Systematic affinities of the lyrebirds (Passeriformes: Menura), with a novel classification of the major groups of passerine birds Per G.P. Ericson, a, * Les Christidis, b Martin Irestedt, a,c and Janette A. Norman b a Department of Vertebrate Zoology and Molecular Systematics Laboratory, Swedish Museum of Natural History, P.O. Box 50007, SE-10405 Stockholm, Sweden b Sciences Department, Museum Victoria, P.O. Box 666E, Melbourne, Vic. 3001, Australia c Department of Zoology, University of Stockholm, SE-106 91 Stockholm, Sweden Received 3 October 2001; received in revised form 19 December 2001 Abstract Phylogenetic relationships of the lyrebirds are investigated using DNA sequence data. The aligned data matrix consists of 4027 bp obtained from three nuclear genes (c-myc, RAG-1 and myoglobin intron II) and two mitochondrial genes (cytochrome b and ND2). Both maximum-likelihood and parsimony analyses show that the lyrebirds unambiguously belong to the oscine radiation, and that they are the sister taxon to all other oscines. The results do not support the suggestion based on DNA–DNA hybridization data (Sibley and Ahlquist, 1990) that the treecreepers and bowerbirds are part of the lyrebird clade. Nevertheless, treecreepers and bowerbirds are sister taxa to all other oscines (except the lyrebirds) and may constitute a monophyletic group, although bootstrap support values for this clade are low. A major disagreement between the present analysis and that based on DNA–DNA hybrid- ization data is that the Corvida (sensu Sibley and Ahlquist, 1990) and Passerida are not reciprocally monophyletic, as we find the latter group be nested within the Corvida. Also, the superfamilies Meliphagoidea and Corvoidea sensu Sibley and Ahlquist (1990), are not recovered as monophyletic in the present study. Within the oscine radiation, all taxa belonging to the earliest splits are confined to the Australo–Papuan region. This suggests strongly that the origins and early radiation of the oscines occurred in the southern supercontinent Gondwana. A new classification of the major groups of passerines is presented following from the results presented in the present study, as well as those published recently on analyses of sequence data from the nuclear c-myc and RAG-1 genes (Ericson et al., 2002; Irestedt et al., 2001). Ó 2002 Elsevier Science (USA). All rights reserved. 1. Introduction The order Passeriformes comprises more than half of all living species of birds. Both the monophyly of the order and its further split into two major clades, the Suboscines and Oscines, are well supported by mor- phological characters (Raikow, 1982), and molecular data (Edwards et al., 1991; Irestedt et al., 2001; Sibley and Ahlquist, 1990). One passerine family, the Acanth- isittidae (New Zealand wrens), falls outside the subos- cine and oscine clades, and thus constitutes the sistertaxon to the other passerines (Ericson et al., 2002). The classification of major groups of oscine birds has long been a contentious issue among avian systematists, most recently reviewed by Voous (1985) and Sibley and Ahlquist (1990). DNA–DNA hybridization data suggest that the oscines consists of two major groups; the Corvida and the Passerida (Harshman, 1994; Sheldon and Gill, 1996; Sibley and Ahlquist, 1990), and mor- phological support for this comes from the presence of a fully developed double pneumatic fossae in the proximal end of the humerus in many oscines, but not in the taxa placed in the Corvida (Bock, 1962). One family of birds that long has been regarded as especially difficult to place phylogenetically is the lyre- birds (Menuridae). Lyrebirds are large, terrestrial Aus- tralian passerines that are particularly renowned for their loud, elaborate songs and spectacular ability to mimic other sounds, both natural and man-made. The two extant species included in the family, the Superb Lyrebird (Menura novaehollandiae) and Albert’s Lyre- bird (M. alberti), are restricted to the forests of south- Molecular Phylogenetics and Evolution 25 (2002) 53–62 MOLECULAR PHYLOGENETICS AND EVOLUTION www.academicpress.com * Corresponding author. Fax: +46-8-51954212. E-mail address: per.ericson@nrm.se (P.G.P. Ericson). 1055-7903/02/$ - see front matter Ó 2002 Elsevier Science (USA). All rights reserved. PII:S1055-7903(02)00215-4