Philosophy of Biology
My primary focus has been on the philosophy of phylogenetic systematics. This is a rich area for philosophers to study, in no small part because biologists working in this area routinely write about and wrestle with philosophical topics in sophisticated and novel ways. Here's a few areas I spend time working on:
Phylogenetic Inference
Reconstructing evolutionary trees (phylogenies) presents several inferential challenges. There are typically going to be multiple, conflicting trees that are consistent with the data. How do we select from among these hypotheses?
More recently, I have become interested in how phylogenetic inference has both become more complicated yet more powerful given the vast amount of genealogical discordance we are discovering across different areas of living systems.
Biological Individuality
An enduring topic in systematics has been an ontological discussion over what kinds of things are species, lineages, and other taxonomic entities. I have defended the view that we ought to regard these things as biological individuals. I am working up what I call the Recursive Account of Biological Individuality to flesh out what it means to be a multilevel, dynamic, and complex biological individual.
Lately, I have been exploring how contemporary process ontology might be a useful stance to adopt in systematics, and how it is reminiscent of a stance adopted by mid-20th century systematists.
Classification and Taxonomy
Systematics includes taxonomy, classification, and nomenclature. These are very old sciences and we still employ tools that predate evolutionary theory to help us understand and categorize the vast biodiversity in the world. How those are reconciled (or not) with evolutionary and phylogenetic thinking is an ongoing discussion in biology, and provides a rich area to draw more general philosophical lessons.
The codes of nomenclature employed by biologists are rules for assigning proper names to taxa. I (and others) have argued that these should be treated as independently developed and empirically tested causal theories of reference. Understanding how they handle hard cases and what conventions have been adopted for them to function well can tell us a lot about reference, logic, language, and metaphysics.
Species
Biologists and philosophers have proposed many different concepts of what it means for a group of organisms to count as a species. There are also a lot of novel responses to this, from pluralism, to eliminativism, to dissolution. I am developing an optimistically skeptical account of this that I call The Species Problem Problem and the No Solution Solution. I think working out how to navigate this sort of situation will generalize across the sciences.
Bioethics
I have a smaller research profile in bioethics. Recently, much of that has been in partnership with students working in my Philosophy of Biology Lab. That includes a project on pediatric clinical ethics and work on the ethics of using AI in healthcare (especially mental health).
Pedagogy and Metaphilosophy
I have a goal to publish work on pedagogy and the tools I've been developing for teaching students both in and outside of the classroom. Additionally, my service as chair and associate dean provided an opportunity to reflect on administrative, organizational, and leadership topics. As areas of research, these are pretty new for me and I look forward to engaging with and contributing to this literature.
Abstract: Positively misleading errors are errors of statistical reasoning in which adding data to an analysis will systematically and reliably strengthen support for an erroneous hypothesis over a correct one. This pattern distinguishes them from other errors of statistical inference and pattern recognition. Here I provide a general account of positively misleading errors by describing an exemplar case from biology along with a candidate case from clinical medicine. Though well known in biology (phylogenetic systematics, to be precise), positively misleading errors are likely more widespread and deserve to be brought to the attention of the wider research community. This will facilitate a better understanding of them and sharpen our ability to assess statistical and probabilistic methods, providing resources for researchers to more effectively identify, diagnose, and dislodge these errors of statistical inference. This reflects the way we have gained a better understanding of scientific reasoning from studying other errors of statistical and probabilistic reasoning.
"Biology's Einstein Moment." (2025) Biological Theory (online only). doi: 10.1007/s13752-024-00475-3. (PDF)
Abstract: We are currently in the midst of what I call biology’s Einstein moment. This is the rejection of absolute biological history, the idea that there is an invariant, privileged biological history against which other histories are measured or deviate from. Instead, biologists must specify theoretically and empirically motivated frames of lineal reference. This is already informing and advancing biological practice, theory, methods, and more, and is a significant and important feature of contemporary biology. Here I argue that it is worth identifying and naming this shift, and encouraging a deeper and broader embrace of it.
“Changing Working Environments in Philosophy: Reflections from a Case Study.” Philosophy of Science. Co-authors: Alison K. McConwell, Magdalena T. Bogacz, Char Brecevic, Jingyi Wu, Sarah M. Roe. (In Press). doi: 10.1017/psa.2025.12. (PDF)
Abstract: There is an "under-representation problem” in philosophy departments and journals. Empirical data suggest that while we have seen some improvements since the 1990s, the rate of change has slowed down. Some posit that philosophy has disciplinary norms making it uniquely resistant to change. We present results from an empirical case study of a philosophy department that achieved and maintained male-female gender parity among its faculty as early as 2014. Our analysis extends beyond matters of gender parity because that is only one, albeit important, dimension of inclusion. We build from the study to reflect on strategies that may catalyze change.
“Justice, Vulnerable Populations, and the Use of Conversational AI in Psychotherapy.” The American Journal of Bioethics. Co-authors: Bennett Knox, Pierce Christoffersen, Kalista Leggitt, Zeia Woodruff. doi: 10.1080/15265161.2023.2191040 (2023). (PDF)
“The Individuality Thesis (3 Ways)” Biology & Philosophy, 31(6):913–930. doi: 10.1007/s10539-016-9548-9. (2016) (PDF)
Abstract: I spell out and update the individuality thesis, that species are individuals, and not classes, sets, or kinds. I offer three complementary presentations of this thesis. First, as a way of resolving an inconsistent triad about natural kinds; second, as a phylogenetic systematics theoretical perspective; and, finally, as a novel recursive account of an evolved character (individuality). These approaches do different sorts of work, serving different interests. Presenting them together produces a taxonomy of the debates over the thesis, and isolates ways it has been (and may continue to be) productive. This goes to the larger point of this paper: a defense of the individuality thesis in terms of its utility, and an update of it in light of recent theoretical developments and empirical work in biology.
“Species in the Age of Discordance.” Philosophy, Theory, and Practice in Biology, 11(21). doi: 10.3998/ptpbio.16039257.0011.021 (2019). (PDF)
Abstract: Biological lineages move through time, space, and each other. As they do, they diversify, diverge, and grade away from and into one another. One result of this is genealogical discordance; i.e., the lineages of a biological entity may have different histories. We see this on numerous levels, from microbial networks, to holobionts, to population-level lineages. This paper considers how genealogical discordance impacts our study of species. More specifically, I consider this in the context of three framing questions: (1) How, if at all, does genealogical discordance challenge, modify, or revise how we conceive of species? (2) How has growing appreciation of genealogical discordance impacted scientific practice? Of systematics in particular? (3) How do lineages at different levels diverge and diversify? All told, genealogical discordance enriches and complicates our taxonomic ontology, as well as the practice of reconstructing phylogenies. This presents both challenges and opportunities for the study of divergence and diversification.
“Multilevel Lineages & Multidimensional Trees: The Levels of Lineage and Phylogeny Reconstruction.” Philosophy of Science, 79(5):609–623. doi: 10.1086/667849. (2012) (PDF)
Abstract: The relation among method, concept, and theory in science is complicated. I seek to shed light on that relation by considering an instance of it in systematics: the additional challenges phylogeneticists face when reconstructing phylogeny not at a single level but simultaneously at multiple levels of the hierarchy. How does this complicate the task of phylogenetic inference, and how might it inform and shape the conceptual foundations of phylogenetics? This offers a lens through which the interplay of method, theory, and concepts may be understood in systematics, which, in turn, provides data for a more general account.
“How To Misidentify A Type Specimen.” Biology & Philosophy, 27(6):767- 784. doi: 10.1007/s10539-012-9336-0. (2012) (PDF)
Abstract: Type specimens are used to designate species. What is the nature of the relation between a type specimen and the species it designates? If species names are rigid designators, and type specimens ostensively define species, then that relation is, at the very least, a close one. Levine (Biol Philos 16(3):325–338, 2001) argues that the relationship of type specimen to a named species is one of necessity—and that this presents problems for the individuality thesis. Namely, it seems odd that a contingently selected specimen should belong to a species of necessity. In considering Levine’s argument, LaPorte (Biol Philos 18:583–588, 2003) suggests that recognizing the distinction between de re and de dicto necessity resolves Levine’s worries. I reconsider the motivating question: does a type specimen belong of necessity to the species that it designates? In light of taxonomic cases and practice the answer is clear: definitively not. This is particularly clear in the case of re-designation of types by taxonomic decree. I explain how this helps reveal how taxonomists prioritize competing (and sometimes conflicting) theoretical commitments, and offer a defense of the individuality thesis as applied to these particular cases. In short, I demonstrate how to misidentify a type specimen.
Abstract: Phylogenetics is the study of the evolutionary history and relationships among individuals, groups of organisms (e.g., populations, species, or higher taxa), or other biological entities with evolutionary histories (e.g., genes, biochemicals, or developmental mechanisms). Phylogenetic inference is the task of inferring this history, and as with other problems of inference, there are interesting and difficult questions regarding how these inferences are justified.
In this entry, we examine what phylogenetic inference is and how it works. In the first section, we briefly introduce the field of phylogenetics and its history. In section 2, we explore how phylogenetic inference provides useful problems for philosophers to examine, and where philosophical approaches have contributed to the scientific examination of phylogenetics. This will help display why phylogenetic inference is not merely a biological research problem, but a philosophical one as well. Finally in section 3, we will move to a discussion of some of the contemporary debates about foundational issues in phylogenetics, and what the future of phylogenetic inference looks like.
“The Biological and the Mereological: Metaphysical Implications of the Individuality Thesis.” In Individuals Across the Sciences, Thomas Pradeu and Alexandre Guay, eds., 295–316. Oxford University Press. doi: 10.1093/acprof:oso/9780199382514.003.0016. (2016) (PDF)
Abstract: Michael Ghiselin and David Hull’s individuality thesis is that biological species are individuals. Philosophers often treat the term “individual” as synonymous with “mereological sum” and characterize it in terms of mereology (an axiomatic system of parts and wholes analogous to set theory). It is easy to see how the biological project (that species are individuals) has been interpreted as a mereological one (that species are mereological sums). This chapter argues that this is a mistake, that biological part/whole relations often violate the axioms of mereology. Conflating these projects confuses the central issues at stake in both, and makes the job of evaluating either unnecessarily burdensome. Clarifying this helps identify the genuine metaphysical implications of the individuality thesis, which serves as an exemplar of scientifically informed metaphysics.
“Colonies Are Individuals: Revisiting the Superorganism Revival.” In From Groups to Individuals: Perspectives on Biological Associations and Emerging Individuality, Frédéric Bouchard and Philippe Huneman, eds., 195– 217 (ch. 9). MIT Press. doi: 10.7551/mitpress/8921.003.0015. (2013) (PDF)
Editorial: Special and Thematic Issues
Thematic Section: “Biological Lineages.” Biological Theory. Co-edited with Javier Suárez. (Forthcoming).
Special Issue: “Species in the Age of Discordance.” Philosophy, Theory, and Practice in Biology, 11. (2019).
Special Issue: “The Edges & Boundaries of Biological Objects.” Biological Theory 4(3). Co-edited with Jay Odenbaugh. (2009)
Articles & Chapters
"Positively Misleading Errors." (Forthcoming) Synthese. doi: 10.1007/s11229-025-05093-5. (PDF)
"Biology's Einstein Moment." (2025) Biological Theory (online only). doi: 10.1007/s13752-024-00475-3. (PDF)
“Changing Working Environments in Philosophy: Reflections from a Case Study.” Philosophy of Science. Co-authors: Alison K. McConwell, Magdalena T. Bogacz, Char Brecevic, Jingyi Wu, Sarah M. Roe. (In Press). doi: 10.1017/psa.2025.12. (PDF)
“Introduction to “Formalization and the Meaning of ‘Theory’ in the Inexact Biological Sciences”.” In Scaffolding: Selected Contributions of James R. Griesemer to History, Philosophy, and Biology, Rachel Ankeny, Sabina Leonelli, and Michael Dietrich, eds. Springer. (2025). (PDF)
“Justice, Vulnerable Populations, and the Use of Conversational AI in Psychotherapy.” The American Journal of Bioethics. Co-authors: Bennett Knox, Pierce Christoffersen, Kalista Leggitt, Zeia Woodruff. doi: 10.1080/15265161.2023.2191040 (2023). (PDF)
“Phylogenetic Inference.” The Stanford Encyclopedia of Philosophy. Co-author: Joel Velasco. link: SEP Entry: Phylogenetic Inference (2021; revised 2022). (PDF)
“Species in the Age of Discordance.” Philosophy, Theory, and Practice in Biology, 11(21). doi: 10.3998/ptpbio.16039257.0011.021 (2019). (PDF)
“Species in the Age of Discordance: Meeting Report and Introduction.” Philosophy, Theory, and Practice in Biology, 11(12). Co-author: Daniel J. Molter. doi: 10.3998/ptpbio.16039257.0011.012 (2019). (PDF)
“The Individuality Thesis (3 Ways)” Biology & Philosophy, 31(6):913–930. doi: 10.1007/s10539-016-9548-9. (2016) (PDF)
“The Biological and the Mereological: Metaphysical Implications of the Individuality Thesis.” In Individuals Across the Sciences, Thomas Pradeu and Alexandre Guay, eds., 295–316. Oxford University Press. doi: 10.1093/acprof:oso/9780199382514.003.0016. (2016) (PDF)
“Mitochondrial Diversity and the Reversal Test” American Journal of Bioethics, 15(6):23–24. Co-author: Madeline Bannon. doi: 10.1080/15265161.2015.1028669. (2015) (PDF)
“Colonies Are Individuals: Revisiting the Superorganism Revival.” In From Groups to Individuals: Perspectives on Biological Associations and Emerging Individuality, Frédéric Bouchard and Philippe Huneman, eds., 195– 217 (ch. 9). MIT Press. doi: 10.7551/mitpress/8921.003.0015. (2013) (PDF)
“Multilevel Lineages & Multidimensional Trees: The Levels of Lineage and Phylogeny Reconstruction.” Philosophy of Science, 79(5):609–623. doi: 10.1086/667849. (2012) (PDF)
“How To Misidentify A Type Specimen.” Biology & Philosophy, 27(6):767- 784. doi: 10.1007/s10539-012-9336-0. (2012) (PDF)
“Reframing the Ethical Issues in Part-Human Animal Research: The Unbearable Ontology of Inexorable Moral Confusion.” The American Journal of Bioethics, 12(9): 17–25. Co-authored with Bryan Benham. doi: 10.1080/15265161.2012.699139. (2012) (PDF)
“Philosophy of Biology” in Philosophy of the Special Sciences. F. Allhoff, ed. Blackwell Press. 184–212. Co-authored with Andrew Hamilton, Jay Odenbaugh and Samir Okasha. (2010)
“The Edges and Boundaries of Biological Objects.” Biological Theory, 4(3):219– 224. Co-authored with Jay Odenbaugh. doi: 10.1162/biot.2009.4.3.219. (2009) (PDF)
“Clade Selection and Levels of Lineage: A Reply to Rieppel.” Biological Theory, 4(2):214–218. Co-authored with Andrew Hamilton. doi: 10.1162/biot.2009.4.2.214. (2009) (PDF)
“Social Insects and the Individuality Thesis: Cohesion and the Colony as a Selectable Individual.” In Organization of Insect Societies: From Genome to Sociocomplexity. J. Gadau and J. Fewell, eds. Harvard University Press. 570–587. Co-authored with Andrew Hamilton and Nathan Robert Smith. (2009) (PDF)
“Moral Confusion and Developmental Essentialism in part-human hybrid research.” The American Journal of Bioethics, 8(12): 42–43. Co-author Bryan Benham. doi: 10.1080/15265160802559195. (2008) (PDF)
“Phylogenetic Inference.” In Blackwell Companion to the Philosophy of History and Historiography, Avi Tucker, ed., 231–242. Blackwell. doi: 10.1002/9781444304916.ch20. (2008) (PDF)
“Clades Are Reproducers." Biological Theory, 1(4): 381–391. Co-authored with Andrew Hamilton. doi: 10.1162/biot.2006.1.4.381. (2006) (PDF)
“Coherence, Consistency and Cohesion: Clade Selection in Okasha and Beyond.” Philosophy of Science, 72(5): 1026–1040. Co-authored with Andrew Hamilton. doi: 10.1086/508101. (2005) (PDF)
“Probability and Systematics: Possibility, Probability, and Phylogenetic Inference.” Systematic Biology, 54: 831–841. doi: 10.1080/106351591007444. (2005) (PDF)
“Morphology, Ultrastructure, and Function of Extrafloral Nectaries in Three Species of Caesalpiniacae.” Jour. Iowa Acad. Sci., 106(4) 82–88. Coauthored with Lenore T. Durkee, Lisa Dorn and Ann Remington. (1999) (PDF)
Book Reviews
“Species Problems.” The Species Problem: A Philosophical Analysis, by Richard A. Richards, Metascience, 22:339–342. doi: 10.1007/s11016-012-9720-z. (2013) (PDF)
Seeking God in Science: An Atheist Defends Intelligent Design, by Bradley Monton, Reports of the National Center for Science Education, 32(5):5.1–5.3 (2012) (PDF)
"Not So Innocent: Methodology and Metaphysics of Evolution.” Doubting Darwin? Creationist Designs on Evolution, by Sahotra Sarkar, BioScience, 58(11): 1088–1089. doi: 10.1641/B581113. (2008)
Evolution and the Levels of Selection, by Samir Okasha, Mind, 117(468):1116–1119. doi: 10.1093/mind/fzn141 (2008)
The Poverty of the Linnaean Hierarchy: A Philosophical Study of Biological Taxonomy, by Marc Ereshefsky, Philosophy of Science, 72(3):491–494. doi: 10.1086/498478 (2005)