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Type: Article
Published: 2019-11-14
Page range: 48-77
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Extralimital Dynastinae (Coleoptera: Scarabaeidae) in Australia

Queensland Museum


Honorary Research Fellow
Department of Primary Industries and Regional Development (Quarantine WA)
Dynastinae introductions exotic species pest species quarantine biosecurity


The identification, origin, introduction, establishment, spread and pest status of extralimital (introduced, those that naturally occur within and outside Australia, and possibly introduced but of uncertain origin) dynastines in Australia and its territories are reviewed. We examine likely introduction pathways and the species’ life histories and attempt to draw out factors that predispose some species to establish, naturalise and then become invasive. Six categories of extralimital dynastines are identified: established and have spread widely (invasive)—Cyclocephala signaticollis Burmeister, 1847, Heteronychus arator (Fabricius, 1775); established but have not spread widely (sleepers)—Oryctes rhinoceros (Linnaeus, 1758), Oryctoderus latitarsus Boisduval, 1835 (possibly), Papuana woodlarkiana (Montrouzier, 1855) (possibly), Temnorhynchus retusus (Fabricius, 1781); known from one or only a few specimens that have arrived but not established—H. parumpunctatus Burmeister, 1847, Or. centaurus Sternberg, 1910, Pa. angusta Arrow, 1914, Pentodon algerinus algerinus (Fuessly, 1778); with distributions that extend from New Guinea into northeastern Australia—Pa. woodlarkiana (possibly), Od. latitarsus (possibly), Xylotrupes carinulus Rowland, 2011; whose records within Australia are doubtful—Dipelicus integriceps (Fairmaire, 1877), D. montrouzieri (Reiche, 1860), Or. nasicornis (Linnaeus, 1758), Pa. tibialis Arrow, 1941, Phyllognathus degener Fairmaire, 1891; species whose identity is unclear—Carne’s (1981) enigmatic species. Characters that make a species an ideal tramp species are: adults feed and are active for an extended period after emergence; larval development is relatively quick; adults and larvae are associated with potted plants and lawn grasses; the native distribution is in the Southern Hemisphere so that life cycles are synchronised between original and introduced localities. Shipping ballast and potted plants were the probable major pathways of introduction, but now rapid air and land transport provide the means for movement into and within Australia of hitchhiker species.


Key words: Dynastinae, introductions, exotic species, pest species, quarantine, biosecurity


  1. Agnew J. R. (Ed.). 1997. Australian Sugarcane Pests. Bureau of Sugar Experiment Stations, Indooroopilly, Australia.

    Allsopp P. G. 1986. Book review. The Dynastinae of the World. S. Endrődi. Journal of the Australian Entomological Society 25: 110.

    Allsopp P. G. 1987. An additional exotic dynastine Temnorrhynchus [sic] retusus (F.) (Coleoptera: Scarabaeidae) in Australia. Journal of the Australian Entomological Society 26: 189–191.

    Allsopp P. G. 1990. Phyllognathus carnei sp. n. and Oryctoderus latitarsis Boisduval (Coleoptera: Scarabaeidae: Dynastinae), two additions to the Australian fauna. Journal of the Australian Entomological Society 29: 229–232.

    Allsopp P. G. & Hitchcock B. E. 1987. Soil insect pests in Australia: control alternatives to persistent organochlorine insecticides. Standing Committee on Agriculture Technical Report Series 21: 1–123.

    Allsopp P. G. & Hutchinson P. M. 2018. The exotic whitegrub Plectris aliena Chapin (Coleoptera: Scarabaeidae: Melolonthinae) is more widely distributed in Australia than previously thought. Australian Entomologist 45: 1–6.

    Allsopp P. G., Samson P. R. & Chandler K. J. 2000. Pest management. In: Manual of Canegrowing (eds DM Hogarth & PG Allsopp) pp. 291–337. Bureau of Sugar Experiment Stations, Brisbane, Australia.

    Anonymous 1993. Relevamiento de los insectos del suelo en cultivos de papa del sudeste bonarerense. Boletín Técnico—Estación Experimental Regional Agropecuaria Balcare 118: 5–18.

    Anonymous 1995. Scarab grubs. Legislative Assembly for the ACT: 1995 Week 2 Hansard (11 May) p. 456. Available from URL: (Accessed 10 July 2018).

    Anonymous 2010. Biosecurity fact sheet. African black beetle (Heteronychus arator). Department of Primary Industries, Parks, Water and Environment, Tasmania, Hobart, Australia.

    Anonymous 2014. Port of Newcastle. Heritage and conservation register. Port of Newcastle and EJE Heritage. Available from URL: (Accessed 20 February 2019).

    Anonymous 2017. Hawaiian Scarab ID. Papuana. Available from URL: (Accessed 1 February 2018).

    Anonymous 2019. Western Australian Organism List. Available from URL: (Accessed 24 January 2019)

    Arrow G. J. 1911. Notes on the coleopterous subfamily Dynastinae, with descriptions of new genera and species. Annals and Magazine of Natural History (8)8: 151–176.

    Arrow G. J. 1914. Some further notes on lamellicorn beetles of the subfamily Dynastinae. Annals and Magazine of Natural History (8)14: 257–276.

    Arrow G. J. 1917. Some systematic notes on melolonthine Coleoptera. Annals and Magazine of Natural History (10)9: 189–197.

    Arrow G. J. 1937a. Systematic notes on beetles of the subfamily Dynastinae, with descriptions of new species in the British Museum collection (Coleoptera). Transactions of the Royal Entomological Society of London 86: 35–57.

    Arrow G. J. 1937b. Scarabaeidae: Dynastinae. In: Coleopterorum Catalogus. Pars 156 (ed S Schenkling), pp. 1–124. Junk, ʼs-Gravenhage, The Netherlands.

    Arrow G. J. 1941. Systematic notes on beetles of the family Dynastidæ and descriptions of a few new species. Annals and Magazine of Natural History 11(8): 273–283.

    Aulmann G. 1911. Neue Pimelopus-Arten (Coleopt.) schädlich an Kokospalmen. Entomologische Rundschau 28: 51–52.

    Barbero E. & Palestrini C. 2003. Coleoptera Trogidae, Geotrupidae, Hybosoridae, Scarabaeidae (Scarabaeinae and Dynastinae) collected in Australia. Monografie, Museo Regionale di Scienze Naturali, Torino 35: 303–328.

    Beard R. L. 1956. Two milky diseases of Australian Scarabaeidae. Canadian Entomologist 88: 640–647.

    Bedford G. O. 1974. Descriptions of the larvae of some rhinoceros beetles (Col. Scarabaeidae, Dynastinae) associated with coconut palms in New Guinea. Bulletin of Entomological Research 63: 445–472.

    Bedford G. O. 1980. Biology, ecology, and control of palm rhinoceros beetles. Annual Review of Entomology 25: 309–339.

    Bedford GO. 2013. Biology and management of palm dynastid beetles: recent advances. Annual Review of Entomology 58: 353–372.

    Beehag G., Kaapro J. & Manners A. 2016. Pest Management of Turfgrass for Sport and Recreation. CSIRO, Clayton, Australia.

    Berón C. M. & Diaz B. M. 2005. Pathogenicity of hyphomycetous fungi against Cyclocephala signaticollis. BioControl 50: 143–150.

    Blackburn T. 1888. Notes on Australian Coleoptera with descriptions of new species. Proceedings of the Linnean Society of New South Wales (2)3: 805–875.

    Blanchard C. É. 1846. Voyage au Pôle Sud et dans l’Océanie sur les Corvettes l’Astrolabe et la Zélée; exécuté par ordre du Roi pendant les années 1837-1838-1839-1840. Atlas zoologique. Gide, Paris, France.

    Boisduval J. B. A. D. de. 1835. Voyage de découvertes de l’Astrolabe exécuté par ordre du Roi, pendant les années 1826-1827-1828-1829, sous le commandement de M. J. Dumont d’Urville. Faune entomologique de l’Océan Pacifique, avec l’illustration des insectes nouveaux recueillis pendant le voyage. Part 2. Coléoptères et autres ordres. J. Tastu, Paris, France.

    Britton E. B. 1951. The nomenclature of Heteronychus sanctae-helenae Blanchard (Coleoptera: Scarabaeidae: Dynastinae). Proceedings of the Linnean Society of New South Wales 76: 133–134.

    Brooks J. G. 1965. New records of Coleoptera in Australia. Journal of the Entomological Society of Queensland 4: 85.

    Brown P. & Daigneault A. 2014. Cost-benefit analysis of managing the Papuana uninodis (Coleoptera: Scarabaeidae) taro beetle in Fiji. Journal of Economic Entomology 107: 1866–1877.

    Bryant L. M. & Krosch M. N. 2016. Lines in the land: a review of evidence for eastern Australia’s major biogeographical barriers to closed forest taxa. Biological Journal of the Linnean Society 119: 238–264.

    Bulinski J. & Matthiessen J. N. 2002. Poor efficacy of the insecticide chlorpyrifos for the control of African black beetle (Heteronychus arator) in eucalypt plantations. Crop Protection 21: 621–627.

    Bulinski J., Matthiessen J. N. & Alexander R. 2006. Development of a cost-effective, pesticide-free approach to managing African black beetle (Heteronychus arator) in Australian eucalyptus plantations. Crop Protection 25: 1161–1166.

    Burmeister H. C. C. 1847. Handbuch der Entomologie. Coleoptera Lamellicornia Xylophila et Pectinicornia. Volume 5. T.C.F. Enslin, Berlin, Germany.

    Camino N. B. & Achinelly M. F. 2012. A new species of the genus Cranifera Kloss, 1960 (Thelastomatidae, Nematoda) parasitizing larvae of Scarabaeidae (Coleoptera) from Argentina. Estudos de Biologia 34: 57–59.

    Camino N. B. & Reboredo G. R. 2005. A new Oxyurida (Thelastomatidae) from Cyclocephala signaticollis Burmeister (Coleoptera: Scarabaeidae) from Argentina. Journal of Parasitology 91: 890–892.

    Carne P. B. 1956. Cyclocephala signaticollis Burmeister, an introduced pasture scarab (Coleoptera). Proceedings of the Linnean Society of New South Wales 81: 217–221.

    Carne P. B. 1957. Systematic Revision of the Australian Dynastinae (Coleoptera: Scarabaeidae). CSIRO, Melbourne, Australia.

    Carne P. B. 1958. A review of the Australian Rutelinae (Coleoptera: Scarabaeidae). Australian Journal of Zoology 6: 162–240.

    Carne P. B. 1971. When was Cyclocephala signaticollis introduced to Australia? News Bulletin of the Australian Entomological Society 7: 131–132.

    Carne P. B. 1981. An undescribed and possibly exotic beetle (Scarabaeidae: Dynastinae) occurring on the south coast of New South Wales. Australian Entomological Magazine 7: 76–78.

    Cassis G. & Weir T. A. 1992. Dynastinae. In: Zoological Catalogue of Australia. Coleoptera: Scarabaeoidea. Volume 9 (ed WWK Houston) pp. 383–425. Australian Government Publishing Service, Canberra, Australia.

    Consolo V. F., Mucci V., Salerno G. L. & Berón C. M. 2010. Pathogenicity of bacterial isolates to Cyclocephala signaticollis (Coleoptera: Scarabaeidae). Biocontrol Science and Technology 20: 475–482.

    Cox J. & Dale P. S. 1977. New records of plant pests in New Zealand. II. New Zealand Journal of Agricultural Research 20: 109–111.

    Crawley M. J. 1987. What makes a community invasible? In: Colonization, Succession and Stability (eds AJ Gray, MJ Crawley & PF Edwards) pp. 429–453. Blackwell, Oxford, United Kingdom.

    Cumpston D. M. 1940. On the external morphology and biology of Heteronychus sanctae-helenae Blanch. and Metanastes vulgivagus Olliff (Col., Scarabaeidae, Dynastinae). Proceedings of the Linnean Society of New South Wales 65: 289–300.

    De Baar M. 1988. Insects collected during a trip to Torres Strait 27 March to 10 April, 1987. News Bulletin, Entomological Society of Queensland 15: 107–117.

    Dechambre R-P. 1986. Insectes Coléoptères Dynastidae. Fauna de Madagascar, 65. Muséum National d’Histoire Naturelle, Paris, France.

    Dechambre R-P. 1991. Désignation de lectotypes de Cyclocephala décrits par Burmeister [Coleoptera, Dynastidae]. Revue Française de Entomologie, Nouvelle Série 13: 123–124.

    Dechambre R-P. 1998. Siralus insularis, nouveau genre, nouvelle espèce de Pentodontini de Nouvelle-Guinée (Coleoptera, Dynastidae). Coléoptères 4: 29–34.

    Dechambre R-P. 1999. Note synonymique sur le genre Phyllognathus Eschscholtz, 1830 (Col. Dynastidae). Bulletin de la Société Entomologique de France 104: 182.

    Dechambre R-P. 2003. Le genre Onychionyx Arrow, 1914 (Coleoptera, Dynastidae). Coléoptères 9: 245–250.

    Dechambre R-P. 2005. Les Coléoptères du Monde. Volume 30: Dynastidae Australiens et Océaniens. Hillside Books, Canterbury, United Kingdom.

    Dechambre R-P. & Lachaume G. 2001. Les Coléoptères du Monde. Vol 27: Dynastidae. Le genre Oryctes. Hillside Books, Canterbury, United Kingdom.

    Dejean P. F. M. A. 1833. Catalogue des Coléoptères de la Collection de M. le Comte Dejean. Volume 2. Méquignon-Marvis, Paris, France.

    Drake J. M. 2004. Allee effects and the risk of biological invasion. Risk Analysis 24: 795–802.

    Elton C. 1958. The Ecology of Invasions by Animals and Plants. Methuen, London, United Kingdom.

    Endrődi S. 1951. Über die Gattung Xylotrupes. Acta Biologica Academiae Scientiarum Hungaricae 2: 239–253.

    Endrődi S. 1960. Coleoptera (Melolonthidae): Dynastinae. In: South African Animal Life. Results of the Lund University Expedition in 1950-1951. Volume 7 (eds B Hanström, P Brinck & G Rudebeck) pp. 34–82. Almqvist & Wiksell, Göteborg, Sweden,

    Endrődi S. 1961. Neue afrikanische Formen der Unterfamilien Dynastinae und Hybosorinae (Coleoptera Scarabaeidae). Revue de Zoologie et de Botanique Africains 63: 378–386.

    Endrődi S. 1963. Clysterius guineensis n. gen. n. sp. (Coleoptera, Melolonthidae, Dynastinae). Bonner zoologische Beiträge 14: 228–233.

    Endrődi S. 1966. Monographie der Dynastinae (Coleoptera, Lamellicornia). I. Teil. Entomologische Abhandlungen staatliches Museum für Tierkunde in Dresden 33: 1–457.

    Endrődi S. 1969. Monographie der Dynastinae. 4. Tribus Pentodontini. 2. Pentodontini der paläarktischen Region. Entomologische Abhandlungen staatliches Museum für Tierkunde in Dresden 37: 147–208.

    Endrődi S. 1971a. Monographie der Dynastinae. 2. Tribus: Oryctoderini (Coleoptera: Lamellicornia: Melolonthidae). Pacific Insects 13: 207–241.

    Endrődi S. 1971b. Monographie der Dynastinae. 4. Tribus: Pentodontini (papuanische und pazifische Inselwelt) (Coleoptera: Lamellicornia: Melolonthidae). Pacific Insects 13: 243–320.

    Endrődi S. 1973. Monographie der Dynastinae. 5. Tribus: Oryctini (Coleoptera, Lamellicornia, Melolonthidae). Entomologische Arbeiten aus dem Museum G. Frey 24: 1–87.

    Endrődi S. 1974a. Monographie der Dynastinae (Col. Lamellicornia, Melolonthidae) 4. Tribus: Pentodontini der äthiopischen Region. Entomologische Arbeiten aus dem Museum G. Frey 25: 4–108.

    Endrődi S. 1974b. Monographie der Dynastinae 4. Tribus Pentodontini von Australien und Neu-Zeeland (Coleoptera). Folia Entomologica Hungarica 27: 17–70.

    Endrődi S. 1976a. Monographie der Dynastinae (Col. Lamellicornia, Melolonthidae) 4. Tribus: Pentodontini der äthiopischen Region, III. Entomologische Arbeiten aus dem Museum G. Frey 27: 118–282.

    Endrődi S. 1976b. Monographie der Dynastinae (Coleoptera) 6. Tribus: Dynastini. Acta Zoologica Academiae Scientiarum Hungaricae 22: 217–269.

    Endrődi, S. 1978. Oryctoderinus walfordorum gen. et sp. n. (Coleoptera: Scarabaeidae: Dynastinae). Journal of the Australian Entomological Society 17: 159–161.

    Endrődi S. 1985. The Dynastinae of the World. Junk, Dordrecht, The Netherlands.

    Fabricius J. C. 1775. Systema Entomologiae, sistens insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Korte, Flensburgi and Lipsiae, Germany.

    Fabricius JC. 1781. Species Insectorum Exhibentes Eorum Differentias Specificas, synonyma auctorum, loca natalia, metamorphosis adiectis observationibus, descritpionibus. C.E. Bohnii, Hamburgi and Kilonii, Germany.

    Fabricius J. C. 1801. Systema Eleutheratorum. Academici Novi, Kiliae.

    Fairmaire L. 1871. Notes sur le coléoptères recueilles par Charles Coquerel à Madagascar et sur les còtes d’Afrique. 3e partie. Annales de la Société Entomologique de France (5)1: 29–78.

    Fairmaire L. 1877a. Trois nouvelles espèces de scarabéides du genre Oryctodus Boisduval. Annales de la Société Entomologique de France (5)7: 158.

    Fairmaire L. 1877b. Diagnoses de coléoptères de la Nouvelle-Bretagne. Petites Nouvelles Entomologiques 2: 185.

    Fairmaire L. 1883. Essai sur les coléoptères de l’Archipel de la Nouvelle-Bretagne. Annales de la Société Entomologique de Belgique 27: 1–58.

    Fairmaire L. 1891. Notes sur quelques coléoptères de l’Afrique intertropicale et descriptions d’espèces nouvelles. Annales de la Société Entomologique de France 60: 231–274.

    Fairmaire L. 1893. Coléoptères des Iles Comores. Annales de la Société Entomologique de Belgique 37: 521–555.

    Faithfull I. 1995. Invasion of Victoria by the African black beetle Heteronychus arator (Fabricius) (Scarabaeidae: Dynastinae) with notes on its dispersal and Australian distribution. Victorian Entomologist 25: 64–72.

    Faithfull I. 1996. Corrigendum. Victorian Entomologist 26: 65.

    Ferrar P., Standfast H. A. & Dyce A. L. 1975. A survey of blood-sucking and synanthropic Diptera and dung insects on Norfolk Island, South Pacific. Journal of the Australian Entomological Society 14: 7–13.

    Forshage M. & Krell F-T. 2016. Two exotic dynastines collected in Sweden (Coleoptera: Scarabaeidae: Dynastinae). Entomologisk Tidskrift 137: 147–149.

    French B. R. 2006. Insect Pests of Food Plants in Papua New Guinea. A Compendium. Available from URL: (Accessed 26 January 2018).

    Fuessly J. C. 1778. Etwas ueber Voets Kaeferwerk. Magazin für die Liebhaber der Entomologie 1: 1–70.

    Ganeshan S. 2000. Biological control of sugarcane pests in Mauritius: current status and future prospects. In: Sugarcane Pest Management in the New Millennium (eds PG Allsopp & W Suasa-ard) pp. 3–9. International Society of Sugar Cane Technologists, Indooroopilly, Australia.

    Ghys M. I. & Favero M. 2004. Espectro trófico de la gaviota capucho café (Larus maculipennis) en agroecosistemas del sudeste de la provincia de Buenos Aires, Argentina. Ornitologia Neotropical 5: 493–500.

    Gordh G. & Headrick D. 2011. A Dictionary of Entomology, 2nd Edition. CABI, Wallingford, United Kingdom.

    Gressit J. L. 1953. The Coconut Rhinoceros Beetle (Oryctes rhinoceros) with particular reference to the Palau Islands. Bulletin 212. Bernice P Bishop Museum, Honolulu, Hawaii.

    Gurney W. B. 1934. Records of some new insect pests. Agricultural Gazette of New South Wales 45: 442–454.

    Hangay G. & German P. 2000. Insects of Australia. Reed New Holland, Sydney, Australia.

    Hangay G. & Zborowski P. 2010. A Guide to the Beetles of Australia. CSIRO Publishing, Clayton, Australia.

    Hawkins C. N. 1942. The insects of Norfolk Island, including a preliminary report on a recent collection. Annals and Magazine of Natural History (11)9: 865–902.

    Heller K. M. 1897. Ueber Camelonotus Fairm., Horonotus Burm., Neodipelicus Rits. und Palmerstonia Blackb. Notes from the Leyden Museum 19: 163–173.

    Heller K. M. 1914. Coleoptera von Seram und Bali. (Zoologische Ergebnisse der II. Freiburger Molukken-Expedition). Entomologische Mitteilungen 3: 293–315.

    Hope F. W. 1837. The Coleopterist’s Manual, Containing the Lamellicorn Insects of Linneus [sic] and Fabricius. Volume 1. H.G. Bohn, London, United Kingdom.

    Howden H. F. 1977. Beetles, beach drift, and island biography. Biotropica 9: 53–57.

    ICZN [International Commission on Zoological Nomenclature]. 1996. Opinion 1838. Temnorhynchus Hope, 1837 (Insecta, Coleoptera): conserved. Bulletin of Zoological Nomenclature 53: 134–135.

    Illiger J. K. W. 1798. Verzeichniss der Käfer Preussens. J.J. Gebauer, Halle, Germany.

    Jack R. W. 1924. On the African species of the dynastid genus Heteronychus. Annals and Magazine of Natural History 9(11): 367–438.

    Jákl S. & Zídek J. 2016. Review of the oryctoderine genus Onychionyx Arrow, with description of a new species from the Moluccas (Coleoptera: Scarabaeidae: Dynastinae). Studies and Reports, Taxonomical Series 12: 109–116.

    Jameson M. L., Oishi D. E., Ratcliffe B. C. & McQuate G. T. 2009. Two additional invasive scarabaeoid beetles (Coleoptera: Scarabaeidae: Dynastinae) in Hawaii. Proceedings of the Hawaiian Entomological Society 41: 25–30.


    Kikkawa J., Monteith G. B. & Ingram G. 1981. Cape York Peninsula: major region of faunal interchange. In: Ecological Biogeography of Australia (ed A Keast) pp. 1697–1742. Junk, The Hague, The Netherlands.

    King P. D., Mercer C. F. & Meekings J. S. 1981a. Ecology of black beetle, Heteronychus arator (Coleoptera: Scarabaeidae)—influence of temperature on feeding, growth, and survival of the larvae. New Zealand Journal of Zoology 8: 113–117.

    King P. D, Mercer C. F. & Meekings J. S. 1981b. Ecology of black beetle, Heteronychus arator (Coleoptera: Scarabaeidae)—population modelling. New Zealand Journal of Agricultural Research 24: 99–105.

    Krajcik M. 2005. Dynastinae of the World. Checklist. Animma.x, Supplement 2: 1–122.

    Krell F-T. 1993. Phylogenetisch-systematische Revision des Genus Temnorhynchus Hope, 1837 (Coleoptera: Scarabaeoidea: Melolonthidae: Dynastinae: Pentondontini). 1. Teil: Phylogenetische Analyse, mit Ammerkungen zur phylogenetish-systematischen Methodologie. Beiträge zur Entomologie 43: 237–318.

    Krell F-T. 1994. Phylogenetisch-systematische Revision des Genus Temnorhynchus Hope, 1837 (Coleoptera: Scarabaeoidea: Melolonthidae: Dynastinae: Pentondontini). 2. Teil: Bestimmungstabelle, Katalog, Bibliographie, Gazetteer und Material-Listen. Beiträge zur Entomologie 44: 83–155.

    Krell F-T. 1995. Historical biogeography of Temnorhynchus species (Insecta: Coleoptera: Scarabaeidae: Dynastinae). Zoologischer Anzeiger 234: 209–226.

    Krell F-T. 2002. On nomenclature and synonymy of Old World Dynastinae (Coleoptera, Scarabaeidae). Entomologische Blätter 98: 37–45.

    Krell F-T. & Hangay G. 1998. The African rhinoceros beetle Temnorhynchus retusus (Fabricius) established in eastern Australia (Coleoptera: Scarabaeidae: Dynastinae). Australian Journal of Entomology 37: 312–314.

    Krell F-T. & Král D. 2017. Order Coleoptera, family Scarabaeidae. Subfamily Dynastinae. Arthropod Fauna of the UAE 6: 169–185.

    Küster H. C. 1852. Die Käfer Europas. Nach der Natur beschrieben. Mit Beiträgen mehrerer Entomologen (Apetz, Erichson, Kisenwetter). Volume 24. Bauer und Raspe, Nürnberg, Germany.

    Landin B-O. 1964. The identity of “Scarabaeus arator Fabricius 1775” (Col. Lamellicornia) with the designation of neotypes of arator Fabr. and arator Illig. and a list of the insects from the Cape Colony described by Fabricius 1775. Opuscula Entomologica 29: 117–142.

    Laporte F. L. de Castelnau. 1840. Histoire naturelle des insectes coléoptères, avec une introduction renfermant l’anatomie et la physiologie des animaux articulés, par M. Brullé. Volume 2. P. Duménil, Paris, France.

    Liebhold A. M. & Tobin P. C. 2008. Population ecology of insect invasions and their management. Annual Review of Entomology 53: 387–408.

    Linnaeus C. 1758. Systemae naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentis, synonymis, locis. Volume 1. 10th Edition, reformata. Laur. Salvii, Holmiae, Sweden.

    Lockwood J. L., Cassey P. & Blackburn T. 2005. The role of propagule pressure in explaining species invasions. Trends in Ecology and Evolution 20: 223–228.

    Lowe S. J., Brown M. & Boudjelas S. 2000. 100 of the World’s Most Invasive Species. IUCN/SSC Invasive Species Specialist Group (ISSG), Auckland, New Zealand.

    Mack R., Simberloff D., Lonsdale M., Evans H., Clout M. & Bazzaz F. 2000. Biotic invasions: causes, epidemiology, global consequences, and control. Ecological Applications 10: 689–710.[0689:BICEGC]2.0.CO;2

    Marshall S. D. G., Moore A., Vaqalo M., Noble A. & Jackson T. A. 2017. A new haplotype of the coconut rhinoceros beetle, Oryctes rhinoceros, has escaped biological control by Oryctes rhinoceros nudivirus and is invading Pacific Islands. Journal of Invertebrate Pathology 149: 127–134.

    Matthews E. G. 1984. A Guide to the Genera of Beetles of South Australia. Part 3. South Australian Museum, Adelaide, Australia.

    Matthews E. G. & Kitching R. L. 1984. Insect Ecology. 2nd edition. University of Queensland Press, St Lucia, Australia.

    Matthiessen J. N. 1999. Late immature mortality is the major influence on reproductive success of African black beetle, Heteronychus arator (Fabricius) (Coleoptera: Scarabaeidae), in a Mediterranean-climate region of Australia. Australian Journal of Entomology 38: 348–353.

    Matthiessen J. N. & Learmonth S. E. 1995. Impact of the soil insects African black beetle, Heteronychus arator (Coleoptera: Scarabaeidae) and whitefringed weevil, Graphognathus leucoloma (Coleoptera: Curculionidae), on potatoes and effects of soil insecticide treatments in south-western Australia. Bulletin of Entomological Research 85: 101–111.

    Matthiessen J. N & Learmonth S. E. 1998. Seasonally contrasting activity of African black beetle, Heteronychus arator (Coleoptera: Scarabaeidae): implications for populations, pest status and management. Bulletin of Entomological Research 88: 443–450.

    Matthiessen J. N. & Ridsdill-Smith T. J. 1990. Populations of African black beetle, Heteronychus arator (Coleoptera: Scarabaeidae) in a Mediterranean climate region of Australia. Bulletin of Entomological Research 81: 85–91.

    Mauco L. & Favero M. 2004. Diet of the common tern (Sterna hirundo) during nonbreeding season in Mar Chiquita Lagoon, Buenos Aires, Argentina. Orntiologia Neotropical 15: 121–131.

    Maynard G. V., Lepschi B. J. & Malfroy S. F. 2018. Norfolk Island quarantine survey 2012-2014—a comprehensive assessment of an isolated subtropical island. Proceedings of the Linnean Society of New South Wales 140: 7–243.

    McKinney M. L. & Lockwood J. L. 1999. Biotic homogenization: a few winners replacing many losers in the next mass extinction. Trends in Ecology and Evolution 14: 450–453.

    Minck P. 1920. Beitrag zur Kenntnis der Dynastiden. 10. Asiatische Xylotrupiden. Archiv für Naturgeschichte 82A: 194–221.

    Miyake Y. & Yamaya S. 1997. New or little known scarabaeid beetles from south east Asia preserved in Nagaoka Municipal Science Museum. Bulletin of the Nagaoka Municipal Science Museum 32: 5–20.

    Mondino E. A., López A. N., Alvarez-Castillo H. A. & Carmona D. M. 1997. Ciclo de vida de Cyclocephala signaticollis Burmeister, 1847 (Coleoptera: Scarabaeidae: Dynastinae) y su relación con los factores ambientales. Elytron 11: 145–156.

    Monteith G. B. 1974. Focus on Cape York. A report from the Fauna Subcommittee of the Entomological Society of Queensland. Entomological Society of Queensland, Brisbane, Australia.

    Monteith G. 2017. The Argentinian scarab: the new kid on the block. News Bulletin, Entomological Society of Queensland 44: 200.

    Montrouzier P. 1855. Essai sur la faune de l’Ile de Woodlark ou Moiou. Annales des Sciences Physiques et Naturelles, d’Agricultue et d’Industrie (2)7: 1–114.

    Montrouzier P. 1860. Essai sur la faune entomologique de la Nouvelle-Calédonie (balade) et des îles de Pins, Art, Lifu, etc. Annales de la Société Entomologique de France (3)8: 229–308.

    Mooney H. A. & Drake J. A. 1989. Biological invasions: a SCOPE program overview. In: Biological Invasions: a Global Perspective. SCOPE 37 (eds JA Drake, HA Mooney, F di Castri et al.) pp. 491–506. Wiley, Chichester, United Kingdom.

    Moore M. R., Cave R. D. & Branham M. A. 2018a. Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1–99.

    Moore M. R., Cave R. D. & Branham M. A. 2018b. Annotated catalog and bibliography of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae, Cyclocephalini). ZooKeys 745: 101–378.

    Moore M. R. & Jameson M. L. 2013. Floral associations of cyclocephaline scarab beetles. Journal of Insect Science 13: 1–43.

    Morelli E. 1991. Descripción de la larva y de la pupa de Cyclocephala signaticollis Burmeister, 1847 (Coleoptera: Scarabaeidae: Dynastinae) y observaciones sobre su biologia. Elytron 5 (Supplement 1): 189–195.

    New T. R. 1994. Exotic insects in Australia. Gleneagles Publishing, Adelaide, Australia.

    Nikitin M. I. 1966. On the seasonal distribution of some scarab beetles (Col., Scarabaeidae) at Cabramatta, N.S.W. Journal of the Entomological Society of Australia (N.S.W.) 3: 34–35.

    Okanjima S. & Nagai S. 2001. New Wide Gakken’s Picture Book (13). Kabuto-mushi—Kuwagata-mushi. Gakken, Tokyo, Japan (in Japanese).

    Olden J. D. 2006. Biotic homogenization: a new research agenda for conservation biogeography. Journal of Biogeography 33: 2027–2039.

    Padin S. B., Dal Bello G. M. & Vasicek A. L. 1996. Potencial bioinsecticida de hongos entomopatógenos de plagas en granos almacenados. Revista de la Facultad de Agronomia 15: 1–7.

    Pearce F. 2015. The New Wild. Why Invasive Species Will Be Nature’s Salvation. Icon Books, London, United Kingdom.

    Péringuey L. 1901. Descriptive catalogue of the Coleoptera of South Africa (Lucanidae and Scarabaeidae). Transactions of the South African Philosophical Society 12: 1–563.

    Plant Health Australia. 2019. Biosecurity. Available from URL: (Accessed 24 January 2019).

    Prins A. J. 1984. Morphological and biological notes on some South African arthropods associated with decaying organic matter. Part 3. The families Dermestidae, Cantharidae, Melyridae, Tenebrionidae, and Scarabaeidae (Coleoptera). Annals of the South African Museum 94: 203–304.

    Queensland Historic Places. 2018. Iron Range USAAF/RAAF Advanced Operational Base. Available from URL: (Accessed 4 March 2918).

    Ramírez L. C. & Alonso C. P. 2016. Two late Pleistocene members of the white-grub complex, one of the most destructive insect pests of turfgrasses. Revista Brasileira de Paleontologia 19: 531–536.

    Ratcliffe B. C., Cave R. D. & Cano E. B. 2013. The dynastine scarab beetles of Mexico, Guatemala, and Belize (Coleoptera: Scarabaeidae: Dynastinae). Bulletin of the University of Nebraska State Museum 27: 1–666.

    Reboredo G. R. & Camino N. B. 2000. Two new rhabditida species (Nematoda: Rhabditidae) parasites of Cyclocephala signaticollis (Coleoptera: Scarabaeidae) in Argentina. Journal of Parasitology 86: 819–821.

    Remedi de Gavotto A. L. 1964. Ciclo biológico de Cyclocephala signaticollis Burm. (Coleoptera, Scarabaeidae) y caracteres específicos de su larva. INTA, Revista de Investigaciones Agropecuarias 1: 151–161.

    Roets F., Allison J. D. & Basson R. J. 2019. Recent records of fruit chafers (Scarabaeidae: Cetoniinae: Cetoniini) in the southwestern Cape region of South Africa suggest that range expansions were facilitated by human-mediated jump-dispersal and pre-adaptation to transformed landscapes. African Entomology 27: 135–145.

    Rouget M. & Richardson D. M. 2003. Inferring process from patterns in plant invasions: a semimechanistic model incorporating propagule pressure and environmental factors. American Naturalist 162: 713–724.

    Rowland J. M. 2003. Male horn dimorphism, phylogeny and systematics of rhinoceros beetles of the genus Xylotrupes (Scarabaeidae, Coleoptera). Australian Journal of Zoology 51: 213–258.

    Rowland J. M. 2006a. Partial revision of the systematics of rhinoceros beetles of the genus Xylotrupes (Scarabaeidae, Coleoptera). Privately published, Albuquerque, NM, United States. 42 pp.

    Rowland J. M. 2006b. Continued revision of the systematics of rhinoceros beetles of the genus Xylotrupes (Scarabaeidae, Coleoptera). Privately published, Albuquerque, NM, United States. 50 pp.

    Rowland J. M. 2011. Notes on nomenclature in Xylotrupes Hope (Scarabaeidae: Dynastinae: Dynastini). Insecta Mundi 176: 1–10.

    Ruiz G. M. & Carlton J. T. 2003. Invasive Species. Vectors and Management Strategies. Island Press, Washington DC, USA.

    Sallam M. N., Allsopp P. G., Chandler K. J. & Samson P. R. 2007. Sugarcane. In: Pests of Field Crops and Pastures. Identification and Control (ed PT Bailey) pp. 305–341. CSIRO, Canberra, Australia.

    San Martín L. E. 1968. Control químico de los gusanos de suelo en cultivos de papas en Balcarce. IDIA (Argentina) 252: 64–72.

    Schaufuss L. W. 1885. Beitrag zur Fauna der niederländischen Besitzungen auf den Sunda-Inseln. Horae Societatis Entomologicae Rossicae 19: 183–209.

    Schoolmeesters P. 2018. Scarabs: World Scarabaeidae database (version Jul 2017). In: Species 2000 & ITIS Catalogue of Life, 20th December 2017 (eds Y Roskov, L Abucay, T Orrell et al. Available at URL: (Accessed 4 January 2018).

    Silvestre G. 1997. Les Xylotrupes de Malaise, Sumatra et Borneo (Coleoptera, Dynastidae). Coléoptères 3: 123–133.

    Silvestre G. 2003. Les Xylotrupes de Nouvelle-Guinée, d’Australie et d’Oceanie (Coleoptera, Dynastidae). Coléoptères 9: 151–162.

    Silvestre G. 2007. Les Dipelicus du groupe integriceps (Coleoptera, Dynastidae). Coléoptères 13: 73–78.

    Silvestre G. 2012. Révision des Dipelicus du groupe triangularis (Montrouzier 1855) (Coleoptera, Dynastidae). Coléoptères 18: 31–55.

    Sternberg C. 1910. Neue Dynastiden-Arten. Annales de la Société Entomologique de Belgique 54: 91–102.

    Szent-Ivany J. J. H. 1969. The distribution, systematics and feeding habits of the elephant beetle, Xylotrupes gideon (L.) in New Guinea and in Solomon Islands. Transactions of the Papua New Guinea Scientific Society 10: 14–18.

    Taylor C. M. & Hastings A. 2005. Allee effects in biological invasions. Ecology Letters 8: 895–908.

    Todd D. H. 1959. Black beetle Heteronychus sanctaehelenae Blanch., in pastures in New Zealand. New Zealand Journal of Agricultural Research 2: 1262–1273.

    Tsatsia H. & Jackson G. 2017. Taro Papuana beetle. Pacific Pests and Pathogens - Fact Sheets. Available at URL: (Accessed 1 February 2018).

    Valentine E. W. 1979. The parasites and predators of Heteronychus arator in South Africa (Coleoptera: Scarabaeidae). Proceedings of the II Australasian Conference on Grassland Invertebrate Ecology 216–219.

    Verdugo A. & Drumont A. 2015. Révision du genre Calicnemis Laporte, 1832: approches morphologique et génétique (Coleoptera, Scarabaeidae, Dynastinae). Revue de l’Association Roussillonnaise d’Entomologie 24 (Supplement): 1–64.

    Vinson J. 1963. Sur l’apparition a l’Maurice d’un scarabee nuisible, Oryctes rhinoceros Linne. Revue Agricole et Sucrière de l’Ile Maurice 42: 173–80.

    Waterhouse D. F. & Norris K. R. 1987. Biological Control. Pacific Prospects. Inkata Press, Melbourne, Australia.

    Weir T. A., Lawrence J. F., Lemann C. & Gunter N. L. 2019. 33. Scarabaeidae: Dynastinae Macleay 1919 [sic]. In: Australian Beetles. Volume 2. Archostemata, Myxophaga, Adephaga, Polyphaga (part) (eds A Ślipiński & JF Lawrence) pp. 516–530. CSIRO, Clayton, Australia.

    Wollaston T. V. 1869. On the Coleoptera of St. Helena. Annals and Magazine of Natural History (4)4: 297–321.

    Wood J. F. 1909. The fauna of the Cocos-Keeling Atoll. Proceedings of the Zoological Society of London 1909: 132–160.