'Species' without species.

Biological science uses multiple species concepts. Order can be brought to this diversity if we recognize two key features. First, any given species concept is likely to have a patchwork structure, generated by repeated application of the concept to new domains. We illustrate this by showing how two species concepts (biological and ecological) have been modified from their initial eukaryotic applications to apply to prokaryotes. Second, both within and between patches, distinct species concepts may interact and hybridize. We thus defend a semantic picture of the species concept as a collection of interacting patchwork structures. Thus, although not all uses of the term pick out the same kind of unit in nature, the diversity of uses reflects something more than mere polysemy. We suggest that the emphasis on the use of species to pick out natural units is itself problematic, because that is not the term's sole function. In particular, species concepts are used to manage inquiry into processes of speciation, even when these processes do not produce clearly delimited species.

Neutral evolution of cellular phenotypes.

Eukaryotes exhibit a great diversity of cellular and subcellular morphologies, but their basic underlying architecture is fairly constant. All have a nucleus, Golgi, cytoskeleton, plasma membrane, vesicles, ribosomes, and all known lineages but one have mitochondrion-related organelles. Moreover, most eukaryotes undergo processes such as mitosis, meiosis, DNA recombination, and often perform feats such as phagocytosis, and amoeboid and flagellar movement. With all of these commonalities, it is obvious that eukaryotes evolved from a common ancestor, but it is not obvious how eukaryotes came to have their diverse structural phenotypes. Are these phenotypes adaptations to particular niches, their evolution dominated by positive natural selection? Or is eukaryotic cellular diversity substantially the product of neutral evolutionary processes, with adaptation either illusory or a secondary consequence? In this paper, we outline how a hierarchical view of phenotype can be used to articulate a neutral theory of phenotypic evolution, involving processes such as gene loss, gene replacement by homologues or analogues, gene duplication followed by subfunctionalization, and constructive neutral evolution. We suggest that neutral iterations of these processes followed by entrenchment of their products can explain much of the diversity of cellular, developmental, and biochemical phenotypes of unicellular eukaryotes and should be explored in addition to adaptive explanations.

A Reappraisal of Charles Darwin's Engagement with the Work of William Sharp Macleay.

Charles Darwin, in his species notebooks, engaged seriously with the quinarian system of William Sharp Macleay. Much of the attention given to this engagement has focused on Darwin's attempt to explain, in a transmutationist framework, the intricate patterns that characterized the quinarian system. Here, I show that Darwin's attempt to explain these quinarian patterns primarily occurred before he had read any work by Macleay. By the time Darwin began reading Macleay's writings, he had already arrived at a skeptical view of the reality of these patterns. What most interested Darwin, as he read Macleay, was not the quinarian system itself. Rather, Darwin's notes on his reading primarily concerned certain background principles animating Macleay's work, in particular: (a) the non-existence of a saltus between human and animal minds, (b) the difficulty of establishing boundaries between species and varieties, and (c) Macleay's method of variation. Darwin's interest in the last of these left a mark on his discussion of taxonomic methodology in the Origin.

On the Origins of the Quinarian System of Classification.

William Sharp Macleay developed the quinarian system of classification in his Horæ Entomologicæ, published in two parts in 1819 and 1821. For two decades, the quinarian system was widely discussed in Britain and influenced such naturalists as Charles Darwin, Richard Owen, and Thomas Huxley. This paper offers the first detailed account of Macleay's development of the quinarian system. Macleay developed his system under the shaping influence of two pressures: (1) the insistence by followers of Linnaeus on developing artificial systems at the expense of the natural system and (2) the apparent tension between the continuity of organic nature and the failure of linear classification schemes (which continuity seemed to require). Against what he perceived as dogmatic indolence on the part of the Linnaeans, Macleay developed a philosophy of science in which hypotheses that exceeded the available evidence should be proposed and subjected to severe tests. He also developed a novel comparative anatomical methodology, the method of variation, to aid in his search for the natural system. Using this method, he developed an intricate system that showed how organic nature could be continuous without being linear. A failure to appreciate these facets of Macleay's thought has led to several misunderstandings of him and his work, most notably that he was an idealist. These misunderstandings are here rebutted.