Abstract. There are over 200 species of nematine sawflies that induce galls on willows (Salix spp.). Most of the species are mono-or oligophagous, and they can be separated into seven or eight different groups based on the type of gall that they induce. We studied the evolution of different gall types and host plant associations by reconstructing the phylogeny of five outgroup and 31 ingroup species using DNA sequence data from the mitochondrial cytochrome b gene. Maximum-parsimony and maximum-likelihood analyses resulted in essentially the same phylogeny with high support for important branches. The results show that: (1) the galling species probably form a monophyletic group; (2) true closed galls evolved only once, via leaf folders; (3) with the possible exception of leaf rollers, all gall type groups are mono-or paraphyletic; (4) similar gall types are closer on the phylogeny than would be expected by a random process; (5) there is an apparent evolutionary trend in galling site from the leaf edge towards the more central parts of the host plant; and (6) many willow species have been colonized several times, which excludes the possiblity of parallel cladogenesis between willows and the gallers; however, there are signs of restrictions in the evolution of host use. Many of the patterns in the evolutionary history of nematine gallers have also been observed in earlier studies on other insect gallers, indicating convergent evolution between the independent radiations.Key words. Coevolution, convergent evolution, evolutionary trends, extended phenotypes, gall morphology, gradualism, insect-plant relationships, Nematinae, phylogeny.Received June 1, 1999. Accepted October 11, 1999.The habit of galling has evolved many times independently in various organisms (Meyer 1987). In insects alone, gall induction has evolved in at least seven different orders, and in most of them several radiations have arisen (Meyer 1987; Dreger-Jauffret and Shorthouse 1992). The multiple origins of galling makes it possible to search for convergent features in these independent evolutionary histories. Indeed, recent phylogenetic analyses of gall-inducing aphids (Stern 1995), wasps (Stone and Cook 1998), and thrips (Crespi and Worobey 1998) have shown that common features exist in separate radiations. For example, in all these taxa gall morphology is determined mainly by the galler, not the host plant, and thus the gall can be considered an extended phenotype (sensu Dawkins 1982) of the galler.The nematine sawflies that induce galls on willows (Salix spp.) also offer a good model system for the study of evolutionary questions, especially questions about the evolution of different gall types and host plant associations (Price 1992). These sawflies belong to the tenthredinid subfamily Nematinae, which is comprised of hundreds of galling and nongalling species (Smith 1979;Gauld and Bolton 1988). There are about 200 species of galling nematines, most of which are mono-or oligophagous (Smith 1970;Price et al. 1994). Like their host plants, thes...