Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

Relationships among felt scale insects (Hemiptera : Coccoidea : Eriococcidae) of southern beech, Nothofagus (Nothofagaceae), with the first descriptions of Australian species of the Nothofagus-feeding genus Madarococcus Hoy

Nate B. Hardy A E , Penny J. Gullan A , Rosa C. Henderson B and Lyn G. Cook C D
+ Author Affiliations
- Author Affiliations

A Department of Entomology, University of California, One Shields Avenue, Davis, California 95616-8584, USA.

B Landcare Research, Private Bag 92170, Auckland, New Zealand.

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

D Present address: School of Integrative Biology, The University of Queensland, Brisbane 4072, Australia.

E Corresponding author. Email:

Invertebrate Systematics 22(3) 365-405
Submitted: 11 July 2007  Accepted: 28 February 2008   Published: 18 June 2008


Species of southern beech (Nothofagus) have been studied extensively because of their importance in understanding southern hemisphere biogeography. Nothofagus species support a diverse assemblage of insect herbivores, including more than 30 described species of felt scales (Eriococcidae). We reconstructed the phylogeny of the Nothofagus-feeding felt scales with nucleotide sequence data and morphology. All but one of the exclusively Nothofagus-feeding species included in the analyses were recovered as a monophyletic group. This clade comprised the genera Chilechiton Hodgson & Miller, Chilecoccus Miller & González, Intecticoccus Kondo, Madarococcus Hoy (except for M. totorae Hoy), Sisyrococcus Hoy and several species of the genus Eriococcus Targioni Tozzetti. The genera Eriococcus and Madarococcus were not recovered as monophyletic. Here we revise Madarococcus. We expand the concept of the genus, provide a key to the adult females of the 31 species of Madarococcus and, for each named species, provide revised synonymies and any new collection or taxonomic information. We recognise the genus from Australia for the first time and describe the adult females of six new Australian species: Madarococcus cunninghamii Hardy & Gullan, sp. nov.; M. meander Hardy & Gullan, sp. nov.; M. megaventris Hardy & Gullan, sp. nov.; M. moorei Hardy & Gullan, sp. nov.; M. occultus Hardy & Gullan, sp. nov., and M. osculus Hardy & Gullan, sp. nov. We also describe the first-instar nymphs of M. cunninghamii, sp. nov., M. meander, sp. nov. and M. moorei, sp. nov. We transfer 17 species into Madarococcus from Eriococcus: M. argentifagi (Hoy), comb. nov.; M. cavellii (Maskell), comb. nov.; M. chilensis (Miller & González), comb. nov.; M. detectus (Hoy), comb. nov.; M. eurythrix (Miller & González), comb. nov.; M. fagicorticis (Maskell), comb. nov.; M. hispidus (Hoy), comb. nov.; M. latilobatus (Hoy), comb. nov.; M. maskelli, (Hoy), comb. nov.; M. montifagi (Hoy), comb. nov.; M. navarinoensis (Miller & González), comb. nov.; M. nelsonensis (Hoy), comb. nov.; M. nothofagi (Hoy), comb. nov.; M. podocarpi (Hoy), comb. nov.; M. raithbyi (Maskell), comb. nov.; M. rotundus (Hoy), comb. nov. and M. rubrifagi (Hoy), comb. nov. We transfer two species from Sisyrococcus into Madarococcus: M. intermedius (Maskell), comb. nov. and M. papillosus (Hoy), comb. nov. One species, M. totarae (Maskell), is excluded from Madarococcus, but cannot at present be placed in another genus and is listed as ‘M.’ totarae incertae sedis. We report the first collection of an eriococcid, M. osculus, sp. nov., on the deciduous beech, Nothofagus gunnii. With respect to biogeography, the results of our phylogenetic analysis are congruent with those obtained from recent analysis of Nothofagus; Australian and New Zealand species of Madarococcus form a monophyletic group to the exclusion of the South American species, suggesting that long-distance dispersal has played an important role in shaping the distributions of both the Nothofagus-feeding felt scales and their hosts.


We gratefully acknowledge the contributions to Australian entomology made by the late Jack Beardsley, who collected specimens of each of the six Australian species described in this paper. We thank Cathy Young (Department of Primary Industries and Water, Tasmania) and Jane Keble-Williams (University of Tasmania) for the loan of Tasmanian eriococcids from Nothofagus. Dug Miller kindly provided information on the USNM holdings of Maskell specimens and arranged the loan of some type specimens. Jon Martin arranged the loan of and provided some information on BMNH specimens of Madarococcus. David Morris kindly provided sequence data for one collection. Many people assisted this research by collecting specimens. We especially thank Nicholas Martin who provided many eriococcid specimens to NZAC, Mike Crisp and David Morris for collecting galls on Nothofagus in Australia, and Demian Kondo for collecting the Chilean specimens used for the DNA work. We acknowledge the Chilean National Forests Corporation (CONAF) for collecting permits (held by PJG in 1996, E. Arias in 2003, T. Kondo in 2006). The Australian National Botanic Gardens (ANBG), Canberra, allowed PJG to collect scale insects and Stuart Donaldson (ANBG) provided assistance. Peter Cranston accompanied PJG on many field trips to collect Nothofagus-feeding eriococcids and Robert Hoare assisted PJG with fieldwork in New Zealand. Graeme Henderson accompanied RCH on field trips to collect scale insects in New Zealand. The following agencies/departments also provided collecting permits to PJG, LGC, RCH and their collaborators: the former Department of Parks, Wildlife and Heritage, and the current Department of Primary Industries and Water, Tasmania; New South Wales National Parks and Wildlife Service; State Forests of New South Wales; and the Department of Conservation, New Zealand. This work was supported by an Australian Biological Resources Study grant (1995–1998) to PJG; a National Science Foundation PEET grant (DEB-0118718) and by Hatch funding from the California Agricultural Experiment Station to PJG; the Foundation for Research, Science and Technology, New Zealand, under contract CO9X0501 to RCH; and an ARC Discovery Project grant to LGC.


Belshaw R., Quicke D. L. J. (1997) A molecular phylogeny of the Aphidiinae (Hymenoptera: Braconidae). Molecular Phylogenetics and Evolution 7, 281–293.
CrossRef | PubMed |

Ben-Dov Y., Miller D. R., and Gibson G. A. P. (2007). ‘ScaleNet, Scales on a Specified Host Query Results.’ Available at [Verified 25 June 2007.]

Brower A. V. Z., DeSalle R. (1998) Patterns of mitochondrial versus nuclear DNA sequence divergence among nymphalid butterflies: the utility of wingless as a source of characters for phylogenetic inference. Insect Molecular Biology 7, 73–82.
CrossRef | PubMed |

Cho S., Mitchell A., Regier J. C., Mitter C., Poole R. W., Friedlander T. P., Zhao S. (1995) A highly conserved nuclear gene for low-level phylogenetics: elongation-factor 1α recovers morphology-based tree for heliothine moths. Molecular Biology and Evolution 12, 650–656.
PubMed |

Cook L. G., Crisp M. D. (2005) Not so ancient: the extant crown group of Nothofagus represents a post-Gondwanan radiation. Proceedings of the Royal Society of London. Series B. Biological Sciences 272, 2535–2544.
CrossRef |

Cook L. G., Gullan P. J. (2004) The gall-inducing habit has evolved multiple times among the eriococcid scale insects (Sternorrhyncha: Coccoidea: Eriococcidae). Biological Journal of the Linnean Society 83, 441–452.
CrossRef |

Cook L. G., Gullan P. J., Trueman H. E. (2002) A preliminary phylogeny of the scale insects (Hemiptera: Sternorrhyncha: Coccoidea) based on nuclear small-subunit ribosomal DNA. Molecular Phylogenetics and Evolution 25, 43–52.
CrossRef | PubMed |

Crosby T. K., Dugdale J. S., Watt J. C. (1998) Area codes for recording specimen localities in the New Zealand subregion. New Zealand Journal of Zoology 25, 175–183.

Deitz L. L., and Tocker M. F. (1980). W.M. Maskell’s Homoptera: species-group names and type material. New Zealand Department of Scientific and Industrial Research. Information Series 146. pp. 1–76.

Fernald M. E. (1903) A catalogue of the Coccidae of the world. Bulletin of the Hatch Experiment Station of the Massachusetts Agricultural College 88, 1–360.

Gibbs R. A., and Cockerill M. (1995). ‘Sequencher. Version 3.0.’ (Gene Codes Corporation: Ann Arbor, MI, USA.)

Gonzalez P. (2004) Eriococcidos (Hemiptera: Coccoidea) encontrados en la Patagonia, Argentina. Revista de la Sociedad Entomológica Argentina 63, 31–32.

Govaerts R., and Frodin D. (1998). ‘World checklist and bibliography of Fagales (Betulaceae, Corylaceae, Fagaceae and Ticodendraceae).’ (The Royal Botanic Gardens: Kew, UK.)

Granara de Willink M. C., Miller D. R. (2004) Two new species of mealybugs (Hemiptera: Coccoidea: Pseudococcidae) from Patagonia, Argentina. Proceedings of the Entomological Society of Washington 106, 140–158.

Gullan P. J. (1984) A revision of the gall-forming coccoid genus Apiomorpha Rübsaaman (Homoptera: Eriococcidae: Apiomorphinae). Australian Journal of Zoology. Supplementary Series 97, 1–203.

Heads M. (2006) Panbiogeography of Nothofagus (Nothofagaceae): analysis of the main species massings. Journal of Biogeography 33, 1066–1075.
CrossRef |

Henderson R. C., Martin N. A. (2006) Review of the gall-inducing scale insects of New Zealand (Hemiptera: Coccoidea), with a guide to field identification. New Zealand Entomologist 29, 59–75.

Hill R. S. (2001) Biogeography, evolution and palaeoecology of Nothofagus (Nothofagaceae): the contribution of the fossil record. Australian Journal of Botany 49, 321–332.
CrossRef |

Hodgson C. J., Miller D. R. (2002) A new genus and two new species of felt scales (Hemiptera: Coccoidea: Eriococcidae) from Chile, with comments on zoogeographical affinities between the eriococcid faunas of southern South America and New Zealand. Systematic Entomology 27, 191–209.
CrossRef |

Hoy J. M. (1962a) Eriococcidae (Homoptera: Coccoidea) of New Zealand. New Zealand Department of Scientific and Industrial Research Bulletin 146, 1–219.

Hoy J. M. (1962b) A new species of Eriococcus (Homoptera: Coccoidea) from Navarino Island, Tierra del Fuego. New Zealand Journal of Science 5, 510–512.

Huelsenbeck J. P. (1991) When are fossils better than extant taxa in phylogenetic analysis? Systematic Zoology 40, 458–469.
CrossRef |

Humphries C. J. (1981). Biogeographical methods amd the southern beeches (Fagaceae: Nothofagus). In ‘Advances in Cladistics: Proceedings of the First Meeting of the Willi Hennig Society’. (Eds V. A. Funk and D. R. Brooks.) pp. 177–207. (The New York Botancial Garden: New York, USA.)

Humphries C. J., Cox J. M., and Nielsen E. S. (1986). Nothofagus and its parasites: a cladistic approach to coevolution. In ‘Coevolution and Systematics’. (Eds A. R. Stone and D. L. Hawksworth.) pp. 55–76. (Claredon Press: Oxford, UK.)

Knapp M., Stöckler K., Havell D., Delsuc F., Sebastiani F., Lockhart P. J. (2005) Relaxed molecular clock provides evidence for long-distance dispersal of Nothofagus (southern beech). PLoS Biology 3, 38–43.
CrossRef |

Kondo T., Hardy N., Cook L., Gullan P. (2006) Descriptions of two new genera and species of Eriococcidae (Hemiptera: Coccoidea) from southern South America. Zootaxa 1349, 19–36.

Lewis P. O. (2001) A likelihood approach to estimating phylogeny from discrete morphological character data. Systematic Biology 50, 913–925.
CrossRef | PubMed |

Li R. Q., Chen Z. D., Lu A. M., Soltis D. E., Soltis P. S., Manos P. S. (2004) Phylogenetic relationships in Fagales based on DNA sequences from three genomes. International Journal of Plant Sciences 165, 311–324.
CrossRef |

Linder H. P., Crisp M. D. (1995) Nothofagus and Pacific Biogeography. Cladistics 11, 5–32.
CrossRef |

Lindinger L. (1933) Beiträge zur Kenntnis der Schildläuse (Hemipt. - Homopt., Coccid.). Entomologischer Anzeiger [In German] 13, 77–166.

Manos P. S. (1997) Systematics of Nothofagus (Nothofagaceae) based on rDNA spacer sequences (ITS): taxonomic congruence with morphology and plastid sequences. American Journal of Botany 84, 1137–1155.
CrossRef |

Manos P. S., Steele K. P. (1997) Phylogenetic analyses of higher Hamamelidae based on plastid sequence data. American Journal of Botany 84, 1407–1419.
CrossRef |

Manos P. S., Soltis P. S., Soltis D. E., Manchester S. R., Oh S. H., Bell C. D., Dilcher D. L., Stone D. E. (2007) Phylogeny of extant and fossil Juglandaceae inferred from the integration of molecular and morphological datasets. Systematic Biology 56, 412–430.
CrossRef | PubMed |

Maskell W. M. (1890). Further notes on Coccidae, with descriptions of new species from Australia, Fiji, and New Zealand. Transactions and Proceedings of the New Zealand Institute 22, 133–156.

Maskell W. M. (1891). Further coccid notes: with descriptions of new species from New Zealand, Australia, and Fiji. Transactions and Proceedings of the New Zealand Institute 23, 1–36.

Maskell W. M. (1892). Further coccid notes: with descriptions of new species, and remarks on coccids from New Zealand, Australia and elsewhere. Transactions and Proceedings of the New Zealand Institute 24, 1–64.

McQuillan P. B. (1993) Nothofagus (Fagaceae) and its invertebrate fauna – an overview and preliminary synthesis. Biological Journal of the Linnean Society 49, 317–354.

Miller D. R., Gimpel M. E. (1996) Nomenclatural changes in the Eriococcidae (Homoptera: Coccoidea). Proceedings of the Entomological Society of Washington 98, 597–606.

Miller D. R., and Gimpel M. E. (2000). ‘A Systematic Catalogue of the Eriococcidae (Felt Scales) (Hemiptera: Coccoidea) of the World.’ (Intercept Ltd: Andover, UK.)

Miller D. R., and Gimpel M. E. (2007). ‘ScaleNet, Madarococcus totarae.’ Available at [Verified 6 July 2007]

Miller D. R., González R. H. (1975) A taxonomic analysis of the Eriococcidae of Chile. Revista Chilena de Entomologia 9, 131–163.

Miller D. R., McKenzie H. L. (1967) A systematic study of Ovaticoccus Kloet and its relatives, with a key to North American genera of Eriococcidae (Homoptera: Coccoidea: Eriococcidae). Hilgardia 38, 471–539.

Posada D., Crandall K. A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics (Oxford England) 14, 817–818.
CrossRef |

Rambaut A. (1998). ‘Se-Al (Sequence Alignment Editor Version 1.0 alpha 1).’ Available at [Verified 12 May 2008.]

Ronquist F., Huelsenbeck J. P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19, 1572–1574.
CrossRef | PubMed |

SanMartín I., Ronquist F. (2004) Southern Hemisphere biogeography inferred by event-based models: plant versus animal patterns. Systematic Biology 53, 216–243.
CrossRef | PubMed |

Springer M. S., Teeling E. C., Madsen O., Stanhope M. J., de Jong W. W. (2001) Integrated fossil and molecular data reconstruct bat echolocation. Proceedings of the National Academy of Sciences of the United States of America 98, 6241–6246.
CrossRef | PubMed |

Sunnucks P., Hales D. F. (1996) Numerous Transposed sequences of mitochondrial chytochrome oxidase I–II in aphids of the genus Sitobion (Hemiptera: Aphididae). Molecular Biology and Evolution 17, 510–524..

Swofford D. L. (2003). ‘PAUP*. Phylogenetic Analysis Using Parsimony (* and Other Methods) Version 4.’ (Sinauer Associates: Sunderland, MA, USA.)

Tautz D., Hancock J. M., Webb D. A., Tautz C., Dover G. A. (1988) Complete sequences of the rRNA genes of Drosophila melanogaster. Molecular Biology and Evolution 5, 366–376.
PubMed |

Torre-Bueno J. la  (1989). ‘The Torre-Bueno Glossary of Entomology.’ Compiled by S. W. Nichols with Supplement A by G. S. Tulloch. (New York Entomological Society: New York, USA.)

von Dohlen C. D., Moran N. A. (1995) Molecular phylogeny of the Homoptera: a paraphyletic taxon. Journal of Molecular Evolution 41, 211–223.
CrossRef | PubMed |

Wiens J. J. (2003) Missing data, incomplete taxa, and phylogenetic accuracy Systematic Biology 52, 528–538.
CrossRef | PubMed |

Williams D. J. (1985a) Some South American scale insects (Homoptera: Coccoidea) on Nothofagus. Journal of Natural History 19, 249–258.
CrossRef |

Williams D. J. (1985b) The British and some other European Eriococcidae (Homoptera: Coccoidea). Bulletin of the British Museum Natural History [Entomology] 51, 347–393.

Williams D. J. (2007) Scale insects of the families Asterolecaniidae and Eriococcidae (Hemiptera: Coccoidea) in New Caledonia. Journal of Natural History 41, 1343–1366.
CrossRef |

Appendix 1.  Collection data, museum depositories and GenBank accession numbers for specimens of species included in the molecular and morphological phylogenetic analyses
The specimen codes allow nucleotide sequences to be matched to slide-mounted voucher specimens in the depositories (see ‘Materials and Methods’ for names of depositories). In the ‘Host’ column, the first cell for each Madarococcus species lists all known host species; full host ranges are not given for outgroups. The ‘Morphology’ column specifies which specimens were used for the phylogenetic analysis of morphological data; c, crawler; e, embryo; f, adult female. Abreviations: Ck, Creek; NP, National Park; PN, Parque Nacional; Ra, Range; RN, Reserva Nacional; SF, State Forest
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Appendix 1a. (continued)
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Appendix 1b. (continued)
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Appendix 1c. (continued)
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