Abstract. A hypothesis has been advanced recently predicting that, in evolution, as higher-level entities arise from associations of lower-level organisms, and as these entities acquire the ability to feed, reproduce, defend themselves, and so on, the lower-level organisms will tend to lose much of their internal complexity (McShea 2001a). In other words, in hierarchical transitions, there is a drain on numbers of part types at the lower level. One possible rationale is that the transfer of functional demands to the higher level renders many part types at the lower level useless, and thus their loss in evolution is favored by selection for economy. Here, a test is conducted at the cell level, comparing numbers of part types in free-living eukaryotic cells (protists) and the cells of metazoans and land plants. Differences are significant and consistent with the hypothesis, suggesting that tests at other hierarchical levels may be worthwhile. Hierarchical transitions occur when a group of organisms combine to form a higher-level entity (Spencer 1904;Campbell 1958;Simon 1962;Pattee 1970;Eldredge and Salthe 1984;Salthe 1985Salthe , 1993Buss 1987;Bonner 1988;Maynard Smith 1988;Wimsatt 1994;Maynard Smith and Szathmáry 1995;McShea 1996 McShea , 2001bPettersson 1996;Valentine and May 1996;Michod 1999). In the history of life, salient examples include the origin of the eukaryotic cell from symbiotic associations of prokaryotic cells, of multicellular individuals from clones of free-living eukaryotic cells, and of colonies from associations of multicellular individuals. A hypothesis has been advanced recently that predicts that, as such transitions occur, and as the new higher-level entities acquire the ability to feed, reproduce, defend themselves, and so on, their component organisms will tend to lose many of these functional capabilities and therefore will tend to lose many of their internal parts (McShea 2001a). More precisely, the expectation is that the component organisms will tend to lose part types at the next hierarchical level down (Fig. 1). (Technical understandings of the terms part and level will be offered shortly; for present purposes, colloquial meanings suffice.)Concretely, the hypothesis predicts that the cells of multicellular organisms will have fewer part types than freeliving eukaryotic cells, that is, nonparasitic protists (hereafter simply protists; sensu Patterson 1999). At a higher level, in colonial marine invertebrates and social insects, the multicellular individuals (e.g., polyps, zooids) that constitute the more individuated colonies-that is, those in which coloniality is more highly developed (Boardman and Cheetham 1973)-should have fewer part types, on average, than those of their less colonial or solitary relatives (Beklemishev 1969). And at a lower level, the former prokaryotes that today function as organelles in eukaryotic cells (e.g., mitochondria, plastids) should have fewer part types than free-living prokaryotes.Attention to hierarchical level is crucial. A considerable body of e...