THE AUSTRALIAN ENTOMOLOGIST

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Cover: Oribius destructor is one of about 50 species of small, flightless beetles

within the genus Oribius. Oribius destructor Marshall is a major pest of horticulture

in the highlands of Papua New Guinea and is particularly damaging to citrus, apples,

capsicums, strawberries and avocados. Damage is caused by the feeding of the adults,

causing leaf shot-holing, stem and fruit scarring, and branch dieback. Illustration by

Amy Carmichael.

Australian Entomologist, 2008, 35 (1): 1-6 l

DISTRIBUTION AND HOST PLANT RECORDS OF

AUSTROPLATYPUS INCOMPERTUS (SCHEDL)

(COLEOPTERA: CURCULIONIDAE: PLATYPODINAE)

DEBORAH S. KENT

Forest Resources Research, New South Wales Department of Primary Industries, Science and

Research, PO Box 100, Beecroft, NSW 2119 (Email: Deborah. Kent(üdpi.nsw.gov.au)

Abstract

The known distribution of Austroplatypus incompertus (Schedl) is extended north to Dorrigo and

west to Styx River State Forest near Armidale, New South Wales. Additionally, some previous

discontinuities in the distribution are filled in with records from the central coast of New South

Wales, Mt Wilson west of Sydney and Mogo and Bodalla on the south coast. The list of known

host plants of A. incompertus is increased to 19 species by the inclusion of Eucalyptus

agglomerata, E. andrewsii, E. cameronii, E. laevopinea and E. resinifera, all economically

important timber species. It is clear from the present study that the current distribution of A.

incompertus represents only a small subset of the range of the known hosts and it is therefore

probable that the importance of this beetle as a potential pest has been underestimated.

Introduction

The majority of Platypodinae attack dead, dying or damaged trees and most

attack hosts from a wide taxonomic range of plants (i.e. are polyphagous).

The tropical platypodine Crossotarsus externedentatus (Fairmaire), for

example, attacks over 100 species in over 60 genera (Beeson 1941). In

general, tropical platypodines attack a greater range of hosts than do

temperate species (Beaver 1979), a trend they share with the Scolytinae. At

first glance this trend would appear to conflict with the fact that there are

many more tropical platypodines than temperate ones (Wood 1993), because

a greater trend towards a reduced host range, as a means of lessening host

overlap and hence competition, would be expected where there is greater

species diversity. It is likely that the large host range in the tropics is related

to the greater plant species diversity and higher decomposition rates of

tropical forests, making host specialisation more difficult due to the lack of

suitable host material (Beaver 1989). However, not all tropical platypodines

are polyphagous and high host specificity can sometimes occur, as for

example in the following species which only attack living trees. Thus, at the

extreme end, Trachyostus ghanaensis Schedl from West Africa is restricted

to just one species, Triplochiton scleroxylon K. Schum. [Sterculiaceae]

(Roberts 1960), while the Malaysian species Dendroplatypus impar (Schedl)

is restricted to the genus Shorea [Dipterocarpaceae] (Browne 1961).

Similarly, Austroplatypus incompertus (Schedl) attacks only living members

of the genus Eucalyptus [Myrtaceae] (Campbell 1969, Browne 1971, Harris

et al. 1973, 1976, Wright and Harris 1974). A. incompertus causes visual

and/or structural defects to timber harvested from these trees. The former

results from the beetle boring and consists of black-stained holes, while the

latter results from weakening of the timber due to fungal attack. Attacks by

the beetle have been recorded only from the coast and tablelands of New

2 Australian Entomologist, 2008, 35 (1)

South Wales and eastern Victoria. Unfortunately, the latest list of known host

species for the beetle (Harris et al. 1976) contains some anomalies, does not

reflect current best knowledge and requires updating.

Given that A. incompertus is a pest species, an incomplete knowledge of its

host plants has the potential to cause severe problems in the control or

management of this beetle. There is little point in attempting, for example, to

limit the numbers of a pest by applying a management strategy to those pests

inhabiting a known host if unknown hosts nearby act as a source of

reinfestation. In a similar vein, while it is known that the distribution of

platypodines must, of necessity, be restricted to the geographic range of their

hosts, there has never been a serious attempt to define the distribution of A.

incompertus and the extent of the problem posed by its attacks is unknown.

The aim of this study was to compile a list of currently known hosts and plot

the present distribution of the species.

Material

Host lists and distributions are based on data obtained from the literature and

the following collections: ANIC - Australian National Insect Collection,

Canberra; AM - Australian Museum, Sydney; NMV - National Museum of

Victoria, Melbourne; NHM - The Natural History Museum, London and

FCNI - Forestry Commission of NSW Insect Collection, Sydney.

Results

Host records

Originally, it was thought that Austroplatypus incompertus was relatively

restricted in its host range, attacking only species of Eucalyptus belonging to

the informal subgenus Monocalyptus (Kent and Simpson 1992). However, as

a result of this study the host range now stands at 19 species, including five

new records. The author collected adults and larvae from E. agglomerata, E.

andrewsii, E. cameronii and E. laevopinea during fieldwork. A single adult

collected from £. resinifera was found in the FCNI collection. Table 1 lists

all hosts for which there are any records — including observations made

during this study, specimens in collections and references in the literature. A

single reference to Eucalyptus eugenioides Sieb. ex Spreng. (Wright and

Harris 1974) is either an error or an exotic planting as the given locality is

outside the range of this eucalypt species; therefore it has not been included

in the table. As a result of taxonomic revision (Chippendale 1976), the

reference to E. scabra Dum.-Cours. (Campbell 1969) has been included

under E. globoidea Blakely and E. gigantea Hook. f. (Browne 1971) under £.

delegatensis R.T. Baker.

It is worth noting that the current list of known hosts reflects a bias towards

commercial timber species, since the beetle has been most studied in these

species. This, combined with the fact that five new host species were

recorded during the current study alone, suggest that the current host list is

Australian Entomologist, 2008, 35 (1)

probably not exhaustive and it is quite likely that further work may identify

other host species.

Table 1. Host records for A. incompertus (all Eucalyptus species [Myrtaceae]).

Host Source of host records

Scientific name (Subgenus!) (Common Literature? Specimens

name”)

E. (M.) agglomerata (Blue leaved Collected this study (*)

stringybark)

E. (M.) andrewsii (New England Collected this study (*)

blackbutt)

E. (M.) baxteri (Brown stringybark) 1,3

E. (S.) botryoides (Southern WS

mahogany)

E. (M.) cameronii (Diehard Collected this study (*)

stringybark)

E. (M.) consideniana (Yertchuk) 123,

E. (M.) delegatensis (Alpine ash) 1,2,3,4,5 FCNI

E. (M.) dives (Broad-leaved 2,3

peppermint)

E. (M.) fastigata (Brown barrel) 1,2,3,5 Collected this study

E. (M.) globoidea (White stringybark) 1,3

E. (C.) gummifera (Red bloodwood) 1,2

E. (M.) laevopinea (Silvertop FCNI + Collected this study

stringybark) (*)

E. (M.) macrorrhyncha (Red 1,2,3

stringybark)

E. (M.) muelleriana (Yellow 1,3

stringybark)

E. (M.) obliqua (Messmate 1923335 FCNI + Collected this study

stringybark)

E. (M.) pilularis (Blackbutt) 1,3,5 FCNI + Collected this study

E. (M.) radiata (Narrow-leaved 1,2

peppermint)

E. (S.) resinifera (Red mahogany) FCNI (*)

E. (M.) sieberi (Silvertop ash) 1,2,3,4 FCNI + Collected this study

' Subgeneric classification used in Pryor and Johnson (1971) - M = Monocalyptus; S =

Symphyomyrtus; C = Corymbia.

? Source of common names from Brooker and Kleinig (1990).

? | = Harris et al. (1973, 1976); 2 = Wright and Harris (1974); 3 = Campbell (1969);

4 = Schedl (1968); 5 = Browne (1971).

(*) = new record.

4 Australian Entomologist, 2008, 35 (1)

Distribution and locality records

The distribution of the known host plants of Austroplatypus incompertus is

considerably more extensive than the areas where the beetle has been

recorded. Reliable locality records of A. incompertus are restricted to areas of

eucalypt forest in the coastal areas and tablelands of New South Wales and

eastern Victoria as shown in Figure 1. Much of the information shown in this

figure is derived from Harris et al. (1976); however, in examining their data

two anomalies were noted. Firstly, the reference to a locality near

Tumbarumba (see Fig. 1 in Harris et al. 1976) is not supported in their list of

known habitats of A. incompertus (see their Table 2) and represents an

anomaly. Accordingly, this location is not included in Figure 1 below.

Secondly, their reference to a ‘Glenboy State Forest’ (Harris et al. 1976) is

probably a reference to Glenbog State Forest north of Bombala, as there is no

known record of a *Glenboy State Forest".

9 & E Range of known hosts

ran

Fig 1. Recorded distribution of Austroplatypus incompertus and its known hosts.

Australian Entomologist, 2008, 35 (1) 5

Just as the current study increased the list of known host species, so it also

extended the known range of A. incompertus. Future work might well extend

this even further, especially considering that the known distribution of the

beetle is only a subset of the known distribution of the current suite of host

plants.

Acknowledgements

I thank Roger Beaver and Gregory Gowing for critically reviewing an earlier

draft of this paper.

References

BEAVER, R.A. 1979. Host specificity of temperate and tropical animals. Nature 281: 139-141.

BEAVER, R.A. 1989. Insect-fungus relationships in the bark and ambrosia beetles. Pp 121-143,

in: Wilding, N., Collins, N.M., Hammond, P.M. and Webber, J.F. (eds), /msect-fungus

interactions. Royal Entomological Society of London Symposium, Academic Press, London; 344

pp.

BEESON, C.F.C. 1941 (reprinted 1961). The ecology and control of the forest insects of India

and the neighbouring countries. Dehra Dun, India; 253 pp.

BROOKER, M.I.H. and KLEINIG, D.A. 1990. Field guide to eucalypts. Vol. 1. South-eastern

Australia. Inkata Press Pty Ltd, Melbourne; 299 pp.

BROWNE, F.G. 1961. The biology of Malayan Scolytidae and Platypodidae. Malayan Forest

Records 22: 1-255.

BROWNE, F.G. 1971. Austroplatypus, a new genus of the Platypodidae (Coleoptera) infesting

living Eucalyptus trees in Australia. Commonwealth Forestry Review 50: 49-50.

CAMPBELL, K.G. 1969. The horizontal borer. Circular of the Entomological Society of

Australia (N.S.W.) 193: 9-11.

CHIPPENDALE, G.M. 1976. Eucalyptus nomenclature. Australian Forest Research 7: 69-107.

HARRIS, J.A., CAMPBELL, K.G. and WRIGHT, G.McK. 1973. Ecological studies on the

horizontal or black borer Austroplatypus (= Platypus) incompertus (Schedl) (Coleoptera:

Platypodidae) in forests of south-eastern Australia. Unpublished report. Forests Commission

Victoria Research Branch Report No. 31: 1-22.

HARRIS, J.A., CAMPBELL, K.G. and WRIGHT, G.McK. 1976. Ecological studies on the

horizontal borer Austroplatypus incompertus (Schedl) (Coleoptera: Platypodidae). Journal of the

Entomological Society of Australia (N.S.W.) 9: 11-21.

KENT, D.S. and SIMPSON, J.A. 1992. Eusociality in the beetle Austroplatypus incompertus

(Coleoptera: Curculionidae). Naturwissenschaften 79: 86-87.

PRYOR, L.D and JOHNSON, L.A.S. 1971. A classification of the eucalypts. Australian National

University, Canberra; 102 pp.

ROBERTS, H. 1960. Trachyostus ghanaensis Schedl, (Col., Platypodidae) an ambrosia beetle

attacking Wawa, Triplochiton scleroxylon K. Schum. Technical Bulletin of the West African

Timber Borer Research Unit No. 3: 1-17.

SCHEDL, K.E. 1968. New platypodid from Australia. Memoirs of the Natural History Museum

of Victoria 28: 15-16.

6 Australian Entomologist, 2008, 35 (1)

WOOD, S.L. 1993. Revision of the genera of Platypodidae (Coleoptera). Great Basin Naturalist

53: 259-281.

WRIGHT, G.McK. and HARRIS, J.A. 1974. Ambrosia beetle in Victoria. Forestry Technical

Papers 21: 47-57. Forests Commission, Victoria.

Australian Entomologist, 2008, 35 (1): 7-17 7

FLOWERING MORPHOLOGY, PHENOLOGY AND FLOWER

VISITORS OF THE AUSTRALIAN RAINFOREST TREE RYPAROSA

KURRANGII (ACHARIACEAE)

BRUCE L. WEBBER!, ALAN S.O. CURTIS’, GERASIMOS CASSIS? and

IAN E. WOODROW’

! Centre d'Ecologie Fonctionnelle et Evolutive, CNRS (UMR 5175), 1919 Route de Mende,

34293 Montpellier Cedex 05, France (email: b.webber@bigfoot.com)

?Daintree Discovery Centre, Cow Bay, Qld 4873, Australia

"School of Biological, Earth and Environmental Sciences, University of New South Wales,

Sydney, NSW 2052, Australia

*School of Botany, The University of Melbourne, Vic 3010, Australia

Abstract

The understorey tree Ryparosa kurrangii B.L.Webber (Achariaceae) is restricted to a limited

number of populations in three distinct valleys of lowland tropical rainforest in northern

Queensland, Australia. To provide baseline information for determining potential pollen vectors

and the opportunity for inter-population gene flow, flowering morphology, phenology and flower

visitation were studied in natural populations over a number of seasons. Contrary to all previous

work, R. kurrangii was found to be monoecious, with temporally separated flushes of staminate

and carpellate flowers. Flower production was primarily basicauliflorous on the main trunk with

small open flowers borne on long racemes. Heavy nocturnal scented nectar production coincided

with peak animal visitation. Predominant flower visitors were adults of an undescribed species of

Monolepta Chevrolat sensu lato (Coleoptera: Chrysomelidae: Galerucinae) and nymphs of

Coridzolon australiense Carvalho & Gross (Heteroptera: Miridae: Mirinae: Hyalopeplini). Now

that the floral biology and morphology as well as the dominant flower visitors of R. kurrangii

have been documented, further work is required to determine the likely contribution of flower

visitors to pollination events and population gene flow.

Introduction

The wet tropical rainforests of northern Queensland comprise just 0.1% of

the land surface yet contain 2596 of all plant genera found in Australia (Keto

and Scott 1986). In tropical rainforests, plant-pollinator and plant-frugivore

interactions are vital for ensuring genetic variation in plant populations

through outcrossing (e.g. Janzen 1983, Bodmer 1991, Mack 1993, Corlett

2001, Brewer and Webb 2002, Peres and van Roosmalen 2002).

Ryparosa kurrangii B.L. Webber (Achariaceae; Flacourtiaceae pro parte) is a

rare sub-canopy tree restricted to tropical lowland rainforest of the Daintree

region. The taxon is likely to depend closely on the endangered cassowary

(Casuarius casuarius johnsonii; a large flightless bird) for long distance seed

dispersal (Webber and Woodrow 2004). However, continued development in

the Daintree lowlands is encroaching on current R. kurrangii populations and

the resulting disturbance appears to be having a detrimental effect on what

might be a significant interaction for R. kurrangii with the increasingly

endangered cassowary (B.L. Webber, pers. obs.) Therefore, any inter-

population gene flow that results from pollination events will become

increasingly important for maintaining genetic diversity, if the disruption of

plant-frugivore interactions continues.

8 Australian Entomologist, 2008, 35 (1)

Previous taxonomic literature has highlighted how little is known about the

life history of flowering in flacourtiaceous genera (van Slooten 1925,

Sleumer 1954). Conflicting statements over monoecy or dioecy within

species (e.g. Woodson and Schery 1968) may be attributed to the

predominance of species descriptions relying largely on herbarium specimens

over field-based observations. Even less is known about pollination vectors in

the family. Thus, the aims of this current study were (1), to provide a detailed

documentation of flowering morphology and phenology in Ryparosa

kurrangii and (2), to determine flower visitors that might act as pollination

vectors and provide opportunity for gene flow during flower fertilisation.

Methods

Field observations on the life history and developmental biology of Ryparosa

kurrangii were made over six years (1998 to 2004), in the Daintree World

Heritage area of northern Queensland, Australia (16°08’S, 145?26'E). The

species is currently classified as rare by the Queensland Nature Conservation

Act (Queensland Government 1992), although a higher conservation status

might be required in light of recent work (Webber and Woodrow 2004, 2006,

Webber 2005). Ryparosa kurrangii has a very restricted distribution and is

only known from three distinct valleys comprising a small strip of coastal

lowland tropical rainforest between the Daintree River and Cape Tribulation.

Within these valleys, R. kurrangii is spatially rare but locally common, often

forming discrete populations near creeks and ephemeral streams. Rainforest

vegetation communities at all sites used in the study can be classed as

complex mesophyll vine forest (CMVF, Type la) or mesophyll vine forest

(MVF, Type 2a; sensu Tracey 1982).

Observations on reproductive biology, life history patterns and plant-animal

interactions were made on eight tagged R. kurrangii populations across the

three valleys, each comprising between 30 and 350 individuals. This allowed

for the long-term monitoring of numerous individual trees (Webber 2005).

More remote untagged populations were periodically examined within the

wider Daintree lowlands region. Flowering phenology and morphology were

documented on an ad hoc basis throughout flowering periods that occurred

during the 6 years of study. Phenological variation in flowering patterns (the

timing of distinct stages of floral development and flower anthesis within and

between populations) was recorded at random intervals spaced throughout the

flowering season. Nectar production was monitored during day visits to

populations and also on night visits while observing flower visitation. Floral

morphological data were collected from fresh flower racemes in the field and

laboratory.

Flower visitation observations for male and female racemes were made both

at night (between dusk to 00:30 and 04:00 to dawn) and during daylight

hours. Visitation was recorded by observing individual trees for periods of at

least 10 minutes, supplemented by random spot sweeps for the presence of

Australian Entomologist, 2008, 35 (1) 9

flower visitors on an individual tree at any one time. These observations were

quantified, where possible, based on the number of individuals, their

interactions with flower racemes (position on flowers, any characteristic

behaviour or foraging activity) and subsequent movements between

individual trees. Invertebrates collected from R. kurrangii flower racemes

were identified from field-collected specimens either preserved in 7096

ethanol or dry killed and mounted.

Voucher specimens for collected invertebrates were lodged at the Australian

Museum, Sydney and the Australian National Insect Collection (ANIC),

Canberra.

Results

Flower morphology and phenology

All eight permanently tagged populations of R. kurrangii were comprised of

trees that were consistently monoecious. On individual trees, staminate and

carpellate flower racemes were produced in distinctly separate temporal

periods on all individuals observed. Staminate flowers appeared first and

were temporally separated from mature carpellate flowers (which appeared as

buds while staminate flowers were open) on the same individual by 3-4

weeks. Occasionally, individual trees produced two flushes of staminate

flowers before a carpellate flush (within the same season), while other

individual trees consistently produced heavier flushes of either staminate or

carpellate flowers over a number of flowering seasons. Within populations

and between populations in close proximity to each other (i.e. in the same

valley) there was temporal overlap between staminate and carpellate racemes

on different individual trees. However, flowering synchrony between

populations in different valleys, although still temporally overlapping, was

not as tightly synchronised.

Flowering position was, for the most part, on the lower portion (2-3 m and

sometimes up to 5 m) of the main trunk from distinctive tubercles (Fig. 1).

On large individuals, these tubercles exceeded 150 x 150 mm and racemes

were produced in fascicles of up to 5 from multiple points on each tubercle

(Fig. 2). While staminate flowers (and very rarely carpellate flowers) were

occasionally observed on small tubercles situated on side branches, flowering

on side branches generally only occurred on younger trees with poorly

developed trunk tubercles. Mature fruits were almost always observed on the

main trunk. Flowering or fruiting was very rarely observed on trees smaller

than 5 m in height and with a diameter at breast height (DBH) of less than

approximately 50-60 mm. Raceme peduncles underwent significant

elongation shortly before the flowers opened and, after flower anthesis,

racemes with staminate flowers were generally 300-400 mm long; those with

carpellate flowers were slightly shorter. On rare occasions, staminate and

carpellate flower racemes in excess of 600 mm long were observed.

10 Australian Entomologist, 2008, 35 (1)

Figs 1-2. Floral raceme position on Ryparosa kurrangii. (1) flower racemes are

generally borne on large distinctive tubercles on the lower portion of the main trunk;

(2) racemes form in fascicles of up to five.

Flowers generally flushed in a spectacular display covering the trunk during

the dry season of June to September (Fig. 3). Staminate flower buds were

globose (Fig. 4), while carpellate buds were pyriform (Fig. 5). The calyx on

both staminate and carpellate flowers was observed to irregularly rupture

during floral anthesis (Fig. 6), to reveal carpellate flowers with short fleshy

staminodes (Figs 7, 8) and staminate flowers with united filaments (Fig. 9).

All recently opened flowers were heavy with nectar at night (Fig. 10). The

fleshy petal scale, a characteristic feature of the genus and other closely

related Pangieae, provided a collection point for nectar droplets during the

night (which then pooled in the keeled petals to a lesser extent during the

day; Fig. 11). The flower scent was strong, particularly at night, and could be

described as somewhat sickly sweet with overtones of wet dog and body

odour. Extrorse dehiscence of the anthers on staminate flowers was observed

to deposit a significant number of pollen grains on the fine hairs surrounding

the anthers, as well as on the filaments (Fig. 12). Mutated carpellate flowers

were occasionally found with deformed anthers borne on some staminodes

(Fig. 13).

Figs 3-13. Floral morphology and habit of Ryparosa kurrangii. (3) flowering takes

place in a spectacular flush covering the trunk during June-September; (4) globose

buds on staminate flower racemes; (5) pyriform buds on carpellate flower racemes;

(6) the calyx (staminate flower shown) irregularly ruptures during floral anthesis; (7)

carpellate flowers with a superior ovary; (8) short fleshy staminodes (indicated by an

arrow) on carpellate flowers; (9) staminate flowers with united filaments; (10) open

flowers (staminate flower shown) are heavy with nectar droplets at night; (11) the

fleshy petal scale provides a common collection point for nectar droplets; (12) pollen

grains (indicated by an arrow) frequently lodge in hairs surrounding the anthers and

on the filaments in staminate flowers; (13) mutated staminode with anther (indicated

by arrow).

Australian Entomologist, 2008, 35 (1) 11

12 Australian Entomologist, 2008, 35 (1)

Flower visitors

All animal interactions observed involved small invertebrates and no flower

visitation by vertebrates such as bats was witnessed. Both staminate and

carpellate flowers had no obvious morphological adaptations to visitation by

specialised insects; however, their small size and relative fragility may

preclude larger animals from settling. Most flower visitation took place at

night and invertebrates were generally concentrated on flowers with ample

nectar. Consistent flower visitation across flowering seasons was observed

for a range of invertebrate species. These included numerous small thrips

(Thysanoptera) concentrated on and around the petal scales and ant species

(Hymenoptera: Formicidae) feeding on flower nectar, including the weaver

or green tree ant Oecophylla smaragdina (Fabricus). However, the two most

abundant invertebrates on R. kurrangii flowers were an undescribed species

of Monolepta Chevrolat sensu lato (Coleoptera: Chrysomelidae: Galerucinae;

Figs 14, 15) and the mirid Coridzolon australiense Carvalho & Gross

(Heteroptera: Miridae: Mirinae: Hyalopeplini; Fig. 16).

Both Monolepta sp. and C. australiense were observed on staminate and

carpellate flowers in all R. kurrangii populations monitored across numerous

flowering seasons. Individuals spent periods of up to 5 minutes on each

flower raceme, coming into contact with both anthers and stigmas (on

staminate and carpellate flowers, respectively). These insects appeared to be

feeding on nectar and perhaps pollen, doing little visible chewing damage to

the actual floral organs during flower visitation. The majority of C.

australiense observed were nymphs, which is a strong indication that R.

kurrangii may be a breeding host plant. Surveys of C. australiense abundance

revealed up to 12 individuals spread over 20 racemes on each tree. In

contrast, all observed specimens of Monolepta sp. were adults that were often

seen flying between staminate and carpellate racemes on nearby trees when

disturbed. The density of Monolepta sp. was also far greater than any other

flower-visiting invertebrate on R. kurrangii, with conservative estimates of

over 300 individuals per tree on some nights.

Discussion

As a genus, Ryparosa is little studied and poorly known from field-based

interactions. It has rarely appeared in the wider scientific literature since it

was first described nearly 200 years ago. Current taxon delimitation within

the genus is largely based on vegetative qualities derived from herbarium

specimens and many Ryparosa species are still imperfectly described in

terms of staminate or carpellate flower form (e.g. Ridley 1936, Sleumer 1954,

Jarvie and Stevens 1998). An overall unfamiliarity with the genus,

particularly in the field, might have led to the situation where the taxon is

widely documented as being dioecious (e.g. Sleumer 1954, Hutchinson 1967,

Renner and Ricklefs 1995, Jarvie and Stevens 1998).

Australian Entomologist, 2008, 35 (1) 13

Figs 14-16. Common invertebrate flower visitors of Ryparosa kurrangii. (14-15) adult

of an undescribed species of Monolepta; (16) nymphs of Coridzolon australiense.

By observing tagged populations of R. kurrangii in Australian rainforests, it

was possible to follow the reproductive biology of individual trees over a

number of seasons. This revealed that R. kurrangii appears to be consistently

monoecious. A distinct confusion over the reproductive details of the

Flacourtiaceae sensu lato in general was described by van Slooten (1925),

who cited examples of Pangium edule Reinw. ‘bearing male flowers and later

on a few fruits’ while trying to describe the species as being dioecious.

Similar observations were made by Woodson and Schery (1968) in other

flacourtiaceous genera. Given that staminate and carpellate flowering periods

in R. kurrangii were temporally distinct, it would be easy to miss this trait in

a once-off sampling of field material. However, Ryparosa species have been

cultivated in the Bogor Botanic Gardens in Indonesia for well over a century,

without any conflicting reports of monoecy (Koorders and Valeton 1900).

Reports of both monoecy and dioecy in Hydnocarpus and other closely

related genera warrant further investigation (e.g. van Slooten 1925).

14 Australian Entomologist, 2008, 35 (1)

For R. kurrangii, nocturnal nectar production and a strong scent, combined

with mass flowering, small open flowers with hairs to trap pollen and a lack

of overall colour, fits closely with a beetle pollination (cantharophily)

syndrome (e.g. Faegi and van der Pijl 1976, Irvine and Armstrong 1988,

1990, Williams and Adam 1994). While the fragile structure of the flower

racemes and the very small size of the flowers would most likely preclude

vertebrates (such as bats) as potential pollinators, they should not be ruled out

altogether (e.g. Hopper 1980, Crome and Irvine 1986).

Flowers under observation were visited by a range of invertebrates dominated

by Monolepta sp., C. australiense and thrips (Thysanoptera). Thrips have

been shown to be effective pollinators for other plant species (Williams et al.

2001, Moog et al. 2002). However, the lack of wind in the rainforest

understorey combined with a weak flying ability means that their role as an

efficient pollination vector for R. kurrangii is doubtful. The problem of

dubious inter-plant visitation also applies to the nymphs of C. australiense,

which are highly unlikely to be a dispersive life stage of the species. It is

generally thought that males of the Miridae are the dispersive phase;

however, no observations were made on the movements of adults. Mirids

have been implicated as potential pollinators, primarily because of their

known pollen feeding strategy and common association with flowers. The

fact that many mirid species are host plant specific increases their capacity as

pollinators, but most studies to date indicate that they are inefficient (Bohart

and Nye 1960) or chance (Wheeler 2001) pollinators. Given that there are

increasing linkages between mirids and flower pollination in other tree

species (Ashton et al. 1988, Corlett 2004), their role as a pollination vector

warrants further investigation.

Perhaps the most effective pollination vector in a seemingly generalist

approach for R. kurrangii may be Monolepta sp. Beetles have increasingly

been considered as potential pollinators (Young 1986, Irvine and Armstrong

1990), with documentation of specialised interactions. Williams and Adam

(1998, 2001) have shown that smooth-bodied invertebrates are inadvertently

capable of transporting pollen. The Galerucinae are known to be

polyphagous, feeding on pollen and nectar as well as flowers and leaves (e.g.

Murray 1982) and observed interactions with R. kurrangii would allow for

pollen transfer between staminate and carpellate flowers. ANIC specimen

records indicate a known distribution range for this Monolepta sp. from

Palmerston to Cape Tribulation in northern Queensland (C.A.M. Reid, pers.

comm.). Therefore, with a distribution much wider than that of R. kurrangii,

it is unlikely that the observed Monolepta taxon is host specific. Studies on

other trees, such as A/phitonia excelsa (Fenzl) Reiss. ex Benth. (Rhamnaceae)

and Syzygium cormiflorum (F. Muell.) B. Hyland (Myrtaceae), also list a

number of Monolepta taxa as flower visitors (Crome and Irvine 1986,

Williams and Adam 2001).

Australian Entomologist, 2008, 35 (1) 15

This study has addressed two areas where a lack of knowledge and a reliance

on herbarium specimens over field observations in plant systematics are

clearly impeding our understanding of plant reproductive biology and plant-

pollinator interactions. Firstly, temporally separated monoecious flowering

may be more widespread in Ryparosa and other closely related taxa than is

currently thought. Further field studies of tagged trees will be required to

confirm if this trait is ubiquitous in R. kurrangii and, perhaps, characteristic

of a wider range of closely related genera. Secondly, the documentation of

what might be an invertebrate pollination system adds to the limited number

of pollination studies from Australian tropical rainforests. Importantly, more

work now needs to be done in extending our knowledge of the pollination

efficiency and dispersal distance of R. kurrangii flower visitors, given their

potential impact on gene flow within and between populations of this rare

tree.

Acknowledgements

Field sampling was conducted under Scientific Purposes permits issued by

the Queensland Department of Environment and Heritage (F1/000233/00/

SAA, WISP01144103, WITK01149403, WISP01889604, WITK01889704)

and with the full permission and support of private landholders when

conducted on freehold land. We acknowledge Chris Reid (Australian

Museum, Sydney) for his assistance in specimen identification, Kate Bugg

and Adrian Hall for their assistance in the field, and Geoff Williams

(Australian Museum, Sydney) for an insight into Australian rainforest

pollination vectors. Comments from Sarah Boulter and Helen Wallace

improved the manuscript. BLW was a recipient of an Australian Postgraduate

Award PhD Scholarship.

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CORLETT, R.T. 2004. Flower visitors and pollination in the Oriental (Indomalayan) Region.

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Pp 135-149, in: Bawa, K.S. and Hadley, M. (eds), Reproductive ecology of tropical forest plants.

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Government Printing Service, Canberra.

KOORDERS, S.H. and VALETON, T. 1900. Bijdrage No. 6 tot de Kennis der Boomsoorten op

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18 Australian Entomologist, 2008, 35 (1)

A NOTE ON BUTTERFLY (LEPIDOPTERA) RECORDS FROM THE

DARWIN REGION, NORTHERN TERRITORY

DANIEL KING

420 Spencer Road, Thornlie, WA 6108

(email: drd_king@hotmail.com or ventnor!@dodo.com.au)

Abstract

Additional records are provided for Eurema herla (W.S. Macleay) and Hypocysta adiante

antirius Butler from Darwin, Northern Territory.

Introduction

Meyer et al. (2006) provided records for 87 butterfly species collected from

the Darwin region between 1990 and 2005 and for an additional 24 species

recorded historically. Further notes on two of these species are provided

below.

New records

PIERIDAE

Eurema herla (W.S. Macleay)

Meyer et al. (2006) noted for this species that ‘Further data are required to

confirm its existence in the Darwin region.’ On 16 November 2006, |

collected two specimens, in freshly emerged condition, at Lee Point [locality

21 on the map of Meyer et al. (2006)]. Many other specimens were present,

suggesting the existence of a colony.

NYMPHALIDAE

Hypocysta adiante antirius Butler

Two specimens, of freshly emerged appearance, were collected at the same

location (Lee Point) and on the same day as those of E. herla noted above.

This is noteworthy since Meyer et al. (2006) did not record this species at this

location or flying during November.

Reference

MEYER, C.E., WEIR, R.P. and WILSON, D.N. 2006. Butterfly (Lepidoptera) records from the

Darwin region, Northern Territory. Australian Entomologist 33(1): 9-22.

Australian Entomologist, 2008, 35 (1): 19-20 19

ADDITIONAL RECORDS OF HAWK MOTHS AND BUTTERFLIES

(LEPIDOPTERA) FROM LIZARD ISLAND, NORTHERN

QUEENSLAND

R.B. LACHLAN

Entomology Department, Australian Museum, 6 College St, Sydney, NSW 2010

Abstract

Additional records are provided for two species of hawk moths and six species of butterflies

from Lizard Island, northern Queensland. Notes on abundance are included.

Introduction

Prior to 2004, only two species of hawk moths and eleven species of

butterflies had been recorded from Lizard Island (Moulds 1985, Duckworth

and McLean 1986). Since then, Lachlan (2004, 2006) has recorded a further

19 species of hawk moths and 21 species of butterflies.

From 2-13 April 2007, the author carried out a third survey of Lizard Island.

It followed a better than average wet season but, once again, continuous

strong winds restricted the survey. This was particularly so in the more

exposed areas of the island. Thus, collecting was primarily confined to the

forested areas around the Australian Museum Research Station.

Voucher specimens are temporarily in the author's collection; duplicates are

held by the Queensland Museum, Brisbane and the Australian Museum,

Sydney.

Discussion

Table 1 details the newly recorded species. Nearly all the butterfly species

previously recorded were netted, identified and released at various times

during the latest survey. Two notable exceptions were Petrelaea tombugensis

(Róber) and Belenois java teutonia (Fabricius), neither of which was seen.

Other lycaenid species recorded for the first time during the 2005 survey,

including Nacaduba berenice berenice (Herrich-Scháffer), Jamides phaseli

(Matthew) and Theclinesthes miskini eucalypti Sibatani & Grund, were much

more common during this latest survey. It is interesting to note that all six

newly recorded species are common species and it is surprising that most, if

not all, were not encountered previously.

Once again, the overall abundance of hawk moths, in terms of both the

number of species and the actual number of individual specimens coming to

light traps, was well down compared with the first survey carried out in early

December 2002 (Lachlan 2004).

Twenty-three species of hawk moths (approximately one third of the total

Australian mainland species) and 38 butterfly species have now been

recorded from Lizard Island.

20 Australian Entomologist, 2008, 35 (1)

Table 1. List of hawk moths and butterflies collected or sighted on Lizard Island

during the April 2007 survey, additional to those recorded by Lachlan (2004, 2006).

All are new species records for the island.

Species Notes

HAWK MOTHS

Sphingidae

Daphnis placida placida (Walker) One male, two females

Acosmeryx anceus anceus (Stoll) Four males

BUTTERFLIES

Papilionidae

Graphium agamemnon ligatum (Rothschild) ^ Not common

Nymphalidae

Cupha prosope prosope (Fabricius) Three clearly sighted; none taken

Hypolimnas alimena lamina Fruhstorfer Three males

Euploea darchia niveata (Butler) One male; two others sighted

Lycaenidae

Zizeeria karsandra (Moore) One male, one female. Not common

Zizina labradus labradus (Godart) One male, two females. Uncommon

Acknowledgements

Once again this survey was supported by the Australian Museum by

provision of facilities at their Lizard Island Research Station. I sincerely

thank the Directors, Dr Anne Hoggett and Dr Lyle Vail, for providing access

to the Research Station for my family and I during the survey period. The

assistance of staff members Tania and Bob Lamb was greatly appreciated. I

also thank the Park Ranger, Alan Clackson (Queensland Parks and Wildlife

Service), for his continued support of this survey, carried out under Permit

number WITK03173805. For comments on the manuscript I sincerely thank

Dr Max Moulds (Australian Museum, Sydney).

References

DUCKWORTH, B.G. and McLEAN, J. 1986. Notes on a collection of butterflies from the

islands of the Great Barrier Reef, Queensland. Australian Entomological Magazine 13(3-4): 43-

48.

LACHLAN, R.B. 2004. An annotated list of the hawk moths and butterflies (Lepidoptera) of

Lizard Island, Queensland. Australian Entomologist 31(1): 1-3.

LACHLAN, R.B. 2006. New records of hawk moths and butterflies (Lepidoptera) from Lizard

Island, northern Queensland. Australian Entomologist 33(3): 133-135.

MOULDS, M.S. 1985. A review of the Australian hawk moths of the genus Macroglossum

Scopoli (Lepidoptera: Sphingidae). Australian Entomological Magazine 12(5): 81-105.

Australian Entomologist, 2008, 35 (1): 21-26 21

THE EXOTIC BEE HALICTUS SMARAGDULUS V ACHAL, 1895

(HYMENOPTERA: HALICTIDAE) IN THE HUNTER VALLEY,

NEW SOUTH WALES: A NEW GENUS IN AUSTRALIA

JOHN R. GOLLAN!”, MICHAEL BATLEY ? and CHRIS A.M. REID!

! Australian Museum, 6 College Street, Sydney, NSW 2010

(Email: john.gollan@austmus.gov.au)

"School of Environmental Sciences and Natural Resources Management, University of New

England, Armidale, NSW 2351

"Department of Biological Sciences, Macquarie University, NSW 2109

Abstract

The emerald furrow bee, Halictus smaragdulus Vachal, 1895, native to the western Palaearctic,

is recorded from the Upper Hunter Valley region, New South Wales, Australia. Halictus

Latreille is a genus previously unknown in Australia. The bee is well established in the area and

has been observed visiting flowers of exotic plants, including Galenia pubescens - a declared

noxious weed.

Introduction

Several specimens of an unfamiliar bee were collected as part of research by

JRG investigating the response of terrestrial invertebrates to riparian habitat

rehabilitation in the Upper Hunter region of New South Wales (Fig. 1 inset).

These small bees were similar in size and colour to Lipotriches flavoviridis

(Cockerell, 19052), but with the divided prepygidial fimbria characteristic of

the subfamily Halictinae (Fig. 2). They were identified as a species of

Halictus Latreille, a genus previously unknown in Australia.

The bees were further identified as belonging to Halictus subgenus Seladonia

Robertson by Dr Ken Walker (Museum Victoria) and Dr Laurence Packer

(York University, Canada) and named as Halictus (Seladonia) smaragdulus

Vachal, 1895 by Fr Andreas Ebmer (Puchenau, Austria), an expert on

Seladonia taxonomy. The species has since been added to the Australian

Faunal Directory (Walker 2006a) and Australia’s Pest and Diseases Image

Library [PaDIL] (Walker 2006b). Diagnostic images can be found on the

PaDIL website (Walker 2006b).

Identification, distribution and biology

Halictus smaragdulus is a metallic green bee, 6-8 mm long. It is native to the

western Palaearctic, from Spain and Portugal in the west, through southern

and central Europe, to Kyrgyzstan and Afghanistan in the east (Dawut and

Tadauchi 2002). In Germany it is known as the Smaragdgrtine Furchenbiene,

which translates as the emerald furrow bee, an appropriate vernacular name

for use in Australia. Although there are few published flower-visiting

records, H. smaragdulus is probably polylectic and has been recorded visiting

Asparagus aphyllus and Thymus capitatus in Europe (Herrera 1988,

Petanidou and Vokou 1993).

22 Australian Entomologist, 2008, 35 (1)

HUNTER RIVER

Fig. 2. Female Halictus (Seladonia) smaragdulus Vachal from the Upper Hunter

Region, NSW, Australia. Scale bar = 2 mm.

Australian Entomologist, 2008, 35 (1) 23

Most of the 360 species of Australian halictids belong to the genus

Lasioglossum Curtis (251 species) or its close relative Homalictus Cockerell

(39 species). Halictus smaragdulus is identifiable as belonging to the

subfamily Halictinae by its wing venation and divided prepygidial fimbria, as

described by Michener (2000). Females may be distinguished from other

Australian halictines by the presence of apical bands of dense setae on the

metasomal tergites and the strongly sclerotised third submarginal crossvein of

the forewing. Males of H. smaragdulus can also be identified by the well

developed lower gonostylus, illustrated by Ebmer (1998) and Dawut and

Tadauchi (2002). It has been suggested the subgenus Seladonia should be

raised to generic rank (Pesenko 2000), but this does not appear to have

received universal acceptance. Dawut and Tadauchi (2002) provided a

detailed redescription of the species.

The family Halictidae includes roughly 20% of all bee species known in

Australia, as estimated from the Australian Faunal Directory (Walker 20062).

Most species nest in the ground (Michener 2000). About 25% of all species,

but probably none of the Australian species (Walker 1986), form eusocial

colonies, while others exhibit varying degrees of parasocial behaviour

(Michener 1990). Fourteen species of subgenus Seladonia are known to be

eusocial (Pesenko 2000). Halictus smaragdulus is probably similar to other

H. (Seladonia) species described by Yanega (1997), with a caste system of

queens and workers. Members of different castes may be indistinguishable

morphologically and an individual female may change her caste during her

lifetime (Michener 1990).

Occurrence in Australia

Invertebrates were surveyed by JRG in the Upper Hunter region, as part of a

PhD investigating responses of invertebrates to riparian habitat rehabilitation.

Yellow pan trapping was undertaken at 12 locations in November 2004 and

January 2006. Four types of habitat were sampled: open grassland; young

revegetation with native saplings planted in the last 4 years; old revegetation

with native saplings planted 10-15 years ago; and natural mature woodland.

Study locations were not regularly spaced but were approximately 5 km apart

along the river (Fig. 1).

Eight yellow pan traps (plastic bowls 17 cm in diameter and 5 cm deep,

Deeko™) were placed approximately 10 m apart at each site. All traps were

half filled with a dilute solution of brine and 2-3 drops of detergent, secured

on the ground using wooden food skewers and collected after 48 hours.

Six hundred and thirty nine bees, representing 17 species, were collected, of

which Halictus smaragdulus comprised 22% of the total abundance and was

the second most abundant species collected. Other abundant species trapped

were Homalictus sphecodoides (Smith, 1853) [32%] and H. caloundrensis

(Cockerell, 1914) [21%].

24 Australian Entomologist, 2008, 35 (1)

Halictus smaragdulus was trapped at seven of the 12 sampling sites (Fig. 1),

with two sites yielding the majority of specimens. One of these sites was an

open grassland habitat, located just northeast of Aberdeen on Fig. 1 (32?08'S

151?55'E), which was dominated by exotic pasture grasses and herbs such as

Galenia pubescens and Foeniculum vulgare. The second site where

abundance of H. smaragdulus was high was an old revegetation site at

Muswellbrook (32?15'S 151°53'E), which had been planted with native trees

typical of riparian zones in the region. Species planted included Eucalyptus

camaldulensis, E. tereticornis, E. melliodora, Casuarina glauca and C.

cunninghamiana subsp. cunninghamiana. The understorey was dominated by

exotic pasture grasses and herbs like those of the other open grassland sites.

The remaining five sites where H. smaragdulus was trapped produced 19

individuals.

Following discovery of the species, MB visited the Upper Hunter region on

two occasions, in 2005 and 2007. At Denman in April 2005, H. smaragdulus

of both sexes were observed visiting roadside flowers of the introduced plant

Galenia pubescens. MB estimated that the density of bees was greater than

10 per m? over an area of at least 30 m^. A similar density of the native

species Ceylalictus perditellus (Cockerell, 1905b) was present. In January

2007, aggregations of H. smaragdulus were found on the exotic flower

Convolvulus mauretanicus at Singleton (32°35'S 151°11'E) and on the exotic

Verbena bonariensis at Glenbawn Dam (32°06'S 150°59'E). Nests were not

found at any of the locations but were probably within 100 m of the observed

bees (Greenleaf et al. 2007).

Discussion

Halictus smaragdulus appears well established in the Upper Hunter region of

New South Wales. In the survey using yellow pan traps, H. smaragdulus was

the second most abundant bee species trapped. Yellow pan traps may be an

efficient method of trapping this exotic bee, particularly where standardised

counts are required.

Given that H. smaragdulus is native to European regions that have a climate

much like temperate Australia, we think that this species has potential to

spread (and may already be present) over a much wider area (north, south,

east and west) than was observed in our study. Furthermore, many of

Australia's introduced weeds, many of which are European, may also provide

a suitable food source for the bees. Halictus smaragdulus was trapped and

observed in habitats that ranged from open grassland to mature woodland,

including roadside verges. Two sites yielded much higher numbers than other

sites. This may denote some preference trend or signify where the bees have

been established for a longer period. However, patterns observed in our

yellow pan survey could also be the result of natural spatial variability. More

dedicated research 1s required to determine the actual extent of the incursion

and on the spatial and temporal variation in abundance.

Australian Entomologist, 2008, 35 (1) 25

While we are unaware of any records of H. smaragdulus causing ecological

problems outside Australia, this species has potential to be very successful in

Australia. The possible undesirable effects of an explosion in the population

of H. smaragdulus would include competition with native species for limited

resources and transmission of parasites or pathogens to native organisms.

Inaction or slow reaction to the discovery of H. smaragdulus may also result

in irreversible environmental damage and species extinctions. Our opinion is

that H. smaragdulus also has great potential to spread exotic plants. Halictus

smaragdulus was observed visiting flowers of several exotic species,

including Galenia pubescens. This plant is a declared noxious weed in NSW

(under the Noxious Weeds Act 1993), requiring management plans to control

its spread in two regions immediately north of the study region (MacDonald

2006). Prioritising exotic species for management or policy actions (control,

removal or prevention) is not an easy task, but guidelines to direct research

are available (e.g. Byers et al. 2002).

Acknowledgements

We thank Dr Ken Walker, Dr Laurence Packer and Fr Andreas Ebmer for

identifications. PhD research was funded by a grant awarded to John R.

Gollan by the NSW Environmental Trust (Project Ref. # 2003/RD/G0001).

Thanks to Scott Ginn and Michael Elliott at the Australian Museum for

imaging and figures.

References

BYERS, J.E., REICHARD, S., RANDALL, J.M., PARKER, I.M., SMITH, C.S., LONSDALE,

W.M., ATKINSON, LA.E., SEASTEDT, T.R., WILLIAMSON, M., CHORNESKY, E. and

HAYES, D. 2002. Directing research to reduce the impacts of nonindigenous species.

Conservation Biology 16(3): 630-640.

COCKERELL, T.D.A. 1905a. New Australian bees of the genus Nomia. Entomologist 38: 217-

223.

COCKERELL, T.D.A. 1905b. Descriptions and records of bees. I. Annals and Magazine of

Natural History (7) 16: 216-225.

COCKERELL, T.D.A. 1914. Descriptions and records of bees. LIX. Annals and Magazine of

Natural History (8) 13: 504-522.

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Australian Entomologist, 2008, 35 (1): 27-35 27

COMPETITION FOR LARVAL FOOD PLANT BETWEEN DELIAS

ARGENTHONA (FABRICIUS) AND DELIAS NIGRINA (FABRICIUS)

(LEPIDOPTERA: PIERIDAE) IN COASTAL WALLUM HABITAT IN

SOUTHERN QUEENSLAND

A.G. ORR

26 Currimundi Road, Caloundra, Qld 4551

Abstract

In parts of their ranges the mistletoe-feeding pierines Delias argenthona (Fabricius) and D.

nigrina (Fabricius) overlap in time and space and compete for larval food plant. The range of

food plants used does not correspond exactly for the two species, which also exhibit different

phenologies in a given locality, D. nigrina preferring the cooler months of the year. In coastal

wallum forest in southern Queensland, larvae of both feed on the mistletoes Dendrophthoe

vitellina and Muellerina celastroides but Diplatia furcata is utilised only by D. argenthona and

Amyema congener only by D. nigrina. This study demonstrates that, in the presence of a

competitor, both Delias Hübner species switch their oviposition patterns, laying more of their

egg clusters on the mistletoe species not used by the competing species. It is also demonstrated

that, in experimental situations, ovipositing females of both species avoid plants carrying egg

masses of their own and the competing species and lay preferentially on plants without eggs. It is

suggested that in nature the decision to oviposit is made only after a thorough visual assessment

of the mistletoe plant and plants bearing eggs are avoided.

Introduction

Larvae of the endemic Australian pierine species Delias argenthona

(Fabricius) and D. nigrina (Fabricius) feed almost exclusively on mistletoes.

Braby (2006) recorded D. argenthona larvae in Australia as feeding on 12

species in five genera of Loranthaceae, as well as a single record of larvae on

Santalum lanceolatum (Santalaceae). For D. nigrina, Braby (2006) listed 15

species in six genera of Loranthaceae as natural food plants. Mistletoes

known to be utilised by both species include Amyema cambagei, A. miquellii,

Dendrophthoe curvata, D. glabrescens, D. vitellina and Muellerina

celastroides.

Throughout their ranges, which largely overlap in eastern Australia from

Cape York to the Victorian-NSW border (Braby 2000), habitat, altitude and

phenology frequently segregate D. argenthona and D. nigrina populations. In

southern Queensland D. argenthona is most common in the coastal lowlands,

where adults are abundant in autumn and spring. It also occurs in moderately

dry inland areas but is generally absent from upland areas and from

rainforest. Adults of Delias nigrina, by contrast, are common in upland

rainforest from late spring to late autumn, being absent from these areas in

winter. In the coastal lowlands they are most common in winter, seldom

appearing before mid autumn. These differences are at least partly

attributable to the different effects of temperature on the development and

survival of the two species, D. nigrina preferring distinctly lower

temperatures than D. argenthona (Nousala 1979, Braby and Lyonns 2003). In

tropical northern Queensland D. nigrina is exclusively a montane species,

28 Australian Entomologist, 2008, 35 (1)

flying mainly in the winter. Delias argenthona is also most commonly found

in high country, being replaced on the coast by the tropical D. mysis

(Fabricius). All recorded food plants of D. mysis are also utilised by D.

argenthona (Braby 2000).

At Currimundi Lake, Caloundra, southern Queensland, there are extensive

stands of low mixed Melaleuca and Banksia forest growing on white sand

adjacent to urban development. This is a climax stage of the vegetation

complex known as ‘wallum’ (Coaldrake 1961). Four mistletoe species occur

in the area: Amyema congener, Dendrophthoe vitellina, Diplatia furcata and

Muellerina celastroides, all at moderate to high abundance and infesting

mainly the dominant tree species. In this habitat, at different times of the

year, D. argenthona, D. nigrina or both are usually exceedingly common and

together utilise all four mistletoe species as larval food plants. However,

Diplatia furcata is utilised only by D. argenthona and Amyema congener

only by D. nigrina.

In 1974, I first noticed that Delias herbivory often defoliated clumps of

mistletoe in nature, with larval cohorts, sometimes including both species of

Delias, dying of starvation. Moreover, it was clear that, although there was an

apparent hyper abundance of mistletoe, much of the growth was very old and

sclerotised and larval survival in captivity on such fodder was invariably

poor. Eggs were almost always laid on the freshest growth available. This

suggested that, although there was an apparent abundance of food plant, the

real amount of high quality food available was often limiting, providing the

conditions under which both intra- and interspecific competition could occur.

The central question was, therefore, what might be the effect of interspecific

competition on food plant utilisation? To answer this, biweekly censuses

were made from February to September 1975 of all Delias early stages on

mistletoes growing along a standard path of 1.5 km. Adult censuses were also

made concurrently. Of the considerable amount of data gathered, the most

significant and easily interpreted were provided by the egg censuses and this

is reported and analysed below. The eggs of D. argenthona and D. nigrina

are particularly easy to census, being laid in large clusters on their small,

discretely clumped food plants. Not only do the eggs of the two species differ

in their fine structure, easily seen with a hand lens, but also the two species

cluster their eggs in quite different patterns. Delias argenthona lays between

15-30 eggs in closely aggregated rows of 2-3 eggs, whereas D. nigrina lays

from 20-60 eggs in an almost circular pattern with the eggs well spaced.

Between 1982 and 1985, as part of a larger study of reproductive physiology

of Delias Hübner (Orr 1988), I conducted, within a large outdoor flight cage,

a short series of experiments on the response of female Delias to the presence

of eggs of their own and other species when choosing plants on which to lay.

These results are reported here in full.

Australian Entomologist, 2008, 35 (1) 29

Area and methods

Field study, 1975

The study area lay mainly along the southern shore of Currimundi Lake

(26°28'S, 153?10'E), then a semi-urbanised area with a broad strip of natural

growth ranging from 20-50 m wide along its southern boundary (Fig. 1).

Dominant trees were Melaleuca quinquinerva and Banksia integrifolia with

some Casuarina equisetifolia, all of which supported mistletoes. Crown

height of the trees was seldom more than six metres. (The area is still largely

intact today, but the trees have grown much taller and many mistletoes have

been removed by misguided environmental vigilantes). Within this area, and

in several nearby gardens, 33 individual mistletoe plants were located within

easy reach (only 5 others were present, beyond reach). A standard sampling

path of 1.5 km, including all reachable plants, was established beginning at

point ‘A’ and ending at point ‘B’ in Fig. 1.

Currimundi Habitat Reserve

Currimundi Lake

Fig. 1. Map of the study area in 1975 showing location of mistletoe plants (solid

circles) relative to Currimundi Lake and major streets. ‘A’ and ‘B’ mark the start and

finish of the standard transect.

When first located in February 1975, mistletoes were identified to species,

numbered and tagged. An estimate was made of the volume of each plant by

measuring, with a metre rule, three principle orthogonal axes. The volume

was calculated by taking half the mean of these three measurements and

calculating the volume of a sphere of this radius. This measurement was

correlated quite closely with total foliage and therefore measured the relative

amount of food available on each plant.

30 Australian Entomologist, 2008, 35 (1)

From 15 February 1975 to 29 September 1975, the plants were examined

every 3-4 days for new egg masses. Once located, the eggs in the mass were

counted and the mass marked as having been recorded by tying a small twist

of fine cotton thread (white for D. argenthona, red for D. nigrina) to the base

of the leaf on which they had been laid. Only new egg batches were recorded

during a sampling session. Because the incubation time of eggs of both

Delias species is about 5 days, this protocol ensured that all Delias eggs laid

on the sample mistletoes would be recorded once and only once.

Oviposition experiments with captive butterflies, 1982-3

Butterflies were maintained in one of two 4 m x 3 m x 2.5 m outdoor

insectaries where they spent their entire lives, having been raised in captivity

and introduced to a cage soon after eclosion. As a rule, no more than 5

females were kept in a cage at any time and mostly these were all of one

species. During oviposition experiments only one female was allowed in the

main cage. On different occasions, 12 individual female D. argenthona and

2] individual female D. nigrina were presented with three sprigs of fresh

Dendrophthoe vitellina placed in different corners of the cage. This was done

at a time when females were known to be highly gravid and ready to oviposit.

Each sprig had approximately 8-12 leaves and, of the three, one bore a single

batch of D. nigrina eggs and one a single batch of D. argenthona eggs, while

one (the control) was free of eggs. Often it was necessary to artificially attach

an egg mass to the sprig in order to obtain the desired experimental material.

The reactions of ovipositing females were noted, especially the first choice of

oviposition site. As cut mistletoe retains its turgor for only 2-3 hours, most

often the females oviposited only once before the plant had to be replaced

and, in this account, only the first oviposition choice of the female was

recorded even if a second egg batch was laid.

Results

Oviposition patterns in the field

Mistletoe species were distributed without any apparent pattern along the

transect, except for some obvious clumping around thickets of food trees

(Fig. 2). Note that Fig. 2 shows the distribution of plants along the path

walked, which was far from being a straight line, and hence the relative

dispersion of mistletoe plants is somewhat approximated.

Of the 33 plants included in the study, Amyema congener and Dendrophthoe

vitellina were present in almost the same numbers but D. vitellina

represented, by a small margin, the greater volume of foliage. Diplatia

furcata, with seven individuals, was present in moderate numbers and

Muellerina celastroides was uncommon, with only three individuals present

(Table 1).

Australian Entomologist, 2008, 35 (1) 31

Delias argenthona actively oviposited from February to late May and again

from late August to late September, when observations ceased (Fig. 3).

Delias nigrina actively oviposited from early April until late September,

peaking in activity during the winter months (Fig. 3). There were thus about 8

weeks during which both species were abundant and reproductively active.

These periods corresponded closely with sightings of adult butterflies.

Fig. 2. Distribution of mistletoe species along the sample path; heights of bars

indicate estimated leaf volume. Mistletoe species are as follows: A, Amyema

congener; D, Dendrophthoe vitellina; F, Diplatia furcata; M, Muellerina celastroides.

The spread of individual plants has been exaggerated slightly in places to

accommodate lettering.

Period of overlap

SS ee

Period of overlap

NUMBER OF EGG MASSES

feb mar apr may jun jul aug sep

TIME OF YEAR (MONTH)

Fig. 3. Phenology of appearance of new egg masses of D. argenthona (—) and D.

nigrina (---) between February and September 1975. Periods of overlap and potential

competition are indicated.

32 Australian Entomologist, 2008, 35 (1)

The dispersion of eggs on the food plants was complex and undoubtedly was

influenced by the availability of fresh growth on each plant, which varied

from plant to plant throughout the season. Although it was thought that egg

masses of each species were dispersed so as to avoid those of their

conspecifics, no simple measurement could demonstrate this conclusively.

However, D. argenthona and D. nigrina showed distinct differences in food

plant utilisation: D. argenthona never oviposited on Amyema congener,

whereas D. nigrina never oviposited on Diplatia furcata (Table 2).

Table 1. Abundance of various mistletoe species and percentage of total foliage

volume represented by each species.

Species number of plants % of total foliage volume

Muellerina celastroides 3 153)

Diplatia furcata 7 14.2

Amyema congener 12 34.0

Dendrophthoe vitellina 11 44.5

Table 2. Numbers of egg batches laid by D. argenthona and D. nigrina on four

mistletoe species in the presence or absence of the other Delias species. Df, Diplatia

furcata, Ac, Amyema congener, Dv, Dendrophthoe vitellina, Mc, Muellerina

celastroides.

Delias spp. Df Ac Dv Mc Total

D. argenthona alone 12 0 58 9 79

D. argenthona: D. nigrina present 31 0 11 3 45

D. nigrina alone 0 67 46 5 118

D. nigrina: D. argenthona present 0 52 9 7 68

In the absence of D. nigrina, D. argenthona oviposited mainly on

Dendrophthoe vitellina and the proportion of egg masses on the three food

plants used was nearly proportional to the amount of foliage available, with a

slight and perhaps insignificant preference for D. vitellina (Table 2).

However, when D. nigrina was present D. argenthona switched to using

mainly Diplatia furcata, the species not attacked by D. nigrina. This

difference is highly significant (Chi? [2df] = 36.9, p « 0.0001).

Similarly, D. nigrina, in the absence of D. argenthona, oviposited almost

equally on Amyema congener and Dendrophthoe vitellina, exhibiting a slight

preference for the former in terms of available foliage. However, in the

presence of D. argenthona, D. nigrina oviposited mainly on Amyema

congener, the species not used by D. argenthona. This shift is highly

significant (Chi? [2df] = 14.75, p « 0.001).

Australian Entomologist, 2008, 35 (1) 33

Oviposition on the uncommon mistletoe species Muellerina celastroides

showed no obvious patterns, it being used in moderation by both species.

Since M. celastroides appeared to be a less favoured plant and was

represented by only three individuals, no conclusions can be drawn from the

data which, however, have been included in the above analyses.

Oviposition experiments with captive butterflies

Table 3 shows the number of egg masses laid by each Delias species on three

classes of Dendrophthoe vitellina sprig offered: ARG included D.

argenthona eggs, CONTROL was free of other eggs and NIG included D.

nigrina eggs.

Table 3. Number of egg masses laid by different captive females on sprigs of

mistletoe free of eggs (CONTROL), or carrying an egg mass of either D. argenthona

(ARG) or D. nigrina (NIG).

Delias spp. ARG CONTROL NIG Total

D. argenthona 2 (1796) 10 (8396) 0 12

D. nigrina 4 (19%) 14 (67%) 3 (1496) 21

For D. argenthona, of 12 egg batches produced 10 (8396) were on the control

sprig and two batches (17 %) were on sprigs that already bore D. argenthona

eggs; none was laid on sprigs with D. nigrina eggs. The avoidance of sprigs

bearing eggs of other females was clearly significant (Chi? [1df] = 14.0, p <

0.001), but the sample size is too small to show if females avoid eggs of D.

nigrina more then those of their own species. However, on one occasion in an

unrelated study, a female D. argenthona laid a second batch of eggs on a

plant with D. nigrina eggs. Before oviposition she scraped away with her

forelegs the D. nigrina eggs, which were freshly laid and not adhering firmly

to the leaf, before placing her own eggs on another leaf. As this was not part

of the choice experiment the result is not included in Table 3.

For D. nigrina, a similar avoidance of leaves already bearing eggs was

evident, with 14 (6796) of egg batches laid on control sprigs. This is also

highly significant (Chi? [1df] = 10.57, p « 0.01), but there was no evidence to

suggest that, when avoiding egg masses already present, females

differentiated between eggs of their own species and those of D. argenthona.

Discussion

The results clearly show that when both Delias species are present, switching

occurs in the food plants utilised, with both D. argenthona and D. nigrina

tending to oviposit more on the mistletoe species which each utilises

exclusively. From experiments in captivity, it is evident that ovipositing

females avoid plants with eggs of their own and other species. It is, however,

unclear if heterospecific eggs are more of a deterrent to oviposition than

34 Australian Entomologist, 2008, 35 (1)

conspecific eggs but this seems likely to be the case. It is almost certain the

butterflies would be able to ascertain the species of eggs present on a plant, as

the visual pattern of the egg clusters is very different for the two species. The

most probable explanation for the food plant switching observed is that it is a

response to visual detection of egg clusters of the competing species on the

shared food plants. The oviposition behaviour of Delias lends support to this

hypothesis; females seldom commit to depositing an egg cluster without

having first closely inspected, visually, every fresh leaf on a plant. On the

other hand, testing of leaf quality with the ovipositor and/or forelegs is rather

perfunctory.

While individual plants varied a good deal in their suitability to sustain

larvae, with a few too small to nourish a large cohort and some completely

defoliated during the course of the study, there was no evidence to suggest

that the switching of D. argenthona from Dendrophthoe vitellina to Diplatia

furcata, or of D. nigrina from Amyema congener to Dendrophthoe vitellina

when D. argenthona finished its season, was related to a change in the quality

of the food plants. Diplatia furcata is probably an inferior food plant for D.

argenthona, as larvae reared in captivity on this food develop more slowly

and survive less well than those reared on Dendrophthoe vitellina (Orr,

unpublished observations). In June, many Dendrophthoe vitellina plants had

been severely eaten by cohorts of both species yet, once D. argenthona

finished its season, D. nigrina still showed a significant switch to this food

plant, possibly because of its essentially superior nutritional qualities.

Such examples of interspecific competition are quite rare in nature; much

more often we are witness to ‘the ghost of competition past’ (Connell 1980).

The fact that D. argenthona and D. nigrina are largely allotopic and

allochronic means that they can maintain populations which never compete

with each other and so seldom come under pressure to diverge ecologically.

This is especially true if populations are fairly open, as seems to be the case

from mark-release-recapture data (Orr, unpublished data). The abundance of

the high-quality mistletoe Dendrophthoe vitellina in the coastal wallum

seems to provide an unusually rich resource over which they compete

intermittently. In the study area, population levels of both Delias species

regularly reached higher densities than at any other locality I know. Such a

situation of an isolated but extremely rich resource is doubtless conducive to

competition.

It is interesting to speculate on what might be the competitive relationships

among the montane Delias of New Guinea. Very often, in a given locality

closely related species pairs tend to show different altitudinal preferences

(Parsons 1998). However, this does not exclude competition among members

of different species groups. In two weeks in December 1973, at one locality

on Mount Kaindi in Morobe Province, I collected 12 species, at least half of

which were abundant and only three of which could be considered rare. Even

Australian Entomologist, 2008, 35 (1) 35

more impressively, during December 1973 and January 1974, the total

number of species I collected on the same mountain at different altitudes

above 1000 m was 21, about a third of the known Delias fauna for mainland

Papua New Guinea. Ecological segregation under these circumstances must

surely be a complex process.

Acknowledgements

I am grateful to the staff of the Queensland Herbarium for identifying

mistletoe species. My parents G.H. and D.A. Orr assisted extensively with

data collection and other logistic help. A pilot study in 1974 was encouraged

and supervised by Peter Dwyer.

References

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

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

BRABY, M.F. 2006. Evolution of larval food plant associations in Delias Hübner butterflies

(Lepidoptera: Pieridae). Entomological Science 9: 383-398.

BRABY, M.F. and LYONNS, K.A. 2003. Effect of temperature on development and survival in

Delias nigrina (Fabricius) (Lepidoptera: Pieridae). Australian Journal of Entomology 42: 138-

143.

COALDRAKE, J.E. 1961. The ecosystem of the coastal lowlands (‘wallum’) of southern

Queensland. CSIRO Bulletin 283. CSIRO, Melbourne; x + 138 pp.

CONNELL, J.H. 1980. Diversity and coevolution of competitors, or the ghost of competition

past. Oikos 35: 131-138.

NOUSALA, A.K. 1979. A study of the temperature-dependent rates of development of two

species of pierid butterfly: Delias nigrina and D. argenthona. BSc thesis, Griffith University,

Brisbane.

ORR, A.G. 1988. Mate conflict and the evolution of the sphragis in butterflies. PhD thesis,

Griffith University, Brisbane; xvi + 348 pp.

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

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

36 Australian Entomologist, 2008, 35 (1)

MISCELLANEOUS NOTES

The following notes on new or interesting distribution and food plant records

for butterflies and moths are abstracted from the News Bulletin of the

Entomological Society of Queensland and were first published during 2006 or

2007 in the volumes and parts indicated.

Pelopidas agna dingo Evans [Hesperiidae] — On 8 November 2001, at a site 10 km

east of the Cradle Mountain turnoff in the northern Tasamanian highlands (called

‘Post Office Tree’), a specimen was collected together with a female Argynnina

hobartia montana L.E. and R. Couchman [Satyridae] resting on a small bush about

100 m from the car park. The specimen was sent to Cliff Meyer in Canberra, who

confirmed its identity. The nearest recorded locality for this species is in northern

New South Wales and it is suggested that the specimen might have been carried to the

site inadvertently by vehicle from Queensland. — A remarkable record of Pelopidas

agna dingo Evans from Tasmania — Ian Knight — 33(8): 158-159 (2006).

Ogyris amaryllis amaryllis (Hewitson) [Lycaenidae] — Established colonies were

found breeding on Amyema cambagei mistletoes on Casuarina Sp. around the shores

of Fennel Bay on Lake Macquarie, near Newcastle, New South Wales. The species

was abundant and both larvae and pupae were located. — Notes from Newcastle — Paul

Bambach — 34(2): 43 (2006).

Delias aganippe (Donovan) [Pieridae] — Established colonies were found breeding on

Amyema cambagei mistletoes on Casuarina sp. around the shores of Fennel Bay on

Lake Macquarie, near Newcastle, New South Wales. The species was abundant and

both larvae and pupae were located. — Notes from Newcastle — Paul Bambach — 34(2):

43 (2006).

Cephonodes spp. [Sphingidae] — The following three species of Cephonodes Hübner

have been recorded from a garden in the suburb of Corinda in Brisbane, SE

Queensland. (1) Cephonodes hylas australis Kitching & Cadiou: collected January

and November, sightings during October, December and March; larvae feeding on

Gardenia augusta [Rubiaceae] during November; possible confusion of sightings with

the more northerly C. picus (Cramer) noted. (2) C. janus janus (Miskin): one adult

sighted and confidently identified on 15 February 1998 flying around flowers of

Duranta repens [Verbenaceae]; a rare species in SE Qld, known previously from

Brisbane by a specimen collected [by the author] at Bowen Hills in December 1956

[illustrated]. (3) C. kingii (W.S. Macleay): common around Duranta and Eucalyptus

flowers, collected October-March; larvae feeding on Gardenia augusta from

November-April; in 1976 bred from Canthium coprosmoides [Rubiaceae] at Long

Pocket, Brisbane. — Bee-hawks and bird-hawks (Sphingidae: Macroglossinae) at

Corinda, Brisbane — Murdoch De Baar — 35(2): 35-37 (2007).

Macroglossum spp. [Sphingidae] — The following two species of Macroglossum

Scopoli have been recorded from a garden in the suburb of Corinda in Brisbane, SE

Queensland. (1) Macroglossum hirundo errans Walker: a regular visitor collected in

February, March and May and sighted on 9 July 2001; previously recorded

September-May in SE Qld. (2) M. micaceum micaceum (Walker): collected from

January-March and sighted in December. — Bee-hawks and bird-hawks (Sphingidae:

Macroglossinae) at Corinda, Brisbane — Murdoch De Baar — 35(2): 35-37 (2007).

Australian Entomologist, 2008, 35 (1): 37-46 37

DIVERSITY OF QUEENSLAND PARADOXOSOMATID

MILLIPEDES (DIPLOPODA: POLYDESMIDA:

PARADOXOSOMATIDAE)

R. MESIBOV

Queen Victoria Museum and Art Gallery, Launceston, Tas 7250

(E-mail: mesibov@southcom.com.au)

Abstract

Paradoxosomatids are the most widespread and abundant native millipedes in mainland

Australia. The Queensland paradoxosomatid fauna currently consists of five inadequately

described species, 28 adequately described species and 199 new and undescribed species. Future

collecting in non-rainforest habitats can be expected to raise the State total by at least another

10% to 250 species.

Introduction

Of the eight native millipede orders in Australia, Polydesmida is the most

diverse and currently accounts for ca 60% of described Australian species

(Mesibov 2006-07). Most polydesmidans in mainland Australia are in

Paradoxosomatidae, the largest of all millipede families and native to every

continent except North America and Antarctica (Hoffman 1982).

Thirty-one species of paradoxosomatids have been described from

Queensland (Table 1). The list is slightly inflated as five of the names refer to

inadequately described species whose types are either missing or the wrong

sex (only mature male polydesmidans can be confidently identified). Of the

remaining 26 species, only eight have been described in the past 65 years.

Two more paradoxosomatids are here added to the State list from the fauna of

northern New South Wales (Table 1), bringing the total of named Queensland

paradoxosomatids to 33 and the total of recognisable named species to 28.

These tallies ignore three subspecies noted in Table 1; two were described at

the same time as the nominate subspecies, while the third was described in

1987. For taxonomic details of all species listed, see Mesibov (2006-07).

Thousands of specimens of Queensland Paradoxosomatidae have been added

to museum collections in recent years, chiefly by Geoff Monteith and

colleagues at the Queensland Museum and by CSIRO entomologists based at

the Australian National Insect Collection. In this article I report on a recent

sorting of that material, which has yielded a remarkably large number of new

species and added many new locality records for known species.

Materials and methods

Results summarised here are from an unpublished study for the Natural

Heritage Assessment Section of the Department of the Environment and

Heritage (now Department of the Environment and Water Resources),

Canberra. The DEH project was carried out by Mark Harvey (Western

Australian Museum), Cathy Car (Charles Sturt University) and myself in

2006 and 2007 and required us to sort to species all native

38 Australian Entomologist, 2008, 35 (1)

Paradoxosomatidae in Australian museums. We also did a limited amount of

additional collecting.

Museum collections contain paradoxosomatids collected by hand, pitfall

trapping, flight intercept trapping, pyrethrum knock-down and Berlese

extraction of leaf litter and moss. The only specimens reliably identifiable to

species are mature males. These are sorted on the structure of the modified

legs known as gonopods, which are used to transfer sperm during mating. For

the DEH project, mature males (and sometimes associated females) were

removed from mixed samples, sorted, relabelled and separately registered.

The Queensland subset of the DEH project results is my own work. Named

species were identified using descriptions and redescriptions in the taxonomic

literature. New species were assigned, where feasible, to existing genera (see

Results). Species were documented with label information, museum

registration numbers, colour images, gonopod drawings, brief notes on key

characters and distribution maps. This documentation, in HTML format, is

available on CD-ROM in the Queensland Museum with restricted access. All

sorted specimen lots have been labelled with name or sorting code.

For project purposes, a small number of locality records with a spatial

uncertainty greater than + 10 km were excluded. The final list of 913 one-

species locality records for Queensland consisted of:

744 records from the Queensland Museum (Brisbane),

95 from the Australian National Insect Collection (Canberra),

27 from the Australian Museum (Sydney),

10 from Museum Victoria (Melbourne),

3 from the Western Australian Museum (Perth),

1 from the South Australian Museum (Adelaide), and

33 from museums overseas (literature records for types and vouchers).

Each locality was defined as a unique latitude/longitude pair. Most pairs were

taken from the collecting-event database maintained by the Queensland

Museum. Other pairs were taken from specimen labels after checking

agreement with locations expressed in words, or were assigned to such

locations using the Geoscience Australia gazetteer, paper maps or the

Australian National Insect Collection online specimen database.

Results

Species diversity

Twenty-one of the 28 adequately described species were sorted from museum

collections (Table 1, Fig. 1). In most cases the sortings were rediscoveries of

taxa not reported since their first description. It might be expected that the list

of 28 would be biased towards relatively abundant and widespread species

and, of the 17 species recorded from 10 or more localities, 15 have names.

Australian Entomologist, 2008, 35 (1) 39

Table 1. List of described species of Queensland Paradoxosomatidae. Under Status, Y

— adequately described, ? — inadequately described (e.g., known from female (F)

only), + = new records, - = no new records since description. Under Ist Year is given

the known or bounded date of first collection; 1910-13 — E. Mjóberg expeditions,

1891-93 — R. Semon expeditions. The two species marked with an asterisk (*) are

NSW species reported here from Qld for the first time. >

Species Status lst year

Atropisoma elegans Silvestri, 1897 ?,Fonly «1897

Aulacoporus affinis Verhoeff, 1924 Y,- 1910-13

Aulacoporus castaneus Verhoeff, 1924 YI 1910-13

Aulacoporus yarrabahnus Verhoeff, 1924 ES 1910-13

Australodesmus divergens Chamberlin, 1920 YE «1920

Brochopeltis mjoebergi Verhoeff, 1924 (2 subspp.) Y 1910-13

Cladethosoma uncinatum Jeekel, 1987 Y, - 1910-13

Desmoxytoides hasenpuschorum Mesibov, 2006 ygs: 1971

‘Eustrongylosoma’ transversefasciatum Silvestri, 1897 ?,Fonly | «1897

Helicopodosoma vittigerum Verhoeff, 1924 Y, - 1910-13

Heterocladosoma asperum (L. Koch, 1867) Y «1867

Heterocladosoma bifalcatum (Silvestri, 1898) Y6 3r 1891

Heterocladosoma hamuligerum (Verhoeff, 1924) Y 1910-13

Heterocladosoma trabeatum Jeekel, 1987 Ner «1987

Heterocladosoma tranversetaeniatum (L. Koch, 1867) Vi <1867;

(2 subspp.) 1924

Mjoebergodesmus annulatus Verhoeff, 1924 YEG 1910-13

Paraustraliosoma malandense (Verhoeff, 1924) YE 1910-13

*Parwalesoma rubriventris (Verhoeff, 1928) YA 1890

Phyllocladosoma annulatipes (Verhoeff, 1924) Y, + 1910-13

Phyllocladosoma broelemanni (Verhoeff, 1941) YA 1936?

‘Polydesmus (Strongylosoma)' dubium L. Koch, 1867 Y <1867

‘Polydesmus (Strongylosoma)’ rubripes L. Koch, 1867 ?,Fonly «1867

Pseudostrongylosoma sjoestedti Verhoeff, 1924 Yat 1910-13

Solaenodolichopus annulatus Verhoeff, 1941 Y, - 1936

(= Solaenodolichopus pruvoti (Broelemann, 1931))

Solaenodolichopus teres (Verhoeff, 1924) Y,- 1910-13

Solaenodolichopus vittatus (Verhoeff, 1924) (2 subspp.) Y, + 1910-13

Streptocladosoma albovittatum Jeckel, 1980 Y,- 1945

Streptocladosoma dissimile Jeekel, 1980 Y, - 1948

Streptocladosoma solum Jeekel, 1987 Y, + 1980

‘Strongylosoma’ semoni Attems, 1898 ?,F only 1891-93

Tholerosoma corrugatum Mesibov, 2006 VEE 1986

Tholerosoma monteithi Mesibov, 2006 Y, + 1983

*Walesoma helmsii Verhoeff, 1928 Y; 1890

40 Australian Entomologist, 2008, 35 (1)

Fig. 1. Some Queensland paradoxosomatid millipedes. Top left: undescribed

Solaenodolichopus, Border Ranges. Top right: Mjoebergodesmus annulatus, Cairns

area. Bottom left: Tholerosoma monteithi, Wet Tropics; note coating of soil particles

on each body ring. Bottom right: undescribed Paraustraliosoma, Massey Range.

Scale bar for Paraustraliosoma = 5 mm, others = 10 mm.

Both Heterocladosoma — transversetaeniatum (L. Koch, 1867) and

Solaenodolichopus vittatus (Verhoeff, 1924) have published subspecies. Both

are represented by a large number of specimens in the collections and

substantial variation is apparent in both, exceeding the variation used to

diagnose subspecies. These two species are probably best regarded as species

complexes pending further work on the taxonomy of the respective genera.

Another 199 species were sorted (Fig. 1), none of which was identified as one

of the well-known ‘tramp’ paradoxosomatids of the Pacific region (Shelley

and Lehtinen 1998). I referred 96 of these new natives to existing genera.

Two of the genera, Antichiropus Attems, 1911 and Notodesmus Chamberlin,

1920, have not previously been reported from Queensland. One monotypic

genus, Brochopeltis Verhoeff, 1924, retains that special status, but all other

monotypic genera expanded during the sorting: Australodesmus Chamberlin,

Australian Entomologist, 2008, 35 (1) 41

1920 (now contains 7 spp.), Mjoebergodesmus Verhoeff, 1924 (8 spp.),

Paraustraliosoma Verhoeff, 1924 (20 spp.), Pseudostrongylosoma Verhoeff,

1924 (2 spp.) and Walesoma Verhoeff, 1928 (5 Queensland spp., 2 New

South Wales spp.) The greatest expansion was in Solaenodolichopus

Verhoeff, 1924, a well-defined genus which has grown in Queensland from

three to 47 sorted species.

The remaining 103 new species have been assigned to codes rather than

genera. The relationships of these species are far from clear and for sorting

purposes I decided not to attempt to 'stretch" diagnoses of existing genera to

accommodate oddities. A large proportion of coded species are much smaller

as adults (6-12 mm long) than any previously described Australian

paradoxosomatids and were collected mainly by the Berlese method.

Fig. 2. Localities for Paradoxosomatidae sorted to species in Queensland.

42 Australian Entomologist, 2008, 35 (1)

Biogeography and sampling

The sole Queensland record for Cladethosoma uncinatum Jeekel, 1987,

‘Christmas Creek’, is geographically ambiguous and is excluded in the

analysis that follows.

Paradoxosomatids were recorded from 431 localities in Queensland, largely

near the coast (Fig. 2). More than half the 226 species (117) were recorded

from one locality only and only 32 species (14%) were recorded from five or

more localities.

n 278 28

H26

No. of species

0 50 100 150

No. of collecting events

Fig. 3. Increase of species recorded with search effort. Each point is from a latitude

class from 11° to 28°S; the latitude class is marked next to the point. No species were

recorded from 14°S. See text for further explanation.

Most species were recorded in the Wet Tropics and in southeastern

Queensland, which were also the most intensively sampled areas. The effect

of this sampling bias is indicated graphically in Fig. 3. Here the number of

species recorded in each 1° latitude class is plotted against search effort,

Australian Entomologist, 2008, 35 (1) 43

approximated as the number of collecting events in that latitude class; each

collecting event is a unique locality and collecting date (the day, month and

year collecting ceased in the case of traps left open for long periods).

Although the relationship looks roughly linear, it should not be concluded

that if every 1? latitude-block near the coast were searched with equal effort,

then the same number of species would be collected. Some parts of

Queensland are undoubtedly more paradoxosomatid-rich than others.

However, as Fig. 3 suggests, collecting to date has not unequivocally

demonstrated that the Wet Tropics and the southeast are dramatically richer

in paradoxosomatid species.

Biogeographical patterns

Heterocladosoma bifalcatum (Silvestri, 1898) was first reported from Cairns

but Jeekel (1987), who described a specimen from Colosseum [85 km S of

Gladstone], suspected the Cairns record to be an error. Recently, Rowe and

Sierwald (2006) redescribed this conspicuous and locally abundant species

from the Sydney area. All other records from the DEH project (Fig. 4) are

from southeastern Queensland, from near Childers south to the New South

Wales border. It seems likely that H. bifalcatum is a southeast Queensland

native that has been introduced elsewhere. It joins another Australian

paradoxosomatid, Akamptogonus novarae (Humbert & de Saussure, 1869), in

the interesting class of ‘native exotics’: species introduced and established

well outside their native range but still within their broader native region. For

more information on A. novarae, believed to be an eastern Australian native

but now resident in Tasmania, Western Australia, Norfolk Island, New

Zealand and Hawaii, see Mesibov (2006-07).

New locality records for the other named species have not greatly extended

their known ranges. However, enough records have now accumulated to

reveal mosaic parapatry between species in Aulacoporus Verhoeff, 1924,

Australodesmus Chamberlin, 1920, Heterocladosoma Jeekel, 1968 (Fig. 4),

Mjoebergodesmus Verhoeff, 1924, Paraustraliosoma Verhoeff, 1924,

Phyllocladosoma Jeekel, 1968, Solaenodolichopus Verhoeff, 1924 and

Walesoma Verhoeff, 1928. Mosaic parapatry (distributions arranged like

adjacent tiles in a mosaic, with very little overlap) is common in millipede

genera (Mesibov 2003) and was first reported for Queensland

paradoxosomatids in Tholerosoma (Mesibov 2006).

Another biogeographical pattern in the new locality data is high species

turnover across the Black Mountain Barrier (BMB) between Cairns and

Mossman in the Wet Tropics. The BMB is regarded as a former dry-forest

barrier between two rainforest refugia (Schneider et al. 1998, Yeates and

Monteith in press). Among the small number of paradoxosomatids occurring

on both sides of the BMB, Mjoebergodesmus annulatus Verhoeff, 1924 is

remarkable for having two colour morphs: the typical dark/light annulated

form in the Cairns area and an all-dark form west of Mossman.

44 Australian Entomologist, 2008, 35 (1)

—

Fig. 4. Localities for Heterocladosoma bifalcatum (crosses) and H. hamuligerum

(squares) in SE Queensland. Scale bar = 200 km.

Discussion

Paradoxosomatidae in Australia have an enormously wide ecological range.

They are abundant in tropical rainforest in northern Queensland and in cool

temperate rainforest in northwestern Tasmania (Mesibov 2000), in sandy

coastal heaths and in tall, dense forest. Their dry-country limits have not yet

been mapped, but specimens collected well inland in Queensland were sorted

in this study. Paradoxosomatids have also been collected by Mark Harvey

(pers. comm.) in semi-arid Western Australia and by Cathy Car (pers.

comm.) in semi-arid New South Wales. To judge from the DEH project

sorting, larger species of Heterocladosoma and Solaenodolichopus (25-40

mm long as adults) may be particularly successful in the Queensland dry

country.

Australian Entomologist, 2008, 35 (1) 45

With further searching in dry, inland areas, it is reasonable to expect the

Queensland paradoxosomatid species total to increase by 10% to ca 250

species. The real total may be much higher if narrow-range endemicity is as

prevalent in western Queensland as it is in the coastal strip. Entomologists

working in the dry country can assist in discovering this diversity by

collecting surface-active millipedes at night and after rain. Specimens can be

killed, preserved and stored in 70-80% ethanol and I recommend that they be

deposited in the Queensland Museum.

While the discovery of millipede diversity is relatively straightforward, the

documentation of that diversity suffers from a lack of specialists. It will be

many years before the backlog of new Australian genera and species created

by the DEH project can be cleared by formal taxonomic publication.

Acknowledgements

I am very grateful to Cameron Slatyer, formerly head of the Natural Heritage

Assessment Section (DEH), for supporting this work with funds and

enthusiasm. For their patience and assistance with work in collections I thank

Graham Milledge (Australian Museum); Natalie Barnett, Matt Colloff and

Jacquie Recsei (Australian National Insect Collection); Peter Lillywhite and

Rachel McBride (Museum Victoria); Mark Harvey and Julianne Waldock

(Western Australian Museum); and especially Robert Raven and Owen

Seeman (Queensland Museum).

References

HOFFMAN, R.L. 1982. Diplopoda. Pp 689-724, in: Parker, S.P. (ed. Synopsis and

classification of living organisms, Vol. 2. McGraw-Hill, New York; 1232 pp.

JEEKEL, C.A.W. 1987. Millipedes from Australia, 11: Australiosomatini from Queensland

(Diplopoda, Polydesmida, Paradoxosomatidae). Beaufortia 37: 11-41.

MESIBOV, R. 2000. An overview of the Tasmanian millipede fauna. Tasmanian Naturalist 122:

15-28.

MESIBOV, R. 2003. Lineage mosaics in millipedes. African Invertebrates 44: 203-212.

MESIBOV, R. 2006. Dirt-encrusted and dragon millipedes (Diplopoda: Polydesmida:

Paradoxosomatidae) from Queensland, Australia. Zootaxa 1354: 31-44.

MESIBOV, R. 2006-07. Millipedes of Australia (website). Accessed 6 August 2007.

http:/Avww.qvmag.tas.gov.au/zoology/millipedes/index.html

ROWE, M. and SIERWALD, P. 2006. Morphological and systematic study of the tribe

Australiosomatini (Diplopoda: Polydesmida: Paradoxosomatidea: Paradoxosomatidae) and a

revision of the genus Australiosoma Brólemann. Invertebrate Systematics 20: 527-556.

SCHNEIDER, C.J., CUNNINGHAM, M. and MORITZ, C. 1998. Comparative phylogeography

and the history of endemic vertebrates in the Wet Tropics rainforests of Australia. Molecular

Ecology 7: 487-498.

SHELLEY, R.M. and LEHTINEN, P.T. 1998. Introduced millipeds of the family

Paradoxosomatidae on Pacific Islands (Diplopoda: Polydesmida). Arthropoda Selecta 7(2): 81-

94.

46 Australian Entomologist, 2008, 35 (1)

SILVESTRI, F. 1898. Note sui Chilopodi e Diplopodi conservati nel Museo Zoologico di

Firenze. I. Alcuni nuovi Diplopodi de Queensland (Cairns). Bullettino della Società

Entomologica Italiana 29: 225-232. PC

YEATES, D.K. and MONTEITH, G.B. (in press). The invertebrate fauna of the Wet Tropics:

diversity, endemism and relationships. In: Stork, N. and Turton, S. (eds), Living in a dynamic

tropical forest landscape. Blackwell Publishing.

Australian Entomologist, 2008, 35 (1): 47-56 47

NEW ANT-LYCAENID ASSOCIATIONS AND BIOLOGICAL DATA

FOR SOME AUSTRALIAN BUTTERFLIES (LEPIDOPTERA:

LYCAENIDAE)

ROD EASTWOOD!, MICHAEL F. BRABY??, DAVID J. LOHMAN* and

ALAN KING?

! Museum of Comparative Zoology, Harvard University, 26 Oxford St., Cambridge MA-02138,

USA

"Biodiversity Conservation Division, Department of Natural Resources, Environment and the

Arts, PO Box 496, Palmerston, NT 0831

*School of Botany and Zoology, The Australian National University, Canberra, ACT 0200

‘Department of Biological Sciences, National University of Singapore, 14 Science Drive 4,

Singapore 117543

ŠPO Box 1302, GPO Townsville, Old 4810

Abstract

Records of 61 ant-lycaenid associations and other ecological data are tabulated for 27 Australian

species of Lycaenidae. Thirty-one of the ant-lycaenid associations and four larval food plant

records are new. Ant-lycaenid records are discussed in further detail for 11 lycaenid species from

the genera Paralucia Waterhouse & Turner, Hypochrysops C. & R. Felder, Ogyris Angas,

Hypolycaena C. & R. Felder, Deudorix Hewitson, Nesolycaena Waterhouse & Turner,

Catopyrops Toxopeus and Theclinesthes Rober.

Introduction

The early stages of many lycaenid butterflies associate with ants. These

associations vary considerably among species, ranging from non-specific

facultative to highly specific obligate myrmecophily or even parasitism

(myrmecophagy) by the butterfly larvae (Pierce et al. 2002). Evolutionary

factors that may have promoted this variation in ant attendance levels are not

well understood. However, differences in the intensity of ant-lycaenid

associations, among butterfly taxa whose evolutionary relationships are

known, can be contrasted in a phylogenetic framework to address questions

such as whether facultative associations evolved before obligate associations

and whether parasitism is derived from mutualism.

By measuring population genetic structure in pairs of lycaenid sister species

that differ in the degree to which they rely on ants as mutualistic partners, it

might be possible to infer the role that ants play in lycaenid demographics

and, hence, their role in lycaenid diversification (e.g. Eastwood et al. 2006).

Underpinning these broad research objectives is the need for complete and

accurate biological and ecological data; thus it is important to publish such

baseline records. This paper builds on earlier works documenting ant-

lycaenid associations (Common and Waterhouse 1981, Eastwood and Fraser

1999, Schmidt 2002), as well as providing new biological data for several

Australian lycaenid species.

48 Australian Entomologist, 2008, 35 (1)

Methods and observations

Observations of lycaenid biology and the collection of their associated ants

were undertaken opportunistically by us and several other lepidopterists

between 1991 and 2007. Voucher specimens of ants identified by A.

Andersen are deposited at CSIRO Sustainable Ecosystems, Darwin (TERC);

Camponotus Mayr identified by A. McArthur are deposited in the South

Australian Museum, Adelaide (SAM); Polyrhachis Smith identified by R.

Kohout are held at the Queensland Museum, Brisbane (QM); and all

remaining species are deposited in the Museum of Comparative Zoology at

Harvard University, USA (MCZ). Butterfly nomenclature largely follows

Braby (2000), with modifications to comply with the International Code of

Zoological Nomenclature. Ant nomenclature follows Shattuck (1999), with

additional nomenclature from R. Kohout (pers. comm.).

In total, 61 ant-lycaenid associations, of which 31 are new, are listed in Table

1, together with four new food plant associations and incidental observations.

Ant-lycaenid associations which duplicate previously published records are

included for spatial and temporal information only. Additional biological and

ecological data are discussed below for selected lycaenid species.

Paralucia pyrodiscus (Doubleday)

P. pyrodiscus has been recorded only in association with Notoncus Emery

ants (Common and Waterhouse 1972, Eastwood and Fraser 1999). Our

records bring the number of attendant species of Notoncus to four, namely N.

capitatus Forel, N. ectatommoides (Forel), N. gilberti Forel and N. enormis

Szabó. The closely related taxa Paralucia aurifera (Blanchard) and P.

spinifera Edwards & Common both associate with Anonychomyrma

Donisthorpe ants; thus it is possible that the specificity of ant association

within the Paralucia Waterhouse & Turner lineage has contributed to their

diversification.

Hypochrysops digglesii (Hewitson)

This obligate myrmecophile has been recorded only in association with

Crematogaster Lund ants (Common and Waterhouse 1972, Eastwood and

Fraser 1999, Schmidt 2002). During a population explosion at Indooroopilly

in 2003, two different Crematogaster species were recorded in attendance.

The specificity of ant association for H. digglesii appears to be constrained at

the generic level (i.e. to Crematogaster).

Hypochrysops polycletus rovena Druce

The attendant ant recorded here, Iridomyrmex sanguineus Forel, is a large

‘meat ant’ in the purpureus species group (Andersen 2000). In addition to

attending H. p. rovena larvae on the food plant, the ants constructed shelters

with sand and bark particles for mature larvae and pupae at the base of the

plant. Although the behaviour of shelter building for butterfly larvae is not

Australian Entomologist, 2008, 35 (1)

Table 1. Ant attendance records and biological data for lycaenid butterfly larvae and

pupae in Australia.

Lycaenid Attendant ant

species

Lucia limbaria Iridomyrmex

(Swainson) rufoniger

(Lowne)

Acrodipsas Crematogaster sp.

cuprea (Sands) near laeviceps*

Smith

Paralucia Anonychomyrma

aurifera sp. (nitidiceps

(Blanchard) group)

(E.André)

P. pyrodiscus —— Notoncus

(Doubleday) enormis*

Notoncus sp.

(enormis

group)*

Hypochrysops Crematogaster

digglesii Spp.

(Hewitson)

H. polycletus Iridomyrmex

rovena Druce — sanguineus*

Iridomyrmex sp.

H. apelles

apelles

(Fabricius)

9 group C

H. ignita Papyrius sp. 1

erythrinus (nitidus group)

(Waterhouse & (Mayr)

Lyell)

Ogyris abrota

(Westwood)

Crematogaster sp.

Voucher

Repository

AA145

(MCZ)

(TERC)

MFB06

(TERC)

00.045

(MCZ)

(TERC)

00.033

(MCZ)

00.050

(MCZ)

00.052

(MCZ)

(TERC)

MFB07

(MCZ)

GFOI

(MCZ)

Date and location

22.11.2005

Darlington Point,

NSW (34°34’02”S,

146*00' 12"E)

19.x.2002

Burrewarra Point,

NSW

19.vii.2004

Tallaganda NP,

9 km E of

Hoskinstown, NSW

(35°24’S, 149*32' E)

25.ix.2001

1.4 km N of Isla

Gorge, Qld

15.1.2005

Kowen Escarpment,

8 km NNE

Queanbeyan, ACT,

650 m (35°17’26"S,

149?15'28"E)

23.iv.2003

Indooroopilly, Qld

5.xii.1999

Mt Stuart,

Townsville, Qld

7.xii.1999

Townsville, Qld

03.vi.2006

Buffalo Creek,

Leanyer Swamp,

Darwin, NT

(12°21719"S,

130?54'20"E)

15.iii.2006

Litchfield Nat Park,

NT (13°02’S,

130°55’E)

13.vii.1997

Mt Mugga Mugga,

ACT

26.11.2006

Tocumwal, NSW

Notes

LFP Oxalis perennans

LFP Acacia mearnsii

Numerous ants

associated with eggs

on trunks of mature

trees

LFP Bursaria spinosa

Numerous ants attending

early stages at base of

small plants

LFP Bursaria spinosa

Numerous ants attending

early stages at base of

small plants

LFP Rhyssopterys

timorensis

LFP Rhyssopterys

timorensis

LFP Lumnitzera

racemosa

Numerous ants attending

three late instar larvae

LFP Planchonia careya

Numerous ants attending

three larvae on small

regenerating plants

LFP Muellerina

eucalyptoides

Table 1. Continued.

Lucia lie eee Ao _ v ao_voe0t e _ .r...«rl

Lycaenid

species

O. oroetes

(Hewitson)

“Arid form’

O. oroetes

oroetes

(Hewitson)

O. olane

(Hewitson)

O. amaryllis

meridionalis

(Bethune-

Baker)

O. amaryllis

amata

(Waterhouse)

Hypolycaena

phorbas

(Fabricius)

Attendant ant

Anonychomyrma

sp. Donisthorpe

Crematogaster sp.

Mongiceps*

Forel

Crematogaster sp.

?longiceps*

Technomyrmex sp.

Mayr

Tetramorium

simillimum*

(Smith)

Crematogaster sp.

Podomyrma

adelaidae*

(Smith)

Iridomyrmex sp.

Crematogaster sp.

laeviceps? group

Smith

Paratrechina sp.*

Tetramorium

bicarinatum*

(Nylander)

Australian Entomologist, 2008, 35 (1)

Voucher Date and location Notes

Repository

00.036 — 16.x.1999 Attending pupa

(MCZ) 11 km S of Amelup,

Stirling Ra, WA

(TERC) 14.x.2005 LFP Amyema miquelii

Finke River, 6 km S Three ants each

Hermannsburg, attending two late

Hermannsburg instar larvae/prepupae

Aboriginal L.T., NT; under loose bark at

560 m (23°59'28"S, base of Eucalyptus

132°46'32"E) camaldulensis

(TERC) 30.vi.2006 LFP Amyema miquelii

Trephina Gorge Nat One or two ants

Park, East Macdonnell attending a few larvae

Ranges, NT; 550m on Eucalyptus with

(23?31'41"S, numerous clumps of

134?22'48"E) mistletoe; most larvae

not attended.

00.046 — 17.viii.1999 -

(MCZ) Brisbane, Qld

00.032 — 8.vi.2000 Ants attending prepupa

(MCZ) Capalaba, Qld

00.035 — 27.1.2001 -

(MCZ) Mt McKenzie, NSW

RE-02- 26.xi.2002 -

A188 l0km W of

(MCZ) Bordertown, SA

00.034 27.xi.1999 -

(MCZ) Kalbarri, WA

(TERC) vii.1997 LFP Amyema cambagei

Murrumbidgee River, A few ants attending

Uriara Crossing, larva in hollow twig

ACT of mistletoe

parasitising

Casuarina

cunninghamii; most

larvae not tended.

00.051 15.xii.2000 LFP Clerodendrum sp.

(MCZ) Upper Ross River,

14 km SW of

Townsville, Qld

00.047 4.iii.2000 LFP Clerodendrum

(MCZ) Aitkenvale, longiflorum

Townsville, Qld

——————————————————— ———

Australian Entomologist, 2008, 35 (1) 51

Table 1. Continued.

Lycaenid Attendant ant Voucher Date and location Notes

species Repository

Deudorix diovis Paratrechina sp. 00.026-29 14.xi.2000 LFP Cupaniopsis sp.

Hewitson (MCZ) East Ballina, NSW

Crematogaster sp. 00.025 13.xi.2000 LFP Cupaniopsis sp.

(MCZ) Broken Head, NSW

Pheidole No voucher Brisbane Attending pupa

variabilis* Mayr

P. megacephala 00.031 14.xi.2000 LFP Cupaniopsis sp.

(Fabricius) (MCZ) East Ballina, NSW

Rhytidoponera 00.030 — 14.xi.2000 LFP Cupaniopsis sp.

metallica* (MCZ) East Ballina, NSW

(Smith)

D. smilis Crematogaster (TERC) 2.vi.2007 LFP Strychnos lucida

dalyensis (Le Group A sp. 3 Lee Point, Casuarina — 1-2 ants associated with

Souéf & (CSIRO)* Coastal Reserve, NT; pupae inside fruits.

Tindale) (12°19°54”S,

130°53’40”E)

Candalides Technomyrmex sp. | RE-03- 17.1.2003 Ants attending larvae

absimilis B118 Chapel Hill,

(Felder) (MCZ) Brisbane, Qld

Nesolycaena Polyrhachis gab* (TERC) 15.iv.2006 LFP Boronia lanceolata

urumelia Forel Litchfield Nat Park, Two ants attending one

(Tindale) NT; 190 m late instar larva; most

(13?07'32"S, larvae not attended

130?48' 11"E)

Monomorium sp. 8 (TERC) 15.iv.2006 LFP Boronia lanceolata

(carinatum Litchfield Nat Park, Four ants attending one

group)* Heterick NT; 190 m early instar larva;

(13°07’32”S, most larvae not

130°48°11”E) attended

Nacaduba Polyrhachis 00.037 19.xii.2000 Attending larvae

berenice (Chariomyrma) (QM) — Fisherman's Island,

berenice aurea* Mayr Brisbane, Qld

(Herrich-

Schaffer)

Catopyrops Iridomyrmex sp.* 99.036 — 9.iii.1999 LFP Trema tomentosa

florinda halys Paratrechina sp.* (MCZ) Griffith Uni.

Waterhouse Nathan Campus, Qld

Paratrechina sp.* 99.001 15.xii.1998 LFP Trema tomentosa

(MCZ) Griffith Uni.

Nathan Campus, Qld

Paratrechina sp.* 00.023 26.iv.2000 LFP Pipturis sp.

(MCZ) Macgregor, Brisbane,

Qld

Tetramorium 00.024 — 26.iv.2000 LFP Pipturis sp.

bicarinatum* (MCZ) Macgregor, Brisbane,

Qld

——————————————————D

Table 1. Continued.

Lycaenid Attendant ant

species

Theclinesthes Polyrhachis

onycha onycha

(Hewitson)

(Hagiomyrma)

lydiae* Forel

T. onycha Monomorium sp.*

capricornia

Sibatani &

Grund

T. miskini Iridomyrmex sp.

miskini

(Lucas)

Iridomyrmex

reburrus*

Shattuck

Iridomyrmex

reburrus*

Iridomyrmex

reburrus*

Iridomymex sp. 21

(gracilis group)*

Iridomyrmex sp.

(mattiroloi

group)* Emery

Iridomyrmex sp.

(anceps group)

(Roger)

T. miskini Iridomyrmex sp.

eucalypti

Sibatani &

Grund

Voucher

Repository

GF02

(QM)

00.039

(MCZ)

RE-02-

A475

(MCZ)

(TERC)

00.040

(MCZ)

Australian Entomologist, 2008, 35 (1)

Date and location

5.iv.2006

Chapel Hill, Qld

13.xii.1991

Cardwell, Qld

13.1.2003

Edungalba Turnoff,

Qld

15.iv.2006

Litchfield Nat Park,

NT (13?0732"S,

130?48' L I"E) 190 m

17.11.2007

Black Point, Cobourg

Peninsula, NT

(11.15515°S,

132.14391°E)

14.1.2007

Berry Springs, NT

(12.67576°S,

131.00876°E)

14.1.2007

Berry Springs, NT

(12.67576°S,

131.00876°E)

3.ii.2007

Cobourg Peninsula,

NT (11.15015°S,

132.16125°E)

28.1.2007

Muirella Park,

Kakadu NP, NT

(12.85462°S,

132.75468°E)

2.xii.199]

Cardwell, Qld

Notes

LFP Cycas media

LFP Acacia difficilis

Numerous ants

attending six

prepupae/pupae

LFP Acacia

auriculiformis

Numerous ants on small

plant with eggs; a few

ants attending | larva

LFP Eucalyptus

Numerous ants

attending larvae and

pupae on small

regenerating plant

LFP Eucalyptus

Numerous ants

attending larvae and

pupae on small

regenerating plant

LFP Acacia holosericea

A few ants attending

larvae

LFP Acacia holosericea

A few ants attending

one early instar larva

and two prepupae

LFP Corynbia

clarksoniana

Seedling about 20 cm

high (record based on

female ovipositing on

plant that was

covered in swarms of

small black ants)

€ cs ee

Australian Entomologist, 2008, 35 (1)

Table 1. Continued.

Lycaenid

species

T. miskini

eucalypti

Sibatani &

Grund (cont.)

T. albocincta

(Waterhouse)

T. serpentata

serpentata

(Herrich-

Schaffer)

T. sulpitius

(Miskin)

Lampides

boeticus

(Linnaeus)

Famegana

alsulus alsulus

(Herrich-

Schaffer)

Freyeria putli

putli (Kollar)

Attendant ant

Iridomyrmex sp.

Camponotus

maculatus

humilior* Forel

Opisthopsis

haddoni* Emery

Iridomyrmex sp.

Iridomyrmex sp.*

Dolichoderus

scrobiculatus*

(Mayr)

Iridomyrmex sp.

(suchieri

group)* Forel

Iridomyrmex

?septentrionalis*

Forel

Iridomyrmex

gracilis group*

(Lowne)

Opisthopsis

haddoni*

Tetramorium

deceptum*

Bolton

Voucher

Repository

00.042

(MCZ)

00.049

(SAM)

00.048

(MCZ)

00.043

(MCZ)

00.038

(MCZ)

00.044

(MCZ)

00.041

(MCZ)

MFB09

(TERC)

MFB04

(TERC)

MFB03

(TERC)

MFB02

(TERC)

MFB08

(TERC)

Date and location

4.iii.1992

Cardwell, Qld

7.1.2001

Mt Stuart,

Townsville, Qld

71.2001

Mt Stuart,

Townsville, Qld

22.xii.1997

Eastern Lookout,

Wyperfield NP,

Vic.

7.iii.1991

Rainbow, Vic.

25.1.1999

Wapengo Lagoon,

11 km NNE of

Tathra, NSW

24.viii.1991

Cleveland Bay Beach,

Townsville, Qld

27.11.1992

Ilbilbie, Qld

29.iv.1992

Campaspe River,

17.5 km NE of

Pentland, Qld

27 .iii.1992

Ilbilbie, Qld

27.11.1992

Cardwell, Qld

17.11.1992

Townsville, Qld

6.11.1992

Townsville, Qld

Notes

LFP Acacia crassicarpa

LFP Xylomelum

scottianum'

LFP Xylomelum

scottianum'

LFP Adriana tomentosa

LFP Rhagodia sp.

LFP Atriplex sp.

A few ants attending

larvae

LFP Suaeda australis

LFP Indigofera

pratensis flowers!

LFP Crotalaria

goreensis flowers

LFP Indigofera

pratensis flowers

LFP Indigofera

pratensis flowers

LFP Indigofera hirsuta

LFP = larval food plant; * = new ant-lycaenid association; t = new larval food plant

record; letters in brackets indicate the repository for ant specimens (see Observations).

24 Australian Entomologist, 2008, 35 (1)

uncommon among attendant ants (e.g. Camponotus with Ogyris genoveva

(Hewitson) and Papyrius Shattuck with Hypochrysops ignita (Leach)), such

behaviour has not been recorded previously in the purpureus species group.

H. p. rovena is likely to be an obligate myrmecophile, as are most species of

Hypochrysops C. & R. Felder whose life histories are known. However, it

appears to have non-specific ant associates, being recorded in the wild with at

least three ant species from two subfamilies (Muller 1998, Eastwood and

Fraser 1999, Table 1).

Ogyris oroetes (Hewitson)

The two new ant association records listed here (Table 1) bring the total

number of attendant ant species for this facultatively ant-associated butterfly

species to 13 (Eastwood and Fraser 1999, Schmidt 2002, Table 1).

Hypolycaena phorbas phorbas (Fabricius)

H. phorbas is considered to be an obligate myrmecophile that is typically

tended by the green tree ant Oecophylla smaragdina (Fabricius) (Common

and Waterhouse 1981). The atypical ant associates (Tetramorium Mayr and

Paratrechina Motschulsky spp.) recorded here were found tending H.

phorbas at two locations near Townsville, Queensland. In addition to

attending the butterfly larvae, both species of ants were found feeding on

phloem exuding from leaf edges damaged by larval feeding by the butterfly.

Although Clerodendrum species (Verbenaceae) are known larval food plants

for H. phorbas (Common and Waterhouse 1981), C. longiflorum Decne. is

previously unrecorded.

Deudorix diovis Hewitson

Larvae are not commonly attended by ants because they feed inside the fruit

of their food plant. The ants recorded here were attending prepupal larvae and

pupae attached to the base of food plants or to accumulated debris in forks or

other convenient locations. Schmidt (2002) recorded at least nine ant species

associated with larvae and pupae of D. diovis in Brisbane, Queensland and

the two additional records here confirm the non-specificity of ant-associates

for this lycaenid. While non-specific ant association is regarded as a defining

characteristic of facultative myrmecophiles (Fiedler 1996), ant-association

during the prepupal and pupal phase may be obligatory for D. diovis (also see

Eastwood et al. 2005).

Deudorix smilis dalyensis (Le Souéf & Tindale)

Two ant species, Crematogaster sp. (Myrmicinae) and Oecophylla

smaragdina (Formicinae) are now recorded in association with the early

stages of this lycaenid. The butterfly larvae and pupae are usually found

without ants, but when ants are present their numbers are low (Eastwood and

Fraser 1999). Unlike D. diovis, described above, D. smilis dalyensis pupates

inside fruit of the larval food plant and it is likely to have only a facultative

association with ants.

Australian Entomologist, 2008, 35 (1) 35

Nesolycaena urumelia (Tindale)

Both of the ant species recorded in attendance are new records (Table 1).

This is another facultative myrmecophile: larvae are not usually tended by

ants, ant-association is non-specific and attendance levels, when present, are

low.

Catopyrops florinda halys Waterhouse

This subspecies is found commonly around Brisbane, Queensland, in autumn.

Ants rarely attend early instar larvae, which rest under young leaves of the

larval food plant alongside the midrib, but later instars are occasionally

attended. At Macgregor, Brisbane, larvae were attended by two ant species

(Paratrechina sp. and Tetramorium sp.) as they fed on two different branches

of a cultivated Pipturis sp. tree (Urticaceae). Similarly at Griffith University

(Nathan Campus), Brisbane, several Paratrechina and a single Iridomyrmex

Mayr species, most likely 7. gracilis (Lowne), were found in attendance on a

small Trema tomentosa (Roxb.) Hara (Ulmaceae).

Theclinesthes onycha (Hewitson)

The two ant associations listed for this species in Table 1 were each recorded

in attendance with two subspecies of T. onycha, namely T. onycha

capricornia Sibatani & Grund in northern Queensland and 7. onycha onycha

in southeastern Queensland. These two new records bring the total number of

attendant ants to nine (Eastwood and Fraser 1999, Table 1) and confirm the

non-specific nature of their ant associations in contrast with the larval food

plant specificity of the two butterfly subspecies, which feed exclusively on

Cycadaceae in the north and on Zamiaceae in the south (Braby 2000).

Theclinesthes miskini (Lucas)

Two new larval food plants for this species are recorded here together with

two new ant associations (Table 1). This brings the total number of food

plants for this taxon to 29 from five families (Braby 2000) and attendant ants

to at least 12 species from eight genera (Eastwood and Fraser 1999). Non-

specific ant association appears to be typical for Theclinesthes Rober species,

all of which are regarded as facultative myrmecophiles. However, all other

Theclinesthes species or subspecies are ecologically specialised in terms of

their food plant range, as are most facultative myrmecophiles (Pierce and

Elgar 1985). The wide larval food plant range coupled with non-specificity of

ant associates in 7. miskini is unusual and may indicate some evolutionary

instability in the taxon.

Acknowledgements

We thank Fabian Douglas, Graham Forbes, Ann Fraser, John Nielsen and

John Moss for their observations and for supplying specimens. Alan

Andersen (CSIRO, Darwin), Chris Burwell (Queensland Museum, Brisbane),

Stefan Cover (Harvard University, Cambridge, USA), Katsuyuki Eguchi

56 Australian Entomologist, 2008, 35 (1)

(Kagoshima University Museum, Japan), Ann Fraser (Kalamazoo College,

Kalamazoo, USA), Rudy Kohout (Queensland Museum, Brisbane), and

Archie McArthur (South Australian Museum, Adelaide) kindly identified

ants for this study.

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THE AUSTRALIAN

Entomologist

Volume 35, Part 1, 10 March 2008

C 0 NTENTS

EASTWOOD, R., BRABY, M.E, LOHMAN, D.J. AND KING, A.

New ant-lycaenid associations and biological data for some Australian

butterflies (Lepidoptera: Lycaenidae).

GOLLAN, J.R., BATLEY, M. AND REID, C.A.M.

The exotic bee Halictus smaragdulus Vachal, 1895 (Hymenoptera:

Halictidae) in the Hunter Valley, New South Wales.

KENT, D.S.

Distribution and host plant records of Austroplatypus incompertus

(Schedl) (Coleoptera: Curculionidae: Platypodinae).

"FL

go MR. e

A tk on buterfly (Lepidoptera) records from the Darwin Region,

Northern Territory.

LACHLAN, R:B.+

Additional records of hawk moths, and bi butterflies (Lepidoptera) from

Lizard island? Northern Qu unu r

MESIBOY, R.

Diversity of Queensland paradoxosomatid millipedes (Diplopoda:

Polydesmida: Paradoxosomatidae).

ORR, A.G.

Competition for larval food plant between Delias argenthona

(Fabricius) and Delias nigrina (Fabricius) (Lepidoptera: Pieridae) in

coastal wallum habitat in southern Queensland.

WEBBER, B.L., CURTIS, A.S.0., CASSIS, G. AND WOODROW, I.E.

Flowering morphology, phenology and flower visitors of the Australian

rainforest tree Ryparosa kurrangii (Achariaceae).

MISCELLANEOUS NOTES

ISSN 1320 6133