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

Abstract

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.