 1 March 424 0.59 94

A. perulata 22 Dec - 9 Nov 28 Feb 292 0.95 113

2 Jan

C. hackeri Diffuse 19 Oct 11 March 93 0.68 144

emergence (= 2307)

A. curvicosta 8 -25 Dec 24 Nov 23 Feb 50 1.15 92

(relatively

diffuse)

Total exuviae - - - 8369 0.980 -

*Period during which singing was heard within and around the census study site (see

text).

Exuvial losses and predation

Two main causes of exuviae losses were observed:

(i) Periodic strong winds often dislodged empty exuviae from trees but, as

they became trapped in surrounding grassland, the effect is believed to be

minimal in terms of the census. Strong winds, however, do cause deformation

of soft wings in freshly emerging adults, being most pronounced during day

emergences (winds at their peak). This clearly will affect mobility but may

not necessarily prevent female mating (e.g. Karban 1984) or affect exuviae

census results.

(ii) Diurnal predation, especially by birds, of both emerging nymphs and

adults was observed. Predators included Noisy Miner (Manorina

melanocephala), Grey and Pied Butcher Birds (Cracticus torquatus, C.

nigrogularis), Magpie-Lark (Grallina cyanoleuca), Torresian Crow (Corvus

orru), Blue-Faced Honeyeater (Entomyzon cyanotis), Brush Wattlebird

(Anthochaera chrysoptera), White-Faced Heron (Egretta novaehollandiae),

and the nocturnal Tawny Frogmouth (Podargus strigoides) (based on faecal

pellets with nymphal and adult cicada remains). These birds hunted

74 Australian Entomologist, 2001, 28 (3)

individually and some also in small groups (<6 birds), searching for emerging

nymphs and especially adults. Moreover, except for the Tawny Frogmouth,

bird predation of emerging nymphs will be minimal at night when nymphal

emergence is at its peak. Sporadic diurnal predation of emerging nymphs by

the Bearded Dragon (Amphibolurus barbatus) was noted. Nymphs that failed

to eclose were very rarely observed and evidently did not represent a

significant mortality factor (cf. White et al. 1979).

Although no quantitative estimates of nymphal losses to predation could be

made, qualitative observations suggest them to be <10%. Total exuvial

numbers collected in this census will, however, provide minimum estimates of

the emerging nymphal population and should provide an estimate of changing

adult population patterns, over successive two-day intervals, during the

summer season.

Exuvial distribution and adult dispersion

Exuviae were not evenly distributed through the census site, although no

obvious differences of vegetation or soils were observed.

Psaltoda and Abricta nymphs emerged on all tree species except the exotic

Norfolk Pines and only very rarely on the Common Paperbark. A. perulata

extensively utilised casuarinas and sporadically the Norfolk Pines. C. hackeri

emerged exclusively on the Common Paperbark. No eclosions were observed

within tidally inundated mangroves, an observation that applies widely along

coastal SEQ and includes nymphs of the mangrove cicada (A. interclusa).

Only where local sand accumulations (above tidal inundation levels) had

occurred within mangroves were occasional nymphal emergences found. The

sites of emergence are significant as P. plaga, after emerging, aggregated

within and adjacent to mangroves, as well as other vegetation adjacent to high

water mark. P. harrisii formed smaller, localised aggregations high within

clumps of casuarinas above the tidal zone, while P. claripennis dispersed

more widely including into surrounding suburban gardens. A. perulata

remained in trees close to the tidal zone, C. hackeri remained in the

paperbarks, while A. curvicosta dispersed widely in low abundances in most

tree types. The above observations are consistent with the habitat preferences

of P. plaga, P. harrisii, A. perulata, and A. curvicosta which were determined

quantitatively by MacNally and Doolan (1986) within a New South Wales

coastal zone.

Emergence patterns

Two types of emergence patterns are illustrated by the data (Figs 7-12):

(i) The Psaltoda species, representing higher abundance, medium to larger-

sized cicadas, had almost ‘explosive’ emergence patterns. The number of

emerging nymphs rapidly increased to a peak over 10-15 days (late

November to early December), followed by a relatively slow but uneven

decline lasting about 75-80 days, terminating in early March. The total

Australian Entomologist, 2001, 28 (3) 75

eclosion period thus lasted about 4 months, with synchronous emergence

occurring between the three species, even at peak emergence. The emergence

patterns did not define smooth curves, with smaller secondary peaks evident

in later December and January. During peak emergence, large numbers

(«100-300) eclosed from localised trees or tree clumps, continuing for 3-4

nights, after which very small numbers emerged («3) for a further 3-5 nights.

Major emergence centres, in the meantime, had shifted to new sites (i.e.

smaller scale emergence patterns were not strictly synchronous). Following

the major emergence phase, the pattern was of small emergence numbers

spread widely throughout the census area, with localised sporadic bursts of

increased eclosion from both previously productive and unproductive trees.

Male/female sex ratios exhibited initial male dominance, the ratios then

decreasing smoothly towards female dominated eclosion immediately

following peak emergence. Ratios then tended to approach unity.

The total sex ratios for P. plaga and P. claripennis were close to unity. The

sex ratio was, however, female biased for the less abundant P. harrisii (Table

2). The reason for this is unclear, but may be one case where significant early

selective predation of male dominant nymphs did occur.

Arunta perulata exhibited a similar, but more symmetrical emergence pattern

than the Psaltoda species, with later peak emergence and lower densities. The

sex ratio changes were similar but less pronounced.

(ii) The second pattern, that of ‘diffuse’ emergence, was exemplified by C.

hackeri. No clear emergence peak occured and no systematic change in sex

ratio was observed. This is an example of a widely distributed, highly cryptic

cicada which exists in relatively low densities, especially along coastal SEQ

in wallum and swamp environments where paperbarks are common. A.

curvicosta also falls into this category and, although exhibiting a poorly

defined peak emergence in December, displayed no systematic sex ratio

changes. It is again a low density species in the census area.

Sex ratio changes during adult emergence, initially male dominated, were

reported in some Odonata (Corbet 1999), moths (Young 1997) and the

American periodical cicada (Williams et al. 1993). In the present case,

although males did not all emerge before females, a systematic ratio change

was apparent during the emergence season for the more abundant species.

The periods during which active singing were noted are shown in Figs 7-10

and 12. For P. harrisii and P. claripennis, the first songs were heard only 10-

15 days after the dates of initial eclosion, whereas the cessation of singing

approximately corresponded to the dates of final nymphal emergence. For P.

plaga, initial singing correlated with initial eclosions but singing continued

sporadically for 26 days after eclosion had ceased, suggesting that a small

number of individuals had survived for over three weeks.

Australian Entomologist, 2001, 28 (3)

= £a Males dominant

as 1

= È Females

fa ST dominant

0.1

1000

100

Total exuviae collected

(Two day averages)

1 | Singing - extended to 5 April

20 40 60 80 100 120 140 160

Nov | Dec | Jan | Feb | March

Days of Observation - 1997/1998

Fig. 7. Emergence pattern and accompanying sex ratio changes, in P. plaga at

southern Golden Beach, Caloundra during 1997/1998. Period during which singing

was heard is shown by bar.

Australian Entomologist, 2001, 28 (3) TI

Males dominant

Females

dominant

Ratios Males/Females

3 PAN

S 5° 100 e,

= 5 e * toa

Q = e

ei e

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1 Singing

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20 40 60 80 100 120 140 160

Nov | Dec | Jan | Feb | March

Days of Observation - 1997/1998

Fig. 8. Emergence pattern and accompanying sex ratio changes in P. claripennis at

southern Golden Beach, Caloundra during 1997/1998. Period of singing is shown by

bar.

Australian Entomologist, 2001, 28 (3)

pa Males dominant

GIA

= [E

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= |=

= = Females

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= =

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Days of Observation - 1997/1998

Fig. 9. Emergence pattern and accompanying sex ratio changes in P. harrisii at

southern Golden Beach, Caloundra during 1997/1998. Period of singing is shown by

bar.

Australian Entomologist, 2001, 28 (3) 719

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È =

2 sS 1

SA >

S ©

SÈ

a E

[aJ —

Females dominant

eum

120 140 160

-1

(CA

Total exuviae collected

(Two day averages)

Singing

20 40 60 80 100 120 140 160

Nov Dec Jan | Feb | March

Days of Observation - 1997/1998

Fig. 10. Emergence pattern and accompanying sex ratio changes in A. perulata at

southern Golden Beach, Caloundra during 1997/1998. Period of singing is shown by

bar.

80 Australian Entomologist, 2001, 28 (3)

$ = Males

g o dominant

SÉ

> 3

S ©

5 E Females

E E dominant

0 20 40 60 80 100 120 140 160

= 10

S 5

9 >

S R

ZÉ

= 1

Cicadetta hackeri

0.5

0 20 40 60 80 100 120 140 160

Oct| Nov | Dec | Jan | Feb | March

Days of Observation - 1997/1998

Fig. 11. Emergence pattern and accompanying sex ratio changes in C. hackeri at

southern Golden Beach, Caloundra during 1997/1998. Singing was heard from early

September to early May in and around the census site (see text).

Australian Entomologist, 2001, 28 (3) 81

Males dominant

= DP

È È

£A

a p>

ZÈ

Females dominant

20 40 60 80 100 120 140 160

Abricta curvicosta

Total exuviae collected

(Two day averages)

Singing

——r, o zu

20 40 60 80 100. 120 140 160

Nov | Dec | Jan | Feb | March

Days of Observation - 1996/1997

Fig. 12. Emergence pattern and accompanying sex ratio changes in A. curvicosta at

southern Golden Beach, Caloundra during 1997/1998. Period of singing is shown by

bar.

82 Australian Entomologist, 2001, 28 (3)

Cicadetta hackeri was heard singing from early September to early May

(including areas peripheral to the study site), although exuviae were found

only between 19 October and 11 March in the study site (Fig. 11). It therefore

seems likely that the emergence season of this cicada lasted 7-8 months.

Exuvial densities

Psaltoda plaga and P. claripennis were the two dominant species, followed

by P. harrisii, A. perulata, C. hackeri and A. curvicosta (Table 2), consistent

with previous qualitative song observations. The total density of collected

exuviae represented 8,370 individuals per ha. This may be compared with

density estimates from comparable overseas studies (Table 3).

Table 3. Comparative overseas cicada population data.

Environment Range Mean Reference

(ha^) (ha!)

American periodical

cicadas (adults)

Magicicada septendecim Hardwood 11,000- - Karban 1984

forest 63,000

M. cassini (dominant) Suburban - 1,670,000 White et al.

1979

M. tredecassini Hardwood - 66,456 Williams et al.

(dominant) forest 1993

M. cassini Flood plain 1,160,000- 3,720,000 Dybas and

forest 8,350,000 (1956)* Davis 1962

M. septendecim Upland forest 20,000- 328,000 Dybas and

1,100,000 Davis 1962

South Africa,

southern Karoo

Quintillia cf. conspersa Arid shrubland 800- 6,080- Dean and

(dominant) 24,800 8,760 Milton 1992;

Milton and

Dean 1992

New Zealand

Six species Subalpine <1,000- - White and

grassland 20,000 Sedcole 1993

(see also Lane

1993)

Italy, Tuscany,

Mediterranean coastal

Cicada orni Pinewoods 4,882- 19,722 Patterson et al.

36,582 1997

Olive groves 2,172- 3,236 Patterson et al.

4,482 1997

* By 1973 this number had dropped to 756,000, following Dutch Elm Disease (White

et al. 1979).

Australian Entomologist, 2001, 28 (3) 83

The most spectacular cicada concentrations occur within the three species of

American Periodical Cicadas, which can reach “super-abundance” levels in

excess of a million per ha. Such numbers are not reported in Australia. The

numbers reported from New Zealand sub-alpine grassland (six species), South

African Karoo (dominated by 1 species), and coastal Italy (1 species) all

encompass the total abundance estimates found in this study. The most

relevant of the overseas estimates is the Mediterranean coastal area of

Tuscany, containing pinewood and olive grove habitats. Pinewood contained

the highest cicada populations (mean 19,722 per ha), compared to a mean of

3,236 per ha for olive groves. Although the pinewood habitat had higher

overall populations per ha than reported here, estimates of exuviae per tree

were 15.3, compared to 5.9 for olive groves (Patterson et al. 1997). The

present study found a mean of 49 exuviae per tree (total data), or 23 and 21

per tree for P. plaga and P. claripennis respectively, higher than the Tuscany

estimates. Overall, the exuviae (= cicada) densities found in this SEQ survey

appear unexceptional.

Synchronous emergence and interspecific song recognition

Young (1980), in a study of peak emergence periods and habitats of cicadas

in Costa Rica, concluded that his data supported the hypothesis that selection

favoured emergence adaptations such as allochrony or habitat non-overlap

amongst species. In the SEQ environment surveyed, synchrony of cicada

emergences was clearly demonstrated, although the six cicada species

occupied localised but still overlapping habitat niches. Such overlaps require

that their mate recognition signals, specifically their songs, are clearly distinct

from the spatially associated species.

The temporal structures (oscillograms/waveform plots) of the songs from

each species were reported by Young and Josephson (1983 [P. plaga is listed

as P. argentata]) and Ewart (1995). These show the distinctive structures of

each song as seen by their pulse and phrase structures and pulse repetition

rates. The differences are reinforced by their dominant frequencies. For P.

plaga, P. claripennis, P. harrisii, A. perulata, C. hackeri and A. curvicosta,

the dominant frequencies are, respectively, 3.6-4.7, 5.9-6.8, 4.3, 6.7-6.8,

10.6-11.1 and 9.5-9.6 kHz (Ewart, unpublished data). Further distinctions are

seen in detailed structures of the frequency bands, i.e. whether broad or

narrow, measured as bandwidths. These are derived from song spectra, in

which the relative sound energy emitted between the lower (25%) and upper

(75%) quartiles is determined. Respective values for the six cicadas species

are 2.7, 2.1, 1.9, 1.4, 1.9 and 2.3 kHz. The songs of each species have their

own uniquely defined acoustic characteristics.

Acknowledgments

Special thanks go to Thelma Ewart for extensive assistance in collecting

exuviae and to Dr G.B. Monteith (Queensland Museum) and two reviewers

for constructive comments on the manuscript.

84 Australian Entomologist, 2001, 28 (3)

References

ANDERSON, D.C. 1994. Are cicadas (Diceroprocta apache) both a “keystone” and a “critical-

link” species in lower Colorado River riparian communities. Southwestern Naturalist 39: 26-33.

COOMBS, M. 1996. Seasonality of cicadas (Hemiptera) on the northern tablelands of New

South Wales. Australian Entomologist 23(2): 55-60.

CORBET, P.S. 1999. Dragonflies: behaviour and ecology of Odonata. Harley Books (B.H. &

A. Harley Ltd.), Colchester, Essex, England; 829 pp.

DEAN, W.R.J. and MILTON, S.J. 1991. Emergence and oviposition of Quintillia cf. conspersa

Karsch (Homoptera: Cicadidae) in the southern Karoo, South Africa. Journal of the

Entomological Society of Southern Africa 54(2): 111-119.

DYBAS, H.S. and DAVIS, D.D. 1962. A population census of seventeen-year periodical cicadas

(Homoptera: Cicadidae: Magicicada). Ecology 43(3): 432-444.

EWART, A. 1995. Cicadas. Pp 79-88, in M. Ryan (ed.), Wildlife of Greater Brisbane.

Queensland Museum, Brisbane; 340 pp.

KARBAN, R. 1984. Opposite density effects of nymphal and adult mortality for periodical

cicadas. Ecology 65(5): 1656-1661.

LANE, D.H. 1993. Can flawed statistics be a substitute for real biology? New Zealand Journal

of Zoology 20: 51-59.

MacNALLY, R.C. and DOOLAN, J.M. 1986. An empirical approach to guild structure: habitat

relationships in nine species of eastern-Australian cicadas. Oikos 47(1): 33-46.

MILTON, S.J. and DEAN, W.R.J. 1992. An underground index of rangeland degradation:

cicadas in arid southern Africa. Oecologia 91: 288-291.

MOULDS, M.S. 1990. Australian cicadas. New South Wales University Press, Kensington,

NSW; 217 pp.

PATTERSON, I.J., MASSEI, G. and GENOV, P. 1997. The density of cicadas Cicada orni in

Mediterranean coastal habitats. Ztalian Journal of Zoology 64: 141-146.

WHITE, J., LLOYD, M. and ZAR, J.H. 1979. Faulty eclosion in crowded suburban periodical

cicadas: Populations out of control. Ecology 60(2): 305-315.

WHITE, E.G. and SEDCOLE, J.R. 1993. A study of the abundance and patchiness of cicada

nymphs (Homoptera: Tibicinidae) in a New Zealand subalpine shrub-grassland. New Zealand

Journal of Zoology 20: 38-51

WILLIAMS, K.S., SMITH, K.G. and STEPHEN, F.M. 1993. Emergence of 13-yr periodical

cicadas (Cicadidae: Magicicada): Phenology, mortality, and predator satiation. Ecology 74(4):

1143-1152.

YOUNG, A.M. 1980. Environmental partitioning in lowland tropical rainforest cicadas. Journal

of the New York Entomological Society 88(2): 86-101.

YOUNG, D. and JOSEPHSON, R.K. 1983. Mechanisms of sound-production and muscle

contraction kinetics in cicadas. Journal of Comparative Physiology 152: 183-195.

YOUNG, M. 1997. The natural history of moths. T. & A.D. Poyser Ltd., London; 271 pp.

Australian Entomologist, 2001, 28 (3): 85-90 85

WHAT IS NACADUBA MALLICOLLO MARKIRA TITE?

A NEW SPECIES OF NACADUBA MOORE FROM THE SOLOMON

ISLANDS (LEPIDOPTERA: LYCAENIDAE)

W. JOHN TENNENT

Biogeography and Conservation Laboratory, Department of Entomology, The Natural History

Museum, London SW7 5BD, UK

(address for correspondence: 38 Colin McLean Road, Dereham, Norfolk NR19 2RY, England)

Abstract

Following recent collection of material in the Solomon Islands, the male holotype and female

‘allotype’ of Nacaduba mallicollo markira Tite from San Cristobal, Solomon Islands, are

reassessed and found to belong to different species. Nacaduba samsoni sp. nov. is described

from Nendo (Santa Cruz group), San Cristobal and Rennell Islands.

Introduction

In a synonymic list of the genus Nacaduba Moore, Tite (1963) paid particular

attention to the south Pacific, describing several new taxa from the region,

including Nacaduba mallicollo markira Tite from the large island of San

Cristobal, at the eastern extremity of the Solomon archipelago. The taxon was

described from a short series, including the male holotype (Figs 1-2) and

female ‘allotype’ (Figs 3-4) collected by A. S. Meek at ‘Markira harbour’, on

the south coast of San Cristobal, in 1908.

The group of polyommatine lycaenid species which includes N. mallicollo

Druce (N. mallicollo, N. kurava Moore, N. berenice Herrich-Scháffer) is in

need of revision in the Solomon and New Hebrides archipelagos. Some

species are notoriously difficult to separate using wing markings and

identification is complicated by the fact that several species display marked

individual variation. For example, either sex of N. mallicollo may be

confused with corresponding sexes of N. kurava from some localities (but see

discussion below regarding the Santa Cruz Islands) and the male genitalia of

these species are similar. The genitalia of male N. berenice, with which N.

mallicollo might also otherwise be confused, are diagnostic. Tite (1963)

examined the genitalia of the N. m. markira holotype (Fig. 9), which

correspond closely to those of nominotypical N. m. mallicollo from Vanuatu.

The upperside of the female allotype of N. m. markira is similar in

appearance to the female holotype of N. m. mallicollo, but the underside

markings are quite dissimilar. Although Tite (1963) did not say so, the

unusual appearance of the female of this pair may have initially attracted

attention. The remaining specimens of the short type series (one male from

Vella Lavella and two females from San Cristobal and Santa Ana) appear to

conform to N. mallicollo.

As part of a study of Solomon Islands butterflies (Tennent 1998), the types of

N. m. markira were examined in 1996. It was concluded that the two

specimens may not be conspecific and that the unusually marked female may

86 Australian Entomologist, 2001, 28 (3)

be aberrant, although a lack of material made it difficult to pursue this

suspicion. A series of nine females, together with a single male, collected on

the island of Nendo in the Santa Cruz group in May 2000, suggested that the

male holotype of N. m. markira was not conspecific with the allotype female

and that the female was not aberrant but represented an undescribed species.

A further female was collected on Rennell I. in August 2000.

Many Nacaduba species have similar underside patterns, characterised by a

series of fine transverse lines. There is also usually a large subtornal spot

surrounded by orange and/or iridescent green or blue-green scales on the

hindwing underside. Differences between some species are minor but despite

marked sexual dimorphism on the upperside, underside markings are

generally of similar appearance in the sexes of the same species. Figures 1-4

illustrate type specimens of N. m. markira, in which significant underside

differences between the sexes (Figs 2, 4) may be observed. In particular, the

subtornal spot of the male is almost completely circled by pale orange and

bordered iridescent green distally (a common Nacaduba feature), whilst the

tornal spot of the female is boldly marked iridescent blue distally with no

trace of orange.

Nacaduba samsoni sp. nov.

(Figs 3-8, 10)

Nacaduba mallicollo markira; Tite, 1963: 82, pl. 1 (allotype 9); misidentification.

Types. Holotype È, SOLOMON ISLANDS: Santa Cruz group, Nendo Island, Lata to

Noipe, 60-140 m, 17.v.2000, W.J. Tennent (gen. prep. BMNH(V) 5974) (in The

Natural History Museum, London [BMNH]). Paratypes: 3 99, same data as holotype;

1 9, same locality, 3.v.2000; 3 99, same locality, 5.v.2000; 3 99, same locality,

9.v.2000; 1 9, Rennell I., Tinggoa and road 10 km east, 8.viii.2000, W.J. Tennent; 1 9

(‘allotype’ of Nacaduba mallicollo markira), San Cristobal, Makira harbour [south

coast], 1-8.v.1908, Meek (all BMNH).

Description. Male (Figs. 5-6) forewing length 16 mm; wing fringes dark

brown, clearly tipped white (uniform muddy brown in N. m. markira);

upperside bright mauve-blue (tinged pinkish in N. m. markira); underside

pale grey-brown, basal markings indistinct; median and postmarginal

markings white, spaces filled pale brown; submarginal area mainly white,

with prominent series of crescent-shaped brown spots; subtornal black spot

large, edged iridescent blue-green distally with no trace of orange (all other

Nacaduba species of the region, including N. mallicollo, have at least a trace

of orange markings associated with the subtornal spot). Genitalia (Fig. 10)

similar to N. mallicollo; valva with hooked apex, directed inwards,

approximately half the length of distal edge of valva (a slightly variable

feature in some associated Nacaduba species; in the male holotype of N. m.

markira [Fig. 9a] the hooked apex is significantly longer); distal edge with 7

(possibly 8) serrated ‘teeth’, larger than those of N. mallicollo; aedeagus

shorter, more squat.

Australian Entomologist, 2001, 28 (3) 87

Figs 1-8. Nacaduba species. (1-2) N. mallicollo markira, holotype male (San

Cristobal): (1) upperside, (2) underside; (3-4) N. samsoni, paratype female (San

Cristobal) [N. m. markira ‘allotype’]: (3) upperside, (4) underside; (5-6) N. samsoni,

holotype male (Nendo): (5) upperside, (6) underside; (7-8) N. samsoni, paratype

female (Nendo): (7) upperside, (8) underside.

88 Australian Entomologist, 2001, 28 (3)

Female (Figs 3-4, 7-8) upperside superficially similar to N. mallicollo;

upperside forewing with broad borders; median area pale blue, almost white,

broken by veins, heavily suffused shining blue basally (less white overall,

blue more dull in associated species); hindwing white, heavily suffused grey-

blue; submarginal and marginal markings prominent; underside highly

distinctive; fundamentally white; basal and median markings obscured;

submarginal and marginal markings prominent, similar to male; subtornal

spot similar to that of male. A female from Rennell is more heavily suffused

blue on the upperside and has more prominent underside markings.

Etymology. The new species is named for Chris Samson, in recognition of his

butterfly studies in the Solomon and New Hebrides archipelagos, including

the Santa Cruz group (Samson 1979, 1980).

Distribution. Solomon Islands: San Cristobal, Rennell and Nendo (Santa

Cruz).

Discussion

The author remained in the Santa Cruz group for several months in 2000 and

collected extensively on Nendo whilst waiting for transport to the more

remote islands of the group. The habits of N. samsoni were dissimilar to other

Nacaduba and associated species present. Females were uncommon and had a

deceptively slow flight. All were taken in flight or whilst feeding at the small

white flowers of Mikania micrantha (Asteraceae). Males were apparently

quite common, but extremely wary and could be seen in places where

Mikania vines covered trees at considerable height, flying around vines 20-30

metres above the ground. Even at a distance, their pale undersides were

distinctive and it was suspected, looking through binoculars, that this was the

male associated with the very pale females already collected. On the few

occasions when males were observed at lower levels they remained only

fleetingly before returning to higher vegetation. Only one male was eventually

collected.

Nacaduba samsoni is unlikely to be confused with any similar species on the

island of Nendo. N. kurava cruzens Tennent occurs on the island; this is a

distinctive subspecies with constant markings by comparison with subspecies

elsewhere (Tennent 2000). N. berenice has not been reported from the Santa

Cruz group. On Rennell, N. kurava has not been reported, whilst N. berenice

has been recorded only from a single pair similar to N. b. korene Druce,

collected in 1953 (Howarth 1962). The female of this pair is quite different in

phenotype to the female N. samsoni recorded here from Rennell. N. samsoni

may also occur on islands of Vanuatu, the type locality of N. mallicollo.

Female N. mallicollo are invariably more blue on the upperside than N.

samsoni, have at least some orange associated with the underside hindwing

tornal spot and are rarely as white overall on the under surface, although some

individuals may be difficult to separate.

Australian Entomologist, 2001, 28 (3) 89

9a 10d

10c

10a 10b

Figs 9-10. Nacaduba species, male genitalia. (9) N. mallicollo markira (BMNH slide

No. 24602; G.E.T 382), a, valvae (posterior view); b, aedeagus (lateral view); (10) N.

samsoni (BMNH(V) No. 5974); a, genitalia (aedeagus removed) (lateral view); b, left

valva (posterior view); c, right valva (posterior view, slightly angled); d, aedeagus

(lateral view).

ĝu Australian Entomologist, 2001, 28 (3)

The island of San Cristobal and its satellites (Ugi, Santa Ana, Santa Catalina)

have a higher proportion of endemic species and subspecies of butterfly taxa

than any other island of the archipelago. The male of N. m. markira is very

similar to nominotypical N. m. mallicollo from Vanuatu. Collection of further

material in due course will no doubt establish whether the name markira is

synonymous with N. m. mallicollo. As already intimated, a detailed revision is

required to fully resolve the identity and distribution of this closely associated

group of lycaenid butterflies.

Acknowledgments

Thanks are due to the Government of the Solomon Islands for continuing to

support the author’s fieldwork, which in the Santa Cruz group Was partially

funded by the Godman Exploration Fund (BMNH) and the Percy Sladen

Fund (Linnean Society of London). Roy Vickery, Department of Botany, The

Natural History Museum, kindly identified Mikania micrantha from a dried

specimen.

References

HOWARTH, T.G. 1962. The Rhopalocera of Rennell and Bellona Islands. The Natural History

of Rennell Island, British Solomon Islands 4: 63-83.

SAMSON, C. 1979. Butterflies (Lepidoptera: Rhopalocera) of the Santa Cruz group of islands,

Solomon Islands. Aurelian, Beckley 1(2): 1-19.

SAMSON, C. 1980. Butterflies (Lepidoptera: Rhopalocera) of the Santa Cruz group of islands,

Solomon Islands. [additional data]. Aurelian, Beckley 1(4): 2.

TENNENT, W.J. 1998. Biodiversity and biogeography of Solomon Islands butterflies. MSc

Thesis, University of Kent at Canterbury, unpublished.

TENNENT, W.J. 2000. Thirteen new butterflies from the Solomon Islands (Lepidoptera:

Lycaenidae). Butterflies 25: 9-22.

TITE, G.E. 1963. A synonymic list of the genus Nacaduba and allied genera (Lepidoptera:

Lycaenidae). Bulletin of the British Museum (Natural History), Entomology 13(4): 67-116.

Australian Entomologist, 2001, 28 (3): 91-96 91

THREE NEW HYPOCHRYSOPS C. 8 R. FELDER TAXA FROM THE

SOLOMON ISLANDS, INCLUDING A NEW SPECIES FROM THE

SANTA CRUZ GROUP (LEPIDOPTERA: LYCAENIDAE)

W. JOHN TENNENT

Biogeography and Conservation Laboratory, Department of Entomology, The Natural History

Museum, London SW7 5BD, UK

(address for correspondence: 38 Colin McLean Road, Dereham, Norfolk NR19 2RY, England)

Abstract

Three new taxa of Hypochrysops C. & R. Felder are described from the Solomon Islands: H.

architas marie subsp. nov. from the New Georgia group, H. julie sp. nov. from the eastern Santa

Cruz group and H. scintillans jamesi subsp. nov. from New Georgia.

Introduction

Hypochrysops C. & R. Felder, 1860, occurs from Malaysia and Thailand to

the Solomon Islands and contains approximately 60 species. Only one species

is confirmed as occurring west of Wallace’s Line (Sands 1986) and the genus

occurs principally from the Moluccas to the Solomons. Sands (1986) divided

Hypochrysops into 20 species-groups, incorporating three of the four

Solomons species (H. architas Druce, 1891, H. scintillans (Butler, 1882) and

H. taeniatus Jordan, 1908) in the anacletus species-group and the remaining

species, H. alyattes Druce, 1891, in the hippuris species-group. The most

easterly member of the genus previously known is H. taeniatus, confined to

the island of San Cristobal. Of the remaining Solomons species, H. architas

occurs in three subspecies from Bougainville to Guadalcanal and Malaita, H.

scintillans occurs on Guadalcanal and Florida (see discussion), and H.

alyattes has been reported from the New Georgia group (Gizo), Santa Isabel,

Guadalcanal, Florida and Malaita. During recent fieldwork in the Solomon

Islands, undescribed subspecies of H. architas and H. scintillans were

collected on New Georgia and an undescribed species was discovered on the

Santa Cruz island of Nendo.

Hypochrysops architas marie subsp. nov.

(Figs 1-4, 13)

Types. Holotype &, SOLOMON ISLANDS: New Georgia group, New Georgia, west,

c. 3 km north of Munda, 100 m, 2.xi.1997, W.J. Tennent (gen. prep. BMNH (V) 4881

(JT349)) (in The Natural History Museum, London [BMNH]). Paratypes: 1 9, Gizo,

xi.1903, Meek; 1 07, Rendova, ii.1904, Meek; 1 9, same data as holotype, 4.xi.1997;

1 9, Rendova, north coast, Mendali Point, 0-160 m, 27.iv.2001, W.J. Tennent (all

BMNH); 1 9, Gizo, 0-140 m, xii.1980, N.L.H. Krauss (Bernice P. Bishop Museum,

Honolulu).

Description. Similar in appearance to other subspecies of H. architas. Male

(Figs 1-2) forewing length 16.5 mm; upperside clear dark blue, similar to H.

narcissus eucletas C. & R. Felder, 1865, from Indonesia (purple or purple-

blue in H. a. architas and H. a. cratevas Druce, 1891); upperside forewing

se Australian Entomologist, 2001, 28 (3)

blue area slightly reduced; underside bands red (orange-red in other H.

architas subspecies). Genitalia (Fig. 13) similar to H. a. architas. Female

(Figs 3-4) similar to other H. architas subspecies on both surfaces.

Etymology. Most of our knowledge of Solomon Islands butterflies stems from

the work of Albert Stewart Meek and Charles Morris Woodford. This taxon is

named after Marie, Albert Meek’s daughter, whom the author was privileged

to meet in Brisbane in 1997, aged 92.

Hypochrysops julie sp. nov.

(Figs 5-8, 11)

Types. Holotype ©, SOLOMON ISLANDS: Santa Cruz group, Nendo Island, ca 4 km

(by road) south of Lata, 160 m, secondary growth on edge of village garden,

11.x.1997, W.J. Tennent (in BMNH). Paratypes: 1 0°, 1 9, same data as holotype; 5

CO", 1 9, same data, 10.x.1997 (o10' including gen. preps. BMNH (V) 4879 & 4880);

3 0, 1 9, same data, 13.x.1997; 1 O”, same data, 14.x.1997; 1 9, Nendo, south-west

central, Forestry camp, 140-160 m, 28.iv.2000, W.J. Tennent; 6 0", 16 99, Nendo,

Lata to Noipe, 60-140 m, 5.v.2000, W.J. Tennent; 17 (0, 6 99, same data, 9.v.2000;

5 OO", 2 99, same data, 12.v.2000; 4 (0, same data, 17.v.2000; 1 o”, 3 99, Santa

Cruz group, Vanikoro, main island, eastern coastal strip, 2.iv.2000, W.J. Tennent; 1 9,

Vanikoro, north-east of Lale village, SL-100 m, 4.iv.2000, W.J. Tennent; 1 9,

Vanikoro, Lale village gardens, 20-140 m, 5.iv.2000, W.J. Tennent; 1 ?, same data,

6.iv.2000 (all BMNH).

Description. Intermediate in appearance between H. architas and H.

taeniatus. Male (Figs 5-6) forewing length 14 mm; upperside similar to H.

taeniatus, ground colour dull purple-blue; upperside forewing apex thinly

lined black; underside similar to H. architas, underside forewing markings

less distinct; underside hindwing median, submedian and postbasal bands

darker orange than in H. taeniatus (red in H. architas), broadly bordered

iridescent emerald green (pale green in H. architas); thin, broken black

` postmedian line, independent from marginal markings. Genitalia (Fig. 11)

typical of anacletus group; posterior of sociuncus deeply indented dorsally;

valva similar to H. taeniatus, less deeply indented anteriorly. Female (Figs 7-

8) upperside dark brown; upperside forewing with indistinct pale discal patch,

tinged violet-blue distad; basal blue suffusion characteristic of H. architas

and H. taeniatus lacking; upperside hindwing unmarked; underside markings

similar to male, metallic green markings less extensive.

Etymology. This attractive new species is named after the author’s wife Julie,

who continues to support his long periods in the field.

Hypochrysops scintillans jamesi subsp. nov.

(Figs 9-10, 12)

Types. Holotype &', SOLOMON ISLANDS: New Georgia group, New Georgia, west,

c. 3 km north of Munda, 100 m, 4.xi.1997, W.J. Tennent (in BMNH). Paratype €,

same data as holotype (gen. prep. BMNH (V) 4882 (JT348)) (BMNH).

Australian Entomologist, 2001, 28 (3) 93

Figs 1-10. New Hypochrysops taxa. (1-4) H. architas marie: (1) male upperside

(Rendova paratype); (2) male underside (holotype); (3) female upperside (Rendova

paratype); (4) female underside (New Georgia paratype). (5-8) H. julie: (5) male

upperside (Vanikoro paratype); (6) male underside (holotype); (7) female upperside

(Nendo paratype); (8) female underside (Nendo paratype). (9-10) H. scintillans

jamesi: (9) male upperside (paratype); (10) male underside (holotype). Scale = 1 cm.

94 Australian Entomologist, 2001, 28 (3)

11b

11d

N Me

12a “« 12b

12c 12d

k v TS Hypochrysops species, male genitalia: a, genitalia; b, sociuncus; c, valva

(left); d, aedeagus. (11) H. julie; (12) H. scintillans jamesi; (13) H. architas marie.

Australian Entomologist, 2001, 28 (3) 95

Description. Male (Figs 9-10) similar to H. s. constancea D’ Abrera, 1971;

larger, forewing length 20 mm; upperside blue darker, less purple; upperside

forewing with distal edge of blue discal patch prominently sagittate;

upperside hindwing blue less extensive at apex than H. s. constancea;

underside red and iridescent gold markings bold; underside ground colour

olive-brown (yellow-brown in H. s. constancea); underside forewing basal

area suffused orange (lacking in the only male of H. s. constancea seen).

Genitalia (Fig. 12) similar to H. s. scintillans. Female unknown.

Etymology. Charles Morris Woodford, the first Resident Commissioner of the

Solomon Islands, was the first person to collect butterflies systematically

there (Tennent 1999). This taxon is named after Jim Woodford, Charles’

great-nephew and traveller/adventurer in his own right, who was generous in

his hospitality during the author’s field visits to the Solomons between 1997

and 2000.

Discussion

The Solomon archipelago, which includes the large island of Bougainville

(politically part of Papua New Guinea), is a significant area of endemism and

New Georgia group populations of a number of butterfly species are distinct

from populations found on adjacent island groups. Discovery of a

Hypochrysops species in the Santa Cruz group, politically part of the

Solomon Islands but faunistically also allied to the islands of Vanuatu to the

south, represents a significant easterly extension of the range of this genus.

Aside from H. s. jamesi, described above, the only subspecies of H.

scintillans known from the Solomon Islands is H. s. constancea. The female

holotype of the latter taxon is from Guadalcanal and was illustrated by

D' Abrera (1971), but appears not to have been labelled as such until Sands’

(1986) revision. It now bears an additional label marked ‘Holotype,

Hypochrysops scintillans constantacea [sic], examined by D. Sands, 1984’.

No paratypes were designated by D’Abrera but, judging from material

available at that time in the BMNH, they comprised a second female with

similar data to the holotype, a further female labelled “Tugela (Woodford)’

and a male labelled ‘Gela, Woodford’, all of which have now been labelled.

‘Gela’, or Nggela, is a name for the island now more usually known as

Florida. The locality known as ‘Tugela’ is more problematic and it is not

certain that an island or place of this name exists, or has ever existed.

Although the name appears on several Solomons labels, usually (but not

exclusively) associated with Woodford material and often (but not in this

case) with Guadalcanal, it does not appear on any map, nor does it appear in

the comprehensive Pacific gazetteers in use at the time of Woodford. The

name is not mentioned in any of the numerous publications of Woodford,

including a book (Woodford 1890) in which he gave an account of his life in

the Solomons and the places he visited. The Solomon Islands Government

Archivist in Honiara has no record of the name (Mr Ishmail Avui, pers.

96 Australian Entomologist, 2001, 28 (3)

comm.). Although less likely, it is possible that the name is a corruption of

‘Tulagi’, a small island in the Florida group and the pre-Second World War

national capital of the Solomon Islands.

The distribution of H. s. constancea was given by D’Abrera (1971, 1978,

1990) as ‘Guadalcanal, Tugela’ and, regardless of whether ‘Tugela’ exists or

existed on Guadalcanal, or whether it in fact refers to Tulagi, the known

distribution of H. s. constancea may be taken as Guadalcanal and Florida.

Acknowledgments

Thanks to Dr Scott Miller for access to the Bernice P. Bishop Museum

collections, Honolulu. Mr Moses Biliki, Ministry of Forests, Environment and

Conservation, Honiara, supported the author’s research and the Ministry of

Education and Human Resources Development, Honiara, issued permits. Dr

Don Sands, Brisbane, commented on New Georgia phenotypes of H. architas

in the BMNH prior to the discovery of further material. The author’s first

field visit to the Solomon Islands in 1996 was partially funded by the

Exploration Board of Imperial College of Science, Technology and Medicine,

London, The Linnean Society, London (Percy Sladen Fund) and the Royal

Entomological Society, London. Significant funding for this and subsequent

field visits was provided by the Trustees of the Godman Exploration Fund

and the Percy Sladen Fund.

References

D’ABRERA, B. 1971. Butterflies of the Australian Region. Lansdowne, Melbourne; 415 pp.

D’ABRERA, B. 1978. Butterflies of the Australian Region. [2nd edition]. Lansdowne,

Melbourne; 415 pp.

D’ABRERA, B. 1990. Butterflies of the Australian Region. [3rd (revised) edition]. Hill House,

Melbourne; 416 pp.

SANDS, D.P.A. 1986. A revision of the genus Hypochrysops C. & R. Felder. Entomonograph

` 7: 1-116.

TENNENT, W.J. 1999. Charles Morris Woodford C.M.G. (1852-1927): Pacific adventurer and

forgotten Solomon Islands naturalist. Archives of Natural History 26(3): 419-432.

WOODFORD, C.M. 1890. A naturalist among the head-hunters, being an account of three

visits to the Solomon Islands in the years 1886, 1887 and 1888. George Philip & Son, London;

249 pp.

Australian Entomologist, 2001, 28 (3): 97-104 97

THE CRAZY ANT ANOPLOLEPIS GRACILIPES (SMITH)

(HYMENOPTERA: FORMICIDAE) IN EAST ARNHEM LAND,

AUSTRALIA

G.R. YOUNG!, G.A. BELLIS”, G.R. BROWN! and E.S.C. SMITH!

! Entomology Branch, Northern Territory Department of Primary Industry and Fisheries,

GPO Box 990, Darwin, NT 0801

?Northern Australia Quarantine Strategy, GPO Box 990, Darwin, NT 0801

(Email: glenn. bellis @ affa. gov.au)

Abstract

Anoplolepis gracilipes (Smith) was first recorded from the Australian mainland in East Arnhem

Land, Northern Territory, in May 1990. In a survey during November 1999, it was found over

five river drainage systems covering an area of approximately 2,500 km? but was mainly

confined to thin strips of monsoon rainforest bordering perennial springs and streams. It was

found only once in dry open Eucalyptus woodland. Highest populations were found in a

disturbed habitat, although the ant was absent from Nhulunbuy town and aboriginal

communities. The large area infested with A. gracilipes suggests that an eradication campaign

would be extremely difficult.

Introduction

Crazy ant, Anoplolepis gracilipes (Smith) (= longipes (Jerdon)), is a tramp

species thought to be native to Africa (Way and Khoo 1992). The. ant has

been spread by commerce throughout East Africa, Asia and the Pacific

(Lewis et al. 1976). A. gracilipes is a pest of agricultural, domestic and

natural environments (Lewis et al. 1976, Haines et al. 1994, Rao and Veeresh

1994) and, like many tramp species, forms unicolonial, polygynous colonies

(Reimer 1994).

Crazy ant is frequently a pest of orchard crops because it nurtures sap-feeding

insects. Copious amounts of honeydew produced by sap-sucking insects

results in the growth of sooty mould on the leaves of fruit trees (Haines and

Haines 1978a, Haines et al. 1994, Young 1996b). The ant often encourages

pest species indirectly by harassing predators and parasites of the pests

(Young 1996a). Additionally, A. gracilipes excavates around the roots of

crops such as sugar cane and coffee, undermining the roots and causing the

plants to collapse (Lewis et al. 1976, Haines and Haines 1978a, Rao and

Veeresh 1994).

A. gracilipes is primarily a scavenger and will enter houses in search of food,

which has led to the ant's reputation as a household pest (Lewis et al. 1976).

It will also pester confined domestic animals, such as poultry (Haines and

Haines 1978b, Haines et al. 1994).

The greatest impact of crazy ant is on the ecology of natural environments

(Lewis et al. 1976, Haines and Haines 1978a, Haines et al. 1994). There are

numerous reports of crazy ants displacing other invertebrate species

(especially ants and spiders), forcing vertebrate species to vacate infested

areas, attacking the young of nesting birds and altering the floral composition

98 Australian Entomologist, 2001, 28 (3)

(Lewis et al. 1976, Haines and Haines 1978a, Gillespie and Reimer 1993,

Rao and Veeresh 1994, O’Dowd et al. 1999). A recent example is on

Christmas Island, where A. gracilipes is having a detrimental effect on

rainforest vegetation, populations of the red land crab Gecarcoidea natalis

Pocock and nesting sea birds (O'Dowd et al. 1999).

A. gracilipes was first recorded from mainland Australia on the Gove

Peninsula, East Arnhem Land, Northern Territory, following a survey by the

Parks and Wildlife Commission of the Northern Territory during May 1990

(Reichel and Andersen 1996, Shattuck 1999). The collection locality was at

Balkbalkbuy, 77 km south-west of Nhulunbuy airport, on the Katherine to

Nhulunbuy road (N. Gambold, pers. comm.) (Fig. 1). Two of us (GAB and

GRY) confirmed the presence of the ant at this site during October 1999. In

view of the importance of A. gracilipes as an agricultural and environmental

pest outside mainland Australia, it was decided to determine the distribution

of the ant in East Arnhem Land and consequently the feasibility of an

eradication campaign.

The vegetation of the Gove Peninsula mainly consists of tall, open woodland

dominated by Eucalyptus tetrodonta and E. minata (Lynch and Wilson 1998).

The woodland is interspersed with small areas of monsoon rainforest

associated with perennial springs and streams (Wilson et al. 1990, Russell-

Smith 1991).

Materials and Methods

During three days of investigation, the Gove Peninsula was searched on foot,

by vehicle and quad bike. Preference was given to accessible areas on or near

“roads and tracks, especially near the upper reaches of watersheds, permanent

watercourses and around aboriginal communities. Each individual inspection

site had the GPS coordinates recorded and was investigated for 0.5 man-hours

or until crazy ants were detected. Ants were visually located by raking leaf

litter with sticks, searching where sooty mould was present on plants or by

placing a small quantity of tuna-based cat food in 20 cm lengths of hollow

bamboo. Representative samples were taken from each site where crazy ants

were present.

Since personal experience of this ant in overseas countries had demonstrated

the requirement for moisture and suitable nesting conditions, investigations

were concentrated in areas where water, either above or below ground, was

accessible. During the survey, accessible water was generally restricted to

watercourses. The Eucalyptus woodland, which covers the great majority of

the watershed areas, was affected by the prolonged absence of rain and the

annual wild fires prevalent in this region.

Forty-nine sites were sampled from a range of habitats on the Gove Peninsula.

Eleven of these were away from creek lines in Eucalyptus woodland.

Australian Entomologist, 2001, 28 (3) 99

Fig. 1. Top End of the Northern Territory showing the Nhulunbuy to Katherine road

and areas searched for A. gracilipes (Bl = presence; A = absence).

Results

Crazy ant was found in five drainage systems on the Gove Peninsula covering

an area of approximately 2,500 km? (Fig. 2). The ant was absent from the port

and town of Nhulunbuy as well as the nearby Yirrkala community. It was

abundant around the first detection point at Balkbalkbuy, which is a

permanent watercourse bordered by a thin strip of monsoon rainforest up to 5

m wide. Balkbalkbuy is used as a camping area and for parking earthmoving

equipment. The resulting refuse and disturbance had encouraged the ant by

providing shelter and nesting sites. Downstream from Balkbalkbuy, the ant

was present and abundant for at least 700 m along the creek and for a further

200 m along a larger adjoining tributary. It was by far the dominant ant

species in parts of the monsoon rainforest along the creek and appeared to

have displaced native species of ant, including the green ant Oecophylla

smaragdina (Fabricius). While crazy ants were found along the creek line, the

ant had not invaded a dense patch of monsoon rainforest surrounding a

spring, which feeds the creek at Balkbalkbuy.

100 Australian Entomologist, 2001, 28 (3)

At Balkbalkbuy, A. gracilipes formed large, interconnected soil colonies.

Nests containing brood were also found under discarded car tyres and rubber

mats. Tuna proved highly attractive to the ant with workers swarming around

the bait within ten minutes of placement in the bamboo tubes. Additionally,

the ant rapidly colonised the tubes; workers, alate and dealate queens, alate

males and brood were present in tubes left out overnight.

The ant was less abundant in the other four drainage systems, being patchily

distributed in shaded areas along creek banks and in the upper reaches of

drainage systems. In one instance nests were found in a clay and shale creek

bank above the normal wet season water level.

There appeared to be an association between the ant and monsoon rainforest

growing along the creeks. This vegetation provided shade and leaf litter,

creating a favourable habitat for the ant. The ant was not found in dry open

woodland away from creek lines, except in one instance where a colony was

found nesting in disturbed rock and soil beside the Katherine to Nhulunbuy

road, possibly indicating that the ant had been transported there by

earthmoving equipment or other vehicles.

Workers were observed climbing the trunks of trees and foraging over foliage

but it was not apparent whether this indicated arboreal nests or ants searching

for either sap-sucking homopterans or nectar. On one occasion A. gracilipes

was observed tending Saissetia sp. (Hemiptera: Coccidae) on Buchanania

obovata (Anacardiaceae), the leaves of which were covered in sooty mould.

However, in other localities where the ant was found sooty mould was not

detected on the vegetation.

Discussion

Anoplolepis gracilipes was found in shaded, moist areas of monsoon

rainforest with a year-round layer of leaf litter and was generally absent from

open Eucalptus woodland. Haines and Haines (1978b) in the Seychelles and

Young (1996b) in Papua New Guinea showed that, while the ant would

forage over 24 hours in tropical climates, maximum foraging activity

occurred at temperatures ranging from 26-30°C and relative humidities from

65-90%. Rao and Veeresh (1991) observed maximum foraging activity at

temperatures between 24 and 28°C. Temperatures in open woodland are often

>34°C, free moisture is unavailable for 4-5 months of the year and the leaf

litter is burnt during annual dry season fires. These conditions make the

woodland an unfavourable habitat for the ant during the dry season.

Conversely, the more permanent leaf litter and mulch layer of the monsoon

rainforest (Bowman and Wilson 1988) provides A. gracilipes with a cooler

and more stable habitat. The failure of crazy ant to colonise dense monsoon

rainforest near Balkbalkbuy is unexplained.

Australian Entomologist, 2001, 28 (3) 101

_Nhulunbuy

ax € Yirrkale

= “Balkbalkbuy

ants present

a no

H yes

ucc eeu d SEA š 20 Kilometres a

Fig. 2. Gove Peninsula, East Arnhem Land, showing the Nhulunbuy - Katherine road,

five drainage systems and survey sites, indicating the presence or absence of A.

gracilipes. Drainage systems: 1 = Goromuru River; 2 = Cato River; 3 = Wonga Creek;

4 = Balkbalkbuy Creek; 5 = Ngabinya Creek.

While both alate queens and males are known to fly, there is no evidence of

mating flights and it appears that colonies reproduce by budding (Haines and

Haines 1978b, Haines et al. 1994). During the wet northwest monsoon,

conditions are probably favourable for sexuals and workers carrying brood to

walk across open Eucalyptus woodland, enabling them to colonise new areas

of monsoon rainforest. The dry season would isolate these new colonies from

the original one. As demonstrated by the rapidity of colonising bamboo tubes,

the ant can be spread readily by human activities. These factors could explain

the patchy distribution of A. gracilipes on the Gove Peninsula.

102 Australian Entomologist, 2001, 28 (3)

Population density of A. gracilipes was greatest at Balkbalkbuy where the ant

was able to construct large nests under refuse. The rapid colonisation of

bamboo tubes suggests that the populations of A. gracilipes are limited by

nesting sites in areas where the environment is favourable to the ant. In the

Seychelles, Haines and Haines (1978b) concluded that population size was

probably limited by the availability of food and nesting sites.

From work in the Seychelles and Christmas Island, it may be assumed that, on

the Gove Peninsula, A. gracilipes obtains its protein by feeding on

invertebrates inhabiting leaf litter in monsoon rainforests (Haines et al. 1994,

O'Dowd et al. 1999). It is not immediately apparent where the ant sources

carbohydrate, although it is likely to be either nectar or other plant exudates

(Haines et al. 1994, Young, 1996a). Contrary to observations of ants tending

homopterans on Christmas Island (O’Dowd et al. 1999), the ant was observed

to tend honey-dew producing homopterans on only one occasion during this

survey.

If A. gracilipes were to spread to tropical horticultural production areas the

ant could damage sugar cane and tree crops as a result of excavating around

root systems, encouraging sap-feeding insects and reducing the effectiveness

of parasites and predators of pest species.

Monsoon rainforest occurs throughout north and north-western Australia as

isolated patches (typically 1-10 ha), usually associated with permanent water,

surrounded by vast areas of savanna woodland (Russell-Smith 1991). These

rainforests have a very significant ant fauna (Reichel and Andersen 1996),

which is an important component of biodiversity in the Northern Territory

(Hoffmann et al. 1999). The exotic ant Pheidole megacephala (F.) has

significantly reduced the richness and abundance of native ants and other

invertebrates in a rainforest patch at Howard Springs near Darwin (Hoffmann

et al. 1999). In view of observations made on the Gove Peninsula, A.

gracilipes can be regarded as an equally serious threat to the invertebrate

fauna of monsoon rainforests in northern Australia.

The detection of the ant in 1990 on an isolated creek bank more than 80 km

from the nearest town and the subsequent discoveries of populations spread

over 2,500 km?, suggest that the ant has been established in the area for at

least several decades. Furthermore, its presence along creeks far removed

from human habitation suggests that the initial introduction could go back to

mining exploration in the last 30 or 40 years, construction and military

activities during the Second World War or even to the annual visits of

Maccassan traders more than a century ago. There are undoubtedly

populations of A. gracilipes on the Gove Peninsula that remain undetected

and in view of the large and inaccessible area known to be infested, an

eradication campaign would be very difficult.

Australian Entomologist, 2001, 28 (3) 103

Further dispersal of A. gracilipes by earth moving equipment and other

vehicles from its current range is a continuing possibility and processes

should be put in place to contain the ant in the Gove Peninsula.

Acknowledgments

We are grateful for the generous support and assistance given us by M. Storrs,

Northern Land Council; K. Leitch, Nanakiya and Mangatjay, Dhimurru Land

Management Aboriginal Corporation; N. Gambold, Central Land Council;

officers of the Laynhapuy Homelands Resource Centre, the Yirkalla Dhanbul

Landcare Group and the Nhulunbuy Corporation. J. Donaldson (Qld

Department of Primary Industries, Brisbane) identified the coccid.

References

BOWMAN, D.M.J.S. and WILSON, B.A. 1988. Fuel characteristics of coastal monsoon forest,

Northern Territory, Australia. Journal of Biogeography 15: 807-817.

GILLESPIE, R.G. and REIMER, N. 1993. The effect of alien predatory ants (Hymenoptera:

Formicidae) on Hawaiian endemic spiders (Araneae: Tetragnathidae). Pacific Science 47: 21-

33.

HAINES, LH. and HAINES, J.B. 1978a. Pest status of the crazy ant, Anoplolepis longipes

(Jerdon) (Hymenoptera: Formicidae) in the Seychelles. Bulletin of Entomological Research 68:

627-638.

HAINES, LH. and HAINES, J.B. 1978b. Colony structure, seasonality and food requirements of

the crazy ant, Anoplolepis longipes (Jerd.), in the Seychelles. Ecological Entomology 3: 109-

118.

HAINES, LH., HAINES, J.B. and CHERRETT, J.M.1994. The impact and control of the crazy

ant, Anoplolepis longipes (Jerd.), in the Seychelles. Pp. 206-218, in D.F. Williams (ed.), Exotic

ants: biology, impact and control of introduced species. Westview Press, Boulder, Colorado,

USA.

HOFFMANN, D.H., ANDERSEN, A.N. and HILL, G.G.E. 1999. Impact of an introduced ant on

native rainforest invertebrates: Pheidole megacephala in monsoonal Australia. Oecologia 120:

595-604.

LEWIS, T., CHERRETT, J.M., HAINES, I., HAINES, J.B. and MATHIAS, P.L. 1976. The

crazy ant (Anoplolepis longipes (Jerd.) (Hymenoptera: Formicidae)) in the Seychelles, and its

control. Bulletin of Entomological Research 66: 97-111.

LYNCH, B.T. and WILSON, P.L. 1998. Land Systems of Arnhem Land. Technical report No.

R97/1, Department of Lands, Planning and Environment, Palmerston Northern Territory,

Australia.

O’DOWD, D.J., GREEN, P.T. and LAKE, P.S. 1999. Status, impact and recommendations for

research and management of exotic invasive ants in Christmas Island National Park.

Unpublished report to Environment Australia, Canberrra.

RAO, N.S. and VEERESH, G.K. 1991. Nesting and foraging habits of crazy ant Anoplolepis

longipes (Jerdon) (Hymenoptera: Formicidae). Environment and Ecology 9: 670-677.

RAO, N.S. and VEERESH, G.K. 1994. Bio-ecology and management of crazy ant Anoplolepis

longipes (Jerdon) - a review. Agricultural Reviews 15: 182-194.

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REICHEL, H. and ANDERSEN, A.N. 1996. The rainforest ant fauna of Australia's Northern

Territory. Australian Journal of Zoology 44: 81-95.

REIMER, N.J. 1994. Distribution and impact of alien ants in vulnerable Hawaiian ecosystems.

Pp. 11-22, in D.F. Williams (ed.), Exotic ants: biology, impact and control of introduced

species. Westview Press, Boulder, Colorado, USA.

RUSSELL-SMITH, J. 1991. Classification, species richness, and environmental relations of

monsoon rain forest in northern Australia. Journal of Vegetation Science 2: 259-278.

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

Collingwood, Australia, 226 pp.

WAY, M.J. and KHOO, K.C. 1992. Role of ants in pest management. Annual Review of

Entomology 37: 419-503.

WILSON, B.A., BROCKLEHURST, P.S., CLARK, M.J. and DICKINSON, K.J.M. 1990.

Vegetation Survey of the Northern Territory, Australia. Technical report No. 49, Conservation

Commission of the Northern Territory, Australia.

YOUNG, G.R. 1996a. An association between the crazy ant Anoplolepis longipes (Jerdon)

(Hymenoptera: Formicidae) and the coconut spathe moth, Tirathaba rufivena (Walker)

(Lepidoptera: Pyralidae) on coconut palms in the Morobe province of Papua New Guinea. 1.

Surveys to determine the extent of crop loss and the incidence of natural enemies of the moth.

Papua New Guinea Journal of Agriculture, Forestry and Fisheries 39; 1-6.

YOUNG, G.R. 1996b. The crazy ant, Anoplolepis longipes (Jerdon) (Hymenoptera: Formicidae)

on coconut palms in New Guinea. Papua New Guinea Journal of Agriculture, Forestry and

Fisheries 39: 10-13.

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THE AUSTRALIAN

Entomologist

Volume 28, Part 3, 25 September 2001

CONTENTS

BRABY, M.F. and ARMSTRONG, J.

A note on the larval food plants of Grapbium weiskei (Ribbe)

(Lepidoptera: Papilionidae) in Papua New Guinea.

EWART, A.

Emergence patterns and densities of cicadas (Hemiptera: Cicadidae)

near Caloundra, south-east Queensland.

TENNENT, W.J.

What is Nacaduba mallicollo markira Tite? A new species of Nacaduba

Moore from the Solomon Islands (Lepidoptera: Lycaenidae).

TENNENT, W.J.

Three new Hypochrysops C. & R. Felder taxa from the Solomon Islands,

including a new species from the Santa Cruz group (Lepidoptera: Lycaenidae).

YOUNG, G.R., BELLS, G.A., BROWN, G.R. and SMITH, E.S.C.

The crazy ant Anoplolepis gracilipes (Smith) (Hymenoptera: Formicidae) in

East Arnhem Land, Australia.

ISSN 1320 6133

Ente Molo GU .