Acta Zoologica Sinica © 2006 Acta Zoologica Sinica S20-1 Supra-generic cladistics of landfowl (Order Galliformes) Timothy M. CROWE 1,2* , Paulette BLOOMER 1,3 , Ettore RANDI 4 , Vittorio LUCCHINI 4 , Rebecca KIMBALL 5 , Edward BRAUN 6 , Jeffrey G. GROTH 2 1. Percy FitzPatrick Institute, University of Cape Town, Rondebosch 7701, South Africa; * tmcrowe@botzoo.uct.ac.za 2. Dept. of Ornithology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024- 5192, USA 3. Dept. of Genetics, University of Pretoria, Pretoria 0001, South Africa 4. Istituto Nazionale per la Fauna Selvatica (INFS), via Cà Fornacetta 9, 40064 Ozzano dell’Emilia (BO), Italy 5. Dept. of Zoology, University of Florida, Gainesville, FL 32611, USA 6. Dept. of Biology, University of New Mexico, Albuquerque, New Mexico 87131, USA Abstract Molecular and organismal evidence bearing on the evolutionary relationships of gamebirds within the order Galliformes is reviewed. The monophyly of most traditionally recognized supra-generic taxa is supported, but the Phasianini (pheasants) and Perdicini (partridges and quails) are shown to be polyphyletic. Contrary to DNA-DNA hybridization data, New World quails (Odontophoridae) did not branch off the galliform evolutionary tree before the guineafowl (Numididae), but rather immediately after; and the cracids (Cracidae) and megapodes (Megapodiidae) are not sister groups of one another. Evidence is presented to support the notion of very early divergence among species currently confined to the New World, Africa and southeast Asia. Evidence from mtDNA sequences (cytochrome b and D-loop), proteins (ovomucoids), life history, behavior, and bones and feathers is analyzed phylogenetically in three ways: separately, combined and combined but differen- tially weighted. Separate analysis always produces less well-resolved trees than those suggested by combined data; and deweighting putatively less informative evidence undermines rather than enhances phylogenetic signal. Furthermore, analysis of combined data produces a tree with a novel, but biogeographically meaningful, topology, with organismal and molecular information more useful at respective more basal and more terminal branches of the tree. Key words Galliformes, Phylogeny, Combined vs separate vs deweighted evidence 1 Introduction The phylogeny of the pheasant, quail and turkey or- der Galliformes is reconstructed for 102 in-group taxa listed in Crowe et al. (1992, Appendix 3) and rooted on two repre- sentatives of the Anseriformes, a duck (Cairina moschata) and a screamer (Chauna torquata). Exemplars represent all putative supra-generic taxa and 55 traditionally recognized genera (after del Hoyo et al., 1994). Character data include a combination of 949 phylogenetical informants, from mor- phology and behavior (n = 47), ovomucoid amino acids (n = 47), and gene sequences in mitochondrial cytochrome b and D-loop (n = 522, 333 respectively). Raw data matrices for these three sources of character variation are available from the senior author or from the web site of the Percy FitzPatrick Institute. 2 Methods Phylogenetic analyses were conducted using WinClada ver. 0.9.99m8.1 BETA (Nixon, 2000). The search- ing strategy employed was the Ratchet Island Hopper (1 000 iterations; 2 trees to hold; ±10% characters to sample; and amb. poly. = option), with organismal characters as additive and all molecular characters nonadditive. Bootstrap analyses were conducted with 1 000 replicates. 3 Results and discussion Analysis of the combined dataset of characters yielded two equally parsimonious trees (CI = 0.21, 7 296 steps), of which a strict consensus application required the collapsing of five nodes, all within the tetraonine grouse (Fig. 1). Traditionally recognized supra-generic taxa sup- ported by this consensus tree with ≥ 50% bootstrap sup- port are: (1) the Superfamily Phasianoidea, including all galliforms except the megapodes and cracids; (2) the megapodes (Megapodiidae); (3) the cracids (Cracidae); (4) the guineafowls (Numididae); and (4) the New World quails (Odontophoridae). Resolution within the Phasianidae is not well supported. Within the phasianids, the grouse (Tetraoninae), the gallo-pheasant subset of the pheasants (Phasianinae, including Syrmaticus , Phasianus , Chrysolophus, Lophura, Catreus and Crossoptilion), and the pavonine peafowl (Pavonini, including Rheinardia, Argusianus, Afropavo and Pavo) are supported. However, although grouping with the gallo-pheasants, the basal pheasant assemblage of Johnsgard (1986), excluding the junglefowls ( Gallus spp.) and peacock-pheasants (Polyplectron spp.), does not have “significant” (≥ 50%) 52(Supplement): 358–361, 2006