Abstract
In order to introduce protists to philosophers, we outline the diversity, classification, and evolutionary importance of these eukaryotic microorganisms. We argue that an evolutionary understanding of protists is crucial for understanding eukaryotes in general. More specifically, evolutionary protistology shows how the emphasis on understanding evolutionary phenomena through a phylogeny-based comparative approach constrains and underpins any more abstract account of why certain organismal features evolved in the early history of eukaryotes. We focus on three crucial episodes of this history: the origins of multicellularity, the origin of sex, and the origin of the eukaryote cell. Despite ongoing uncertainty about where the root of the eukaryote tree lies, and residual questions about the precise endosymbioses that have produced a diversity of photosynthesizing eukaryotes, evolutionary protistology has illuminated with considerable clarity many aspects of protist evolution. Our main message in light of evolutionary protistology is that these ‘other eukaryotes’ are in fact the organisms through which the rest of the eukaryotes should be understood.


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Notes
The ‘five kingdom system’ is often attributed to Whittaker (1969). However, his version intentionally included polyphyletic (not sharing a recent common ancestor) higher kingdoms and thus differs significantly in structure from the five-kingdom system popularized between the 1970s and 1990s. This more familiar version was proposed by Lynn Margulis (1971), who respectfully modified Whittaker’s scheme so that animals, plants and fungi were each potentially monophyletic, in effect by moving several groups to Protista from either fungi or plants.
Encephalitozoon intestinalis (a microsporidian) is the parasite with the smallest known protist genome; Gonyaulax polyedra (a dinoflagellate) has the largest reliably estimated genome (Gregory 2007). Even amongst parasitic protists (expected to have smaller genomes), there is considerable variation from just over two million base pairs to 160 million (Zubáčová et al. 2008). However, the size range of protist genomes is currently understood from very limited data (Gregory et al. 2007), and some earlier reports of extraordinarily large genomes in amoeba seem to be incorrect (Gregory 2005).
Roughly half of the dinoflagellates are photosynthetic. The remainder are free-living or parasitic heterotrophs.
Permanent anaerobiosis, the loss of the ability to use oxygen as a terminal electron acceptor during energy metabolism, has evolved a number of times in eukaryote evolution. There are particularly successful lineages of anaerobes in Excavata, but the property is not unique to this supergroup.
However, see Dickinson et al. (2012) for a suggestion that the various instances of multicellularity in Amorphea, usually considered ‘independent’, can be collapsed into a common origin. The current understanding of the deep-level diversity and phylogeny of Amorphea (e.g., Kim et al. 2006; Katz et al. 2012) makes this idea relatively unparsimonious.
HJC published as James-Clark; his actual surname was Clark even though this was his mother’s family name (his father’s was Porter).
Saville Kent also hyphenated his name occasionally; Saville was a second forename but he used it as the first part of his surname.
Integrin-related proteins have been found in bacteria as well as protists. In the former, their role is tentatively hypothesized as intracellular signaling (Chouhan et al. 2011).
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Acknowledgments
We thank Mark Olson (UNAM) for detailed comments that greatly clarified our argument. MAO acknowledges funding from the Australian Research Council and University of Sydney in the form of a Future Fellowship; AGBS is supported by the Canadian Institute for Advanced Research program in Integrated Microbial Biodiversity, and a Discovery grant from the Natural Sciences and Engineering Research Council of Canada; AJR is supported by the Canada Research Chairs Program and a Discovery grant from the Natural Sciences and Engineering Research Council of Canada.
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O’Malley, M.A., Simpson, A.G.B. & Roger, A.J. The other eukaryotes in light of evolutionary protistology. Biol Philos 28, 299–330 (2013). https://doi.org/10.1007/s10539-012-9354-y
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DOI: https://doi.org/10.1007/s10539-012-9354-y


