Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

Taxonomy, ecology, genetics and conservation status of the pale imperial hairstreak (Jalmenus eubulus) (Lepidoptera : Lycaenidae): a threatened butterfly from the Brigalow Belt, Australia

Rod Eastwood A D E , Michael F. Braby B C , Daniel J. Schmidt A and Jane M. Hughes A
+ Author Affiliations
- Author Affiliations

A Australian Rivers Institute, Griffith School of Environment, Griffith University, Nathan, Queensland 4111, Australia.

B Biodiversity Conservation Division, Department of Natural Resources, Environment and the Arts, PO Box 496, Palmerston, NT 0831, Australia.

C School of Botany and Zoology, The Australian National University, Canberra, ACT 0200, Australia.

D Present address: Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA.

E Corresponding author. Email:

Invertebrate Systematics 22(4) 407-423
Submitted: 25 July 2006  Accepted: 23 May 2008   Published: 14 October 2008


The taxonomic status of Jalmenus eubulus Miskin stat. rev. is revised and considered to be specifically distinct from J. evagoras (Donovan) based on fundamental differences in morphology, ecology and genetics. Miskin’s holotype is fixed by monotypy and illustrated, with type locality Rockhampton, Queensland. Fixed differences in the mitochondrial genomes of J. eubulus and J. evagoras in which the mean pairwise divergence is only 0.85% indicate absence of matrilineal gene flow, whereas allozyme data show significant structure within and between populations of both species consistent with recent diversification. Underlying causes for the observed genetic patterns are investigated. The two species are parapatric, with a narrow range of overlap along the Great Escarpment in south-eastern Queensland. Jalmenus eubulus is restricted to vegetation communities comprising brigalow-dominated old-growth open-forests and woodlands in the Brigalow Belt (with larvae monophagous on Acacia harpophylla F. Muell. Ex Benth), whereas J. evagoras occurs in a range of disturbed eucalypt woodlands/open-forests predominantly in montane and coastal areas east of this bioregion (with larvae polyphagous on Acacia species other than A. harpophylla). The conservation status of J. eubulus is considered to be vulnerable nationally and critically endangered in New South Wales according to International Union for Conservation of Nature (IUCN) criteria. Nationally, the geographic range has an estimated area of occupancy of less than 2000 km2, is severely fragmented, and the extent or quality of its habitat, which is poorly conserved, continues to decline. It is recommended that the taxon be used as an indicator for identification of remnant old-growth forest for conservation planning, as well as a flagship for the conservation of invertebrate biodiversity associated with this threatened ecological community.

Additional keywords: Acacia harpophylla, allozymes, biological species concept, Haldane’s rule, hybrid zone, mitochondrial DNA, parapatry.


We thank M. Breitfuss and C. Hill for allowing us to cite their unpublished observations of Jalmenus eubulus, B. Wilson for vegetation data on the extent of cleared and remnant areas in the Brigalow Belt bioregion, and J. Edwards for assistance with Graphical Information System analysis and figures depicting the distribution maps. We also thank N. E. Pierce for helpful comments on the manuscript. The following curators/collection managers kindly provided assistance with access to specimens held in their care: G. Daniels (University of Queensland Insect Collection), E. D. Edwards (Australian National Insect Collection), C. Burwell and G. Monteith (Queensland Museum), D. R. Britton (Australian Museum), B. Done (Museum of Tropical Queensland) and J. Forrest (South Australian Museum). G. Thompson assisted with photography of specimens in the Queensland Museum. A. F. Atkins, M. DeBaar, P. Hendry, J. F. R. Kerr, R. Mayo, J.St.L. Moss, D. P. A. Sands, B. Wilson, P. Wilson and J. Woinarski provided additional specimens for examination or literature. Butterflies sampled in this study were collected under Department of Environment and Heritage permit WISP00490302, Queensland National Parks permit WITK0040602 and State Forests permit TWB/17/2001–3 issued to the Queensland Entomological Society. This project was funded in part by an Australian Geographic Science Sponsorship to RE.


Accad A. , Neldner V. J. , Wilson B. A. , and Niehus R. E. (2006). Remnant vegetation in Queensland: analysis of remnant vegetation 1997–1999–2000–2001–2003, including regional ecosystem information. Queensland Herbarium, Environmental Protection Agency, Brisbane, Australia.

Akaike H. (1974) A new look at the statistical model identification. IEEE Transactions on Automatic Control 19, 716–723.
CrossRef | open url image1

Atkins A. (1976) New records for butterflies in southern, central and northern Queensland. Australian Entomological Magazine 3, 1–4. open url image1

Bailey A. (Ed.) (1984). ‘The Brigalow Belt of Australia. The Proceedings of a Symposium held at the John Kinder Memorial Theatre, Queensland Institute of Technology, 23 October 1982.’ (The Royal Society of Queensland: Brisbane, Australia.)

Birky C. W., Fuerst P., Maruyama T. (1989) Organelle gene diversity under migration, mutation, and drift: equilibrium expectations, approach to equilibrium, effects of heteroplasmic cells, and comparison to nuclear genes. Genetics 121, 613–627.
PubMed |
open url image1

Boland D. J. , Brooker M. I. H. , Chippendale G. M. , Hall N. , Hyland B. P. M. , Johnston R. D. , Kleinig D. A. , and Turner J. D. (1985). ‘Forest Trees of Australia.’ (Thomas Nelson: Melbourne, Australia.)

Braby M. F. (2000). ‘Butterflies of Australia: Their Identification, Biology and Distribution.’ (CSIRO Publishing: Melbourne, Australia.)

Braby M. F. (2004). ‘The Complete Field Guide to Butterflies of Australia.’ (CSIRO Publishing: Melbourne, Australia.)

Breitfuss M. J., Hill C. J. (2003) Field observations on the life history and behaviour of Jalmenus evagoras eubulus Miskin (Lepidoptera: Lycaenidae) in the southern brigalow belt of Queensland. Australian Entomologist 30, 135–138. open url image1

Burns A. N. , and Rotherham E. R. (1969). ‘Australian Butterflies in Colour.’ (Reed: Sydney, Australia.)

Burns J. M., Janzen D. H., Hajibabaei M., Hebert P. D. (2007) DNA barcodes of closely related (but morphologically and ecologically distinct) species of skipper butterflies (Hesperiidae) can differ by only one to three nucleotides. Journal of the Lepidopterists’ Society 61, 138–153. open url image1

Common I. F. B. (1964). ‘Australian Butterflies.’ (Jacaranda Press: Brisbane, Australia.)

Common I. F. B. , and Waterhouse D. F. (1972). ‘Butterflies of Australia.’ (Angus and Robertson: Sydney, Australia.)

Common I. F. B. , and Waterhouse D. F. (1981). ‘Butterflies of Australia,’ revised edn. (Angus and Robertson: Sydney, Australia.)

Costa J. T., McDonald J. H., Pierce N. E. (1996) The effect of ant associations on the population genetics of the Australian butterfly Jalmenus evagoras (Lepidoptera: Lycaenidae). Biological Journal of the Linnean Society 58, 287–306. open url image1

Covacevich J. (2000). Brigalow belt blues. Wildlife Australia. 36(4), 15–19.

Coyne J. A., Orr H. A. (1989) Patterns of speciation in Drosophila. Evolution 43, 362–381.
CrossRef | open url image1

Coyne J. A., Orr H. A. (1997) Patterns of speciation in Drosophila revisited. Evolution 51, 295–303.
CrossRef | open url image1

Coyne J. A. , and Orr H. A. (2004). ‘Speciation.’ (Sinauer Associates: Sunderland, MA, USA.)

D’Abrera B. (1971). ‘Butterflies of the Australian Region.’ (Lansdowne Press: Melbourne, Australia.)

DeBaar M. (1977a) Butterflies from an area between the Bunya Mountains and Archookoora State Forest, Queensland. Australian Entomological Magazine 3, 115–119. open url image1

DeBaar M. (1977b) New records for butterflies in Queensland and northern New South Wales. Australian Entomological Magazine 4, 11–12. open url image1

Dobzhansky T. (1935) A critique of the species concept in biology. Philosophy of Science 2, 344–355.
CrossRef | open url image1

Doyle J. J., Doyle J. L. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemistry Bulletin 19, 11–15. open url image1

Dunn K. L. , and Dunn L. E. (1991). ‘Review of Australian Butterflies: Distribution, Life History and Taxonomy. Parts 1–4.’ (Privately published by the authors: Melbourne, Australia.)

Dunn K. L. , Kitching R. L. , and Dexter E. M. (1994). The conservation status of Australian butterflies. Unpublished report, Australian National Parks and Wildlife Service, Canberra, Australia.

Eastwood R. (1999) An aberrant food-plant record for Jalmenus evagoras (Donovan) (Lepidoptera: Lycaenidae). Australian Entomologist 26, 13–14. open url image1

Eastwood R., Fraser A. M. (1999) Associations between lycaenid butterflies and ants in Australia. Australian Journal of Ecology 24, 503–537.
CrossRef | open url image1

Eastwood R., Pierce N. E., Kitching R. L., Hughes J. M. (2006) Do ants enhance diversification in the Lycaenidae (Lepidoptera)? Predictive patterns from the model myrmecophile Jalmenus evagoras. Evolution 60, 315–327.
PubMed |
open url image1

Edwards E. D. (1996). Lycaenidae. In ‘Checklist of the Lepidoptera of Australia. Monographs on Australian Lepidoptera, Volume 4’. (Eds E. S. Nielsen, E. D. Edwards and T. V. Rangsi.) pp. 249–254, 360–363. (CSIRO Publishing: Melbourne, Australia.)

Edwards E. D. , Newland J. , and Regan L. (2001). ‘Lepidoptera: Hesperioidea, Papilionoidea. Zoological Catalogue of Australia. Vol. 31.6’ (CSIRO Publishing: Melbourne, Australia.)

Excoffier L., Smouse P. E., Quattro J. M. (1992) Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data. Genetics 131, 479–491.
CAS | PubMed |
open url image1

Excoffier L. , Laval G. , and Schneider S. (2005). Arlequin ver. 3.0: An integrated software package for population genetics data analysis. Evolutionary Bioinformatics Online 1, 47–50. Available from

Fensham R. J., McCosker J. C., Cox M. J. (1998) Estimating clearance of Acacia-dominated ecosystems in central Queensland using land-system mapping data. Australian Journal of Botany 46, 305–319.
CrossRef | open url image1

Folmer O., Black M., Hoeh W., Lutz R., Vrijenhoek R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3, 294–299.
CAS | PubMed |
open url image1

Fraser A. M., Tregenza T., Wedell N., Elgar M. A., Pierce N. E. (2002) Oviposition tests of ant preference in a myrmecophilous butterfly. Journal of Evolutionary Biology 15, 861–870.
CrossRef | open url image1

Guo S. W., Thompson E. A. (1992) Performing the exact test of Hardy-Weinberg proportion for multiple alleles. Biometrics 48, 361–372.
CrossRef | CAS | PubMed | open url image1

Hajibabaei M., Janzen D. H., Burns J. M., Hallwachs W., Hebert P. D. N. (2006) DNA barcodes distinguish species of tropical Lepidoptera. Proceedings of the National Academy of Sciences of the United States of America 103, 968–971.
CrossRef | PubMed | open url image1

Haldane J. B. S. (1922) Sex ratio and unisexual sterility in animal hybrids. Journal of Genetics 12, 101–109. open url image1

Hancock D. L. (1995) The butterfly types of W. H. Miskin in the Queensland Museum (Lepidoptera). Memoirs of the Queensland Museum 38, 519–528. open url image1

Hebert P. D. N., Cywinska A., Ball S. L., deWaard J. R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London. Series B. Biological Sciences 270, 313–322.
CrossRef | CAS | open url image1

International Commission of Zoological Nomenclature (1999). ‘International Code of Zoological Nomenclature.’ (International Trust for Zoological Nomenclature: London, UK.)

IUCN (2001). ‘IUCN Red List Categories: Version 3.1.’ (IUCN Species Survival Commission: Gland, Switzerland.)

Johnson N. K., Cicero C. (2002) The role of ecological diversification in sibling speciation of Empidonax flycatchers (Tyrannidae): multigene evidence from mtDNA. Molecular Ecology 11, 2065–2081.
CrossRef | CAS | PubMed | open url image1

Kerr J. F. R., Macqueen J. (1967) Notes on the genus Jalmenus (Lepidoptera: Lycaenidae) with a description of a new species. Journal of the Australian Entomological Society 6, 45–48.
CrossRef | open url image1

Lloyd P. L. (1984). Agricultural and pastoral land use in the brigalow belt of Queensland. In ‘The Brigalow Belt of Australia’. (Ed. A. Bailey.) pp. 81–96. (The Royal Society of Queensland: Brisbane, Australia.)

Mayr E. (1942). ‘Systematics and the Origin of Species.’ (Columbia University Press: New York, USA.)

McCubbin C. (1971). ‘Australian Butterflies.’ (Thomas Nelson Australia Limited: Melbourne, Australia.)

Miskin W. H. (1876). Descriptions of new species of Australian diurnal Lepidoptera. Transactions of the Entomological Society of London 1876, 451–457.

Miskin W. H. (1883) On Ogyris genoveva, Hewitson, and its life-history. Transactions of the Entomological Society of London 1883, 343–345. open url image1

Miskin W. H. (1891) A synonymical catalogue of the Lepidoptera Rhopalocera (butterflies) of Australia with full bibliographical reference; including descriptions of some new species. Annals of the Queensland Museum 1, 1–93. open url image1

Monteith G. B., Yeates D. K. (1988) The butterflies of Mount Moffatt and Carnarvon National Parks, Queensland. Queensland Naturalist 28, 14–25. open url image1

Musgrave A. (1932). ‘Bibliography of Australian Entomology 1775–1930. With Biographical Notes on Authors and Collectors.’ (Royal Zoological Society of New South Wales: Sydney, Australia.)

Peters J. V. (1971). ‘A Catalogue of the Type Specimens of the Hesperioidea and Papilionoidea (Lepidoptera) in the Australian Museum.’ (Australian Entomological Press: Sydney, Australia.)

Pierce N. E. (1984). Amplified species diversity: a case study of an Australian lycaenid butterfly and its attendant ants. In ‘The Biology of Butterflies’. (Eds R. I. Vane-Wright and P. R. Ackery.) pp. 197–200. (Academic Press: London, UK.)

Pierce N. E. , and Nash D. R. (1999). The Imperial Blue, Jalmenus evagoras (Lycaenidae). In ‘Monographs on Australian Lepidoptera, 6: Biology of Australian Butterflies’. (Eds R. L. Kitching, E. Scheermeyer, R. E. Jones and N. E. Pierce.) pp. 279–315. (CSIRO Publishing: Melbourne, Australia.)

Pierce N. E., Braby M. F., Heath A., Lohman D. L., Mathew J., Rand D. B., Travassos M. A. (2002) The ecology and evolution of ant association in the Lycaenidae (Lepidoptera). Annual Review of Entomology 47, 733–771.
CrossRef | CAS | PubMed | open url image1

Posada D., Crandall K. A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics 14, 817–818.
CrossRef | CAS | PubMed | open url image1

Presgraves D. C. (2002) Patterns of postzygotic isolation in Lepidoptera. Evolution 56, 1168–1183.
PubMed |
open url image1

Pulsford I. F. (1984). Conservation status of brigalow Acacia harpophylla (Mimosaceae) in New South Wales. In ‘The Brigalow Belt of Australia’. (Ed. A. Bailey.) pp. 160–175. (The Royal Society of Queensland: Brisbane, Australia.)

QPWS (1994). ‘Queensland Nature Conservation (Wildlife) Regulation (1994).’ (Government Printer: Brisbane, Australia.)

Raymond M., Rousset F. (1995) GENEPOP, version 1.2: population genetics software for exact tests and ecumenism. The Journal of Heredity 86, 248–249. open url image1

Recher H. (2002) The past, present and future of biodiversity conservation in Australia. Pacific Conservation Biology 8, 8–11. open url image1

Richardson B. J. , Baverstock P. R. , and Adams M. (1986). ‘Allozyme Electrophoresis. A Handbook for Animal Systematics and Population Studies.’ (Academic Press: Sydney, Australia.)

Sands D. P. A. , and New T. R. (2002). ‘The Action Plan for Australian Butterflies.’ (Environment Australia: Canberra, Australia.)

Sands D. P. A., New T. R. (2008) Irregular diapause, apparency and evaluating conservation status: anomalies from the Australian butterflies. Journal of Insect Conservation 12, 81–85.
CrossRef | open url image1

Sattler P. S. , and Webster R. J. (1984). The conservation status of brigalow (Acacia harpophylla) communities in Queensland. In ‘The Brigalow Belt of Australia’. (Ed. A. Bailey.) pp. 149–160. (The Royal Society of Queensland: Brisbane, Australia.)

Simon C., Frati F., Beckenbach A., Crespi B., Liu H., Flook P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87, 651–701.
open url image1

Standards and Petitions Working Group (2006). ‘Guidelines for using the IUCN Red List Categories and Criteria: Version 6.1.’ (Standards and Petitions Working Group for the IUCN Species Survival Commission Biodiversity Assessments Subcommittee: Gland, Switzerland.)

State of Queensland (1994). Nature Conservation (Wildlife) Regulation 1994. SL No. 474. Reprint number 1. Government Printer, Brisbane, Australia.

Swofford D. L. (2002). ‘PAUP*: Phylogenetic Analysis Using Parsimony (*and Other Methods) version 4.0b10 for Macintosh.’ (Sinauer Associates: Sunderland, MA, USA.)

Taylor M. F. J., McKechnie S. W., Pierce N. E., Kreitman M. (1993) The lepidopteran mitochondrial control region: structure and evolution. Molecular Biology and Evolution 10, 1259–1272.
CAS | PubMed |
open url image1

Travassos M. A., Pierce N. E. (2000) Acoustics, context and function of vibrational signalling in a lycaenid butterfly-ant mutualism. Animal Behaviour 60, 13–26.
CrossRef | PubMed | open url image1

Valentine P. S. (2004) The demise of mass migration of the brown awl Badamia exclamationis (Fabricius 1775) (Lepidoptera: Hesperiidae): a consequence of land clearing in Queensland? Pacific Conservation Biology 10, 67–69. open url image1

Waterhouse G. A. (1903a) Notes on Australian Rhopalocera: Lycaenidae. Part III Revisional. Proceedings of the Linnean Society of New South Wales 28, 132–275. open url image1

Waterhouse G. A. (1903b) A catalogue of the Rhopalocera of Australia. Memoirs of the New South Wales Naturalists’ Club No. [i–ii.] 1, 1–49. open url image1

Waterhouse G. A. (1932). ‘What Butterfly is That?’ (Angus and Robertson: Sydney, Australia.)

Waterhouse G. A. , and Lyell G. (1914). ‘The Butterflies of Australia. A Monograph of the Australian Rhopalocera. Introducing a Complete Scheme of Structural Classification, and Giving Descriptions and Illustrations of All the Butterflies Found in Australia, Including a Number Now Recorded For the First Time.’ (Angus and Robertson: Sydney, Australia.)

White M. E. (1997). ‘Listen... Our Land is Crying. Australia’s Environment: Problems and Solutions.’ (Kangaroo Press: Sydney, Australia.)

Woinarski J. C. Z., Catterall C. P. (2004) Historical changes in the bird fauna at Coomooboolaroo, northeastern Australia, from the early years of pastoral settlement (1873) to 1999. Biological Conservation 116, 379–401.
CrossRef | open url image1

Young P. A. R. , Wilson B. A. , McCosker J. C. , Fensham R. J. , Morgan G. , and Taylor P. M. (1999). Brigalow Belt. In ‘The Conservation Status of Queensland’s Bioregional Ecosystems’. (Eds P. S. Sattler and R. D. Williams.) pp. 11/1–11/81. (Environmental Protection Agency: Brisbane, Australia.)

Appendix 1.  Distribution of Jalmenus eubulus collection sites
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