The concept of individuality as applied to species, an important advance in the philosophy of evolutionary biology, is nevertheless in need of refinement. Four important subparts of this concept must be recognized: spatial boundaries, temporal boundaries, integration, and cohesion. Not all species necessarily meet all of these. Two very different types of pluralism have been advocated with respect to species, only one of which is satisfactory. An often unrecognized distinction between grouping and ranking components of any (...) class='Hi'>species concept is necessary. A phylogenetic species concept is advocated that uses a grouping criterion of monophyly in a cladistic sense, and a ranking criterion based on those causal processes that are most important in producing and maintaining lineages in a particular case. Such causal processes can include actual interbreeding, selective constraints, and developmental canalization. The widespread use of the biological species concept is flawed for two reasons: because of a failure to distinguish grouping from ranking criteria and because of an unwarranted emphasis on the importance of interbreeding as a universal causal factor controlling evolutionary diversification. The potential to interbreed is not in itself a process; it is instead a result of a diversity of processes which result in shared selective environments and common developmental programs. These types of processes act in both sexual and asexual organisms, thus the phylogenetic species concept can reflect an underlying unity that the biological species concept can not. (shrink)

Species serve as both the basic units of macroevolutionary studies and as the basic units of taxonomic classification. In this paper I argue that the taxa identified as species by the Phylogenetic Species Concept (Mishler and Brandon 1987) are the units of biological organization most causally relevant to the evolutionary process but that such units exist at multiple levels within the hierarchy of any phylogenetic lineage. The PSC gives us no way of identifying one of (...) these levels as the privileged level on which taxonomic classifications can be based. (shrink)

The phylogenetic species concept is applied for the first time to a major radiation of birds, the birds-of-paradise (Paradisaeidae) of Australasia. Using the biological species concept, previous workers have postulated approximately 40–42 species in the family. Of these, approximately 13 are monotypic and 27 are polytypic with about 100 subspecies. Phylogenetic species are irreducible (basal) clusters of organisms (terminal taxa) that are diagnosably distinct from other such clusters. Within the context of this concept, approximately (...) 90 species of paradisaeids are postulated to have diversified within Australasia. The phylogenetic species concept more accurately describes evolutionary diversity within the family and provides a better theoretical and empirical framework for analysing speciation, historical biogeography and patterns of morphological, behavioral and ecological diversification within this group than does the biological species concept. (shrink)

A natural starting place for developing a phylogenetic species concept is to examine monophyletic groups of organisms. Proponents of “the” Phylogenetic Species Concept fall into one of two camps. The first camp denies that species even could be monophyletic and groups organisms using character traits. The second groups organisms using common ancestry and requires that species must be monophyletic. I argue that neither view is entirely correct. While monophyletic groups of organisms exist, they should (...) not be equated with species. Instead, species must meet the more restrictive criterion of being genealogically exclusive groups where the members are more closely related to each other than to anything outside the group. I carefully spell out different versions of what this might mean and arrive at a working definition of exclusivity that forms groups that can function within phylogenetic theory. I conclude by arguing that while a phylogenetic species concept must use exclusivity as a grouping criterion, a variety of ranking criteria are consistent with the requirement that species can be placed on phylogenetic trees. (shrink)

Frank E. Zachos offers a comprehensive review of one of today's most important and contentious issues in biology: the species problem. After setting the stage with key background information on the topic, the book provides a brief history of species concepts from antiquity to the Modern Synthesis, followed by a discussion of the ontological status of species with a focus on the individuality thesis and potential means of reconciling it with other philosophical approaches. More than 30 (...) different species concepts found in the literature are presented in an annotated list, and the most important ones, including the Biological, Genetic, Evolutionary and different versions of the Phylogenetic Species Concept, are discussed in more detail. Specific questions addressed include the problem of asexual and prokaryotic species, intraspecific categories like subspecies and Evolutionarily Significant Units, and a potential solution to the species problem based on a hierarchical approach that distinguishes between ontological and operational species concepts. A full chapter is dedicated to the challenge of delimiting species by means of a discrete taxonomy in a continuous world of inherently fuzzy boundaries. Further, the book outlines the practical ramifications for ecology and evolutionary biology of how we define the species category, highlighting the danger of an apples and oranges problem if what we subsume under the same name ("species") is in actuality a variety of different entities. A succinct summary chapter, glossary and annotated list of references round out the coverage, making the book essential reading for all biologists looking for an accessible introduction to the historical, philosophical and practical dimensions of the species problem. (shrink)

Biologists and philosophers have long recognized the importance of species, yet species concepts serve two masters, evolutionary theory on the one hand and taxonomy on the other. Much of present-day evolutionary and systematic biology has confounded these two roles primarily through use of the biological species concept. Theories require entities that are real, discrete, irreducible, and comparable. Within the neo-Darwinian synthesis, however, biological species have been treated as real or subjectively delimited entities, discrete or nondiscrete, (...) and they are often capable of being decomposed into other, smaller units. Because of this, biological species are generally not comparable across different groups of organisms, which implies that the ontological structure of evolutionary theory requires modification. Some biologists, including proponents of the biological species concept, have argued that no species concept is universally applicable across all organisms. Such a view means, however, that the history of life cannot be embraced by a common theory of ancestry and descent if that theory uses species as its entities.These ontological and biological difficulties can be alleviated if species are defined in terms of evolutionary units. The latter are irreducible clusters of reproductively cohesive organisms that are diagnosably distinct from other such clusters. Unlike biological species, which can include two or more evolutionary units, these phylogenetic species are discrete entities in space and time and capable of being compared from one group to the next. (shrink)

Many phylogenetic systematists have criticized the Biological Species Concept (BSC) because it distorts evolutionary history. While defenses against this particular criticism have been attempted, I argue that these responses are unsuccessful. In addition, I argue that the source of this problem leads to previously unappreciated, and deeper, fatal objections. These objections to the BSC also straightforwardly apply to other species concepts that are not defined by genealogical history. What is missing from many previous discussions is the (...) fact that the Tree of Life, which represents phylogenetic history, is independent of our choice of species concept. Some species concepts are consistent with species having unique positions on the Tree while others, including the BSC, are not. Since representing history is of primary importance in evolutionary biology, these problems lead to the conclusion that the BSC, along with many other species concepts, are unacceptable. If species are to be taxa used in phylogenetic inferences, we need a history-based species concept. (shrink)

This chapter contains sections titled: Introduction Early Species Concepts—Linnaeus Biological Species Concepts Phylogenetic Species Concepts Species Concepts and Speciation Conclusions Postscript: Counterpoint References.

This paper isolates a hard, long-standing species problem: developing a comprehensive and exacting theory about the constitutive conditions of the species category, one that is accurate for most of the living world, and which vindicates the widespread view that the species category is of more theoretical import than categories such as genus, sub-species, paradivision, and stirp. The paper then uncovers flaws in several views that imply we have either already solved that hard species problem or (...) dissolved it altogether – so-called We Are Done views. In doing so the paper offers new criticisms of the general lineage species concept (GLSC), evolutionary species concept (EvSC), biological species concept (BSC), other similar concepts, Ereshefsky’s eliminative pluralism about the species category, and both Mishler’s pessimism and Wilkins’ phenomenalism about that category. Opposed to We Are Done views, the paper argues for a Revving Up view, on which we are nearly ready to begin the hard species problem in earnest. To help work towards these conclusions, the paper begins with an outline of a new kind of view of species (Barker 2019a), which proposes they are feedback systems of a mathematically specifiable and empirically testable sort. (shrink)

Recent work in viral genomics has shown that bacteriophages exhibit a high degree of mosaicism, which is most likely due to a long history of prolific horizontal gene transfer (HGT). Given these findings, we argue that each of the most plausible attempts to properly classify bacteriophages into distinct species fail. Mayr's biological species concept fails because there is no useful viral analog to sexual reproduction. Phenetic species concepts fail because they obscure the mosaicism and the rich (...) reticulated viral histories. Phylogenetic species concepts, even when extended to take into account reticulation, fail because there is no non-arbitrary distinction between recombination events that create a new viral species and those that do not. There is good reason to think that bacteriophages, arguably the Earth's most abundant biological agent, evolve without forming species.1. Introduction2. The Biology of Viruses2.1. Bacteriophage life cycles2.2. Mechanisms of HGT3. The Species Problem and Species Concepts3.1. Phenetic species concepts3.2. The biological species concept3.3. Phylogenetic species concepts3.4. The ecological species concept3.5. Homeostatic property cluster species4. Viruses and Species Taxonomy5. Reticular Phylogenies6. Conclusion. (shrink)

In the last thirty years systematics has transformed itself from a discipline concerned with classification into a discipline concerned with reconstructing the evolutionary history of life. This transformation has been driven by cladistic analysis, a set of techniques for reconstructing evolutionary trees. Long interested in the large-scale structure of evolutionary history, cladistically oriented systematists have recently begun to apply "tree thinking" to problems near the species level. ¶ In any local ("non-dimensional") situation species are usually well-defined, but across (...) space and time the grouping of populations into species is often problematic. Three views of species are in common use today, the biological species concept, the evolutionary species concept, and the phylogenetic species concept. Each of these has strengths and weaknesses, but no matter which is applied, exact counts of the number of species in any extended area will always be ambiguous no matter how much factual information is available. This ambiguity arises because evolution is an historical process, and the grouping of organisms into species always depends to some extent upon expectations of the future behavior of those organisms and their descendants, expectations that cannot be evaluated in the present. The existence and special character of the species problem is itself one of the central pieces of evidence for evolution. (shrink)

An examination of the post-Darwinian history of biological taxonomy reveals an implicit assumption that the definitions of taxon names consist of lists of organismal traits. That assumption represents a failure to grant the concept of evolution a central role in taxonomy, and it causes conflicts between traditional methods of defining taxon names and evolutionary concepts of taxa. Phylogenetic definitions of taxon names (de Queiroz and Gauthier 1990) grant the concept of common ancestry a central role in the definitions (...) of taxon names and thus constitute an important step in the development of phylogenetic taxonomy. By treating phylogenetic relationships rather than organismal traits as necessary and sufficient properties, phylogenetic definitions remove conflicts between the definitions of taxon names and evolutionary concepts of taxa. The general method of definition represented by phylogenetic definitions of clade names can be applied to the names of other kinds of composite wholes, including populations and biological species. That the names of individuals (composite wholes) can be defined in terms of necessary and sufficient properties provides the foundation for a synthesis of seemingly incompatible positions held by contemporary individualists and essentialists concerning the nature of taxa and the definitions of taxon names. (shrink)

College of Medicine, University of South Alabama Mobile, AL 36688-0002, USA wbp501{at}jaguar1.usouthal.edu ' + u + '@' + d + ' '//--> Abstract Recent work in viral genomics has shown that bacteriophages exhibit a high degree of mosaicism, which is most likely due to a long history of prolific horizontal gene transfer (HGT). Given these findings, we argue that each of the most plausible attempts to properly classify bacteriophages into distinct species fail. Mayr's biological species concept fails because (...) there is no useful viral analog to sexual reproduction. Phenetic species concepts fail because they obscure the mosaicism and the rich reticulated viral histories. Phylogenetic species concepts, even when extended to take into account reticulation, fail because there is no non-arbitrary distinction between recombination events that create a new viral species and those that do not. There is good reason to think that bacteriophages, arguably the Earth's most abundant biological agent, evolve without forming species. Introduction The Biology of Viruses 2.1 Bacteriophage life cycles 2.2 Mechanisms of HGT The Species Problem and Species Concepts 3.1 Phenetic species concepts 3.2 The biological species concept 3.3 Phylogenetic species concepts 3.4 The ecological species concept 3.5 Homeostatic property cluster species Viruses and Species Taxonomy Reticular Phylogenies Conclusion CiteULike Connotea Del.icio.us What's this? (shrink)

The species problem is often described as the abundance of conflicting definitions of _species_, such as the biological species concept and phylogenetic species concepts. But biologists understand the notion of species in a non-definitional as well as a definitional way. In this article I argue that when they understand _species_ without a definition in their mind, their understanding is often mediated by the notion of _good species_, or prototypical species, as the idea of (...) ``prototype'' is explicated in cognitive psychology. This distinction helps us make sense of several puzzling phenomena regarding biologists' dealing with species, such as the fact that in everyday research biologists often behave as if the species problem is solved, while they should be fully aware that it is not. I also briefly discuss implications of this finding, including that some extant attempts to answer what the nature of species is have an inadequate assumption about how the notion of species is represented in biologists' minds. (shrink)

In addition to the distinction between species as a category and speciesas a taxon, the word species is ambiguous in a very different butequally important way, namely the temporal distinction between horizontal andvertical species. Although often found in the relevant literature, thisdistinction has thus far remained vague and undefined. In this paper the use ofthe distinction is explored, an attempt is made to clarify and define it, andthen the relation between the two dimensions and the implications of (...) thatrelation are examined. Using Darwin's analogy of language evolution forspeciesevolution, and by appealing to a major change in the conception of languagebetween 19th- and 20th-century linguistics, it is argued that the horizontaldimension has priority (pragmatic, epistemological, logical, and ontological)over the vertical dimension. This has immense ramifications for the modernspecies problem. Fundamentally, it favors horizontal species concepts oververtical ones. In so doing it places species realism on a much more securefoundation and largely undercuts species pluralism. In addition it raises aserious problem for the increasingly popular family of phylogenetic speciesconcepts, which generally suffer from a dimensionality confusion. However,thereis a recent trend within this family that attempts to restore the priority ofthe horizontal dimension. It is concluded that this trend should be affirmedandthat the species-as-individuals view should be abandoned. (shrink)

The long-awaited revision of the industry standard on phylogenetics Since the publication of the first edition of this landmark volume more than twenty-five years ago, phylogenetic systematics has taken its place as the dominant paradigm of systematic biology. It has profoundly influenced the way scientists study evolution, and has seen many theoretical and technical advances as the field has continued to grow. It goes almost without saying that the next twenty-five years of phylogenetic research will prove as fascinating (...) as the first, with many exciting developments yet to come. This new edition of Phylogenetics captures the very essence of this rapidly evolving discipline. Written for the practicing systematist and phylogeneticist, it addresses both the philosophical and technical issues of the field, as well as surveys general practices in taxonomy. Major sections of the book deal with the nature of species and higher taxa, homology and characters, trees and tree graphs, and biogeography—the purpose being to develop biologically relevant species, character, tree, and biogeographic concepts that can be applied fruitfully to phylogenetics. The book then turns its focus to phylogenetic trees, including an in-depth guide to tree-building algorithms. Additional coverage includes: Parsimony and parsimony analysis Parametric phylogenetics including maximum likelihood and Bayesian approaches Phylogenetic classification Critiques of evolutionary taxonomy, phenetics, and transformed cladistics Specimen selection, field collecting, and curating Systematic publication and the rules of nomenclature Providing a thorough synthesis of the field, this important update to Phylogenetics is essential for students and researchers in the areas of evolutionary biology, molecular evolution, genetics and evolutionary genetics, paleontology, physical anthropology, and zoology. (shrink)

In this paper, we discuss the following four alternative ways of understanding the outcomes of resurrection biology. Implications of each of the ways are discussed with respect to concepts of species and extinction. Replication: animals created by resurrection biology do not belong to the original species but are copies of it. The view is compatible with finality of extinction as well as with certain biological and ecological species concepts. Re-creation: animals created are members of the (...) original species but, despite their existence, the species remains extinct. The view is incompatible with all species concepts presented. Non-extinction: animals produced belong to the original species which actually never went extinct. The view may be consistent with phenetic and phylogenetic species concepts as well as with finality of extinction. According to literal resurrection, resurrection biology is successful in reversing extinction through the creation of new members of species that once went extinct. This view presupposes non-finality of extinction and it is compatible with phenetic species concepts. It is notable that no species or extinction concept is consistent with all possible views of resurrection biology nor is any view of resurrection biology consistent with all species or extinction concepts. Thus, one’s views regarding species and extinction determine which views one can adopt regarding resurrection biology and vice versa. (shrink)

Many groups of closely related species have reticulate phylogenies. Recent genomic analyses are showing this in many insects and vertebrates, as well as in microbes and plants. In microbes, lateral gene transfer is the dominant process that spoils strictly tree‐like phylogenies, but in multicellular eukaryotes hybridization and introgression among related species is probably more important. Because many species, including the ancestors of ancient major lineages, seem to evolve rapidly in adaptive radiations, some sexual compatibility may exist among (...) them. Introgression and reticulation can thereby affect all parts of the tree of life, not just the recent species at the tips. Our understanding of adaptive evolution, speciation, phylogenetics, and comparative biology must adapt to these mostly recent findings. Introgression has important practical implications as well, not least for the management of genetically modified organisms in pest and disease control. (shrink)

An examination of the post-Darwinian history of biological taxonomy reveals an implicit assumption that the definitions of taxon names consist of lists of organismal traits. That assumption represents a failure to grant the concept of evolution a central role in taxonomy, and it causes conflicts between traditional methods of defining taxon names and evolutionary concepts of taxa. Phylogenetic definitions of taxon names (de Queiroz and Gauthier 1990) grant the concept of common ancestry a central role in the definitions (...) of taxon names and thus constitute an important step in the development of phylogenetic taxonomy. By treating phylogenetic relationships rather than organismal traits as necessary and sufficient properties, phylogenetic definitions remove conflicts between the definitions of taxon names and evolutionary concepts of taxa. The general method of definition represented by phylogenetic definitions of clade names can be applied to the names of other kinds of composite wholes, including populations and biological species. That the names of individuals (composite wholes) can be defined in terms of necessary and sufficient properties provides the foundation for a synthesis of seemingly incompatible positions held by contemporary individualists and essentialists concerning the nature of taxa and the definitions of taxon names. (shrink)

This chapter contains sections titled: Historical and Current Views of Species Return to a Darwinian View of Species Practical Implications Postscript: Counterpoint References.

This afterword to Species and Beyond provides some reflections on species, with special attention to what I think the most significant developments have been in the thinking of biologists and philosophers working on species over the past 25 years, as well as some bad jokes.

A common reaction to functional diversity is to group entities into clusters that are functionally similar. I argue here that people are diverse with respect to reasoning-related processes, and that these processes satisfy the basic requirements for evolving entities: they are heritable, mutable, and subject to selective pressures. I propose a metric to quantify functional difference and show how this can be used to place psychological processes into a structure akin to a phylogenetic or evolutionary tree. Three species (...) concepts are repurposed from biology and used to understand relationships in that tree. (shrink)

Species, or ‘the Species Problem’, is a topic in science, in the philosophy of science, and in general philosophy. There is not one, but many, species problems, and these are dealt with in this volume. Species are often thought of as units of biology, to be used in ecology, conservation, classification, and theory. The chapters in this book present opposing views on the current philosophical and conceptual issues of the Species Problem in biology. -/- Divided (...) into four sections Theories and Concepts, Practice and Methods, Ranks, Trees and Names, and Metaphysics and Epistemologies, the book is authored by systematists, philosophers, historians and working biologists, many leaders in their fields. Topics include Essentialism and individuals, speciation modes, theory and the species rank, nomenclature and diagnosis, scale of species, and ecology, conservation, and species. -/- Species Problems and Beyond aims to clarify the contemporary issues of the Species Problem. It is ideal for use in upper-level seminars and courses in Evolutionary Biology, Philosophy of Science, Philosophy of Biology, Systematics and Taxonomy, and Phylogenetics/Cladistics, and for any scholar in these fields. (shrink)

In biological systematics, as well as in the philosophy of biology, species and higher taxa are individuated through their unique evolutionary origin. This is taken by some authors to mean that monophyly is a (relational) property not only of higher taxa, but also of species. A species is said to originate through speciation, and to go extinct when it splits into two daughter species (or through terminal extinction). Its unique evolutionary origin is said to bestow identity (...) on a species through time and change, and to render species names rigid designators. Species names are thus believed to function just like names of supraspecific taxa. However, large parts of the Web of Life are composed of species that do not have a unique evolutionary origin from a single population, lineage or stem-species. Further, monophyly is an ambiguous concept if it is defined simply in terms of ‘unique evolutionary origin’. Disambiguating the concept by defining a monophyletic taxon as ‘a taxon that includes the ancestor and all, and only, its descendant’ renders monophyly inapplicable to species. At the heart of the problem lies a fundamental distinction between species and monophyletic taxa, where species form mutually exclusive reticulated systems, while higher taxa form inclusive hierarchical systems. Examples are given both at the species level and below to illustrate the problems that result from the application of the monophyly criterion to species. The conclusion is that the concepts of exclusivity and monophyly should be treated as non-overlapping: exclusivity marks out a species synchronistically, i.e. in the present time. Monophyly marks out clades (groups of species) diachronistically, i.e. within an historical dimension. (shrink)

A. W. F. Edwards is one of the most influential mathematical geneticists in the history of the discipline. One of the last students of R. A. Fisher, Edwards pioneered the statistical analysis of phylogeny in collaboration with L. L. Cavalli-Sforza, and helped establish Fisher's concept of likelihood as a standard of statistical and scientific inference. In this book, edited by philosopher of science Rasmus Grønfeldt Winther, Edwards's key papers are assembled alongside commentaries by leading scientists, discussing Edwards's influence on their (...) own research and on thinking in their field overall. In an extensive interview with Winther, Edwards offers his thoughts on his contributions, their legacy, and the context in which they emerged. This book is a resource both for anyone interested in the history and philosophy of genetics, statistics, and science, and for scientists seeking to develop new algorithmic and statistical methods for understanding the genetic relationships between and among species both extant and extinct. (shrink)

We argue that the logical outcome of the cladistics revolution in biological systematics, and the move towards rankless phylogenetic classification of nested monophyletic groups as formalized in the PhyloCode, is to eliminate the species rank along with all the others and simply name clades. We propose that the lowest level of formally named clade be the SNaRC, the Smallest Named and Registered Clade. The SNaRC is an epistemic level in the classification, not an ontic one. Naming stops at (...) that level because there is no currently acceptable evidence for clades within it, not because no smaller clades exist. Later, included clades may be named. They would then become the SNaRCs, while the original SNaRC would keep its original name. We argue that all theoretical tasks of biology, in evolution and ecology, as well as practical tasks such as conservation assessment, are better approached using this rankless phylogenetic approach. (shrink)

Treating species as individuals and not classes has been crucial to the integration of evolutionary theory with modern systematics. Despite the theoretically important role the concept of individuality plays in modern phylogenetic systematics and in evolutionary theory more generally, many have been content to rely on common-sense intuitions about what counts as an individual. One of the most often cited intuitions is that individuals should be defined intrinsically. Unfortunately, common-sense intuitions like this one have proven to be inadequate (...) for identifying and characterizing historical individuals (like species). An examination of real-world biological examples shows that our common-sense intuitions are equally inadequate when applied to at least some biological organisms--the paradigm individuals. (shrink)

The project of this paper is to understand what a phylogenetic tree represents and to discuss some of the implications that this has for the practice of systematics. At least the first part of this task, if not both parts, might appear trivial—or perhaps better suited for a single page in a textbook rather than a scholarly research paper. But this would be a mistake. While the task of interpreting phylogenetic trees is often treated in a trivial way, (...) their interpretation is tied to foundational conceptual questions at the heart of systematics—questions whose answers are hotly disputed. I have previously argued that widely shared ideas about the meaning and interpretation of phylogenetic trees are inconsistent with species concepts other than some genealogical version of a phylogenetic species concept (Velasco 2008). Here I rely on a similar approach and concentrate on the implications of the necessary conditions underlying the inferences that we make using phylogenetic trees. I argue that common practices for the interpretation and use of trees are in conflict and that unacceptable principles about species as units of phylogeny must be given up. According to the view that I will develop, all phylogenetic trees depict the history of populations. The branches on trees represent collections of population lineages through time and the splits represent population lineage splits. This is true regardless of whether the tips of the trees are themselves populations, or are species or higher taxa. Although this conclusion might be paired naturally with a view that species must be monophyletic groups, this population-centric view of trees is independent of that view of species. If we still want to have species that are paraphyletic groups of populations, this is permissible as long as we also do not treat species as the units of phylogeny. This population-centric view opposes a species-centric view of phylogeny and might be called a “rank-free” approach since it entails that we do not need to determine which groups are species (which is partly a ranking question) in order to build a tree. This conclusion and the argument for it are meant to be consistent with, but not require, acceptance of the conclusions of Velasco (2008) regarding species. (shrink)

In recent years, the research field of the evolution of art has witnessed contributions from a wide range of disciplines across the "three cultures". In this thesis, I make both a critical review of existing explanations, and try to do elucidate the evolution of art by employing insights, methods and concepts from different disciplines. First, I critically evaluate the evidentiary criteria from standard evolutionary psychology some accounts employ to demonstrate that art qualifies as a human biological adaptation. I argue (...) that these criteria do not suffice to make that claim. Furthermore, I make the case for a cultural evolutionary approach to the arts. One commonly used evidentiary criterion for art as adaptation is the “tight fit” between artnbsp;human cognition, which is interpreted as suggesting that cognition has undergone selection to produce and appreciate art. However, disentangling cultural and genetic evolution allows formulating thenbsp;hypothesis that art has evolved culturally to match human cognition. Second, I observe that in ancient and modern hunter gatherer societies, artistic activities are virtually alwaysnbsp;with ritual. I discuss what this implies for the adaptive significance of art. Specifically, if the arts have culturally evolved in function of ritualistic purposes, their adaptive value has depended on the adaptiveness of ritual. Third, because architecture is characterized by the combination of utilitarian and aesthetic aspects, it is particularly interesting from an evolutionary perspective and therefore I devote special attention to it. Employing the cross-species comparative approach, I investigate two main purposes human architecture has in common with nonhuman animal constructions: protection andnbsp;Based on the phylogenetic approach I establish that protection may have been the primary function of building aptitudes in humannbsp;and that building was later co-opted for signaling purposes. Subsequently, I comparatively evaluate thenbsp;of signaling models in the cultural evolution of architectural aesthetics. Furthermore I combine insights from environmental psychology, niche construction and cultural evolution theory to develop the hypothesis that a function of religious monumental architecture may have been to support nbsp;or even to galvanize nbsp;social learning. Finally, I connect the account I developed to the cognitive science of religion. I complement the frequently held view that religious monumental architecture is a costly signal with the mechanism of sensory exploitation. I attempt to demonstrate that by exploiting thenbsp;response of awe in religious followers, religious monumental architecture promotes and regulates prosocial behavior and creates in them an openness to adopt supernatural beliefs. Fourth, I dealnbsp;the evolution of visual art, paying again special attention to cultural evolution andnbsp;explanatory value of sensory exploitation. I assert that the relatively late appearance of figurative art in the course of human evolution may be attributed to an increase in the number of social learning opportunities within and between human populations at that time. I support this assertion with archaeological data that indicate overall population growth and increased population densities at that time. These demographic changes, I argue, have allowed the retention and accumulation of innovations required for figurative art making. Contrary to previous accounts, I furthermore aim to show that this process doesnbsp;require figurative art to have served adaptive purposes. Employing sensory exploitation, I establish that figurative art could have evolved by exploiting pre-existing biases of evolved psychology. The dominant themes of upper Paleolithic and late stone age figurative art, animals and humans, seem to support this contention. Fifth, I present a number of experimental studies I conducted to verify whether contemporary art experts would have socially learned to resist exploitation of evolved psychology. One aspect ofrsquo;s exploitative power may be that it elicits aesthetic pleasure by pushing “pleasure buttons” that evolved for other purposes. Consequently, spectators may trade-off rewards from indulging in art and biological activities, resulting in less effort being allocated to reproduction. I contend that, while the generalnbsp; may indeed be exploited, experts, who are typically being exposed to high doses of art, may have socially learned to resist exploitation by selectively preferring art from prestigious art contexts. The latter would be in line with the claim that prestige bias may trump content biases of evolved psychology. The results of three experimental studies supportnbsp;contention. I find that laypeople's art appreciation is positively affected by a content bias for attractive faces, mediated by aesthetic pleasure, whereas experts' appreciation is positively affected by prestige and mediated by admiration for the artist. Moreover, experts confer lower appreciation to attractive compared to moderately attractive content, which is consistentnbsp;the contention that expertise and the use of prestige are associatednbsp;resistance against beautiful content that exploits evolved preferences. This research thus suggests that expertise moderates content and context biases, which may be of relevance to an ongoing debate about their relative importance as drivers of cultural evolution. In addition, this study provides a tentative but novel explanation for the fact, famously established by philosopher Arthur Danto, that beauty lost its central position in Western art during the 20th century. However my findings cannot exclude alternative explanations for the fact that art expert appreciation deviates from evolved aesthetic preferences. Further research may experimentally assess the role of prestige bias and resistance as drivers of this deviation versus other, previously suggested, potential processes. For example, preferences of experts may deviate on account of the fact that art serves as a badge of elite group membership and thus expert art appreciation as a means to distinguish oneself from the masses. Alternatively, art experts may seek intellectual challenges instead of pushing pleasure buttons. Future research may experimentally verify the respective roles of these different hypothesized mechanisms. In addition, it may be investigated how insights from sexual selection research may help elucidating the different processes at play in cultural evolution and particularly in the cultural evolution of art appreciation, given the commonalities between sexual selection and cultural selection models. (shrink)

Some plausibly necessary a posteriori theoretical claims include ‘water is H 2 O’, ‘gold is the element with atomic number 79’, and ‘cats are animals’. In this paper I challenge the necessity of the third claim. I argue that there are possible worlds in which cats exist, but are not animals. Under any of the species concepts currently accepted in biology, organisms do not belong essentially to their species. This is equally true of their ancestors. In (...) class='Hi'>phylogenetic systematics, monophyletic clades such as the animal kingdom are composed of an ancestral stem species and all of its descendants. If the stem species had not existed, neither would the clade. Thus it could have been the case that all the organisms which actually belong to the animal kingdom might have existed yet not have been animals. (shrink)

The recent conception of biodiversity proposed by James Maclaurin and Sterelny was developed mostly with macrobiological life in mind. They suggest that we measure biodiversity by dividing life into natural units (typically species) and quantifying the differences among units using phenetic rather than phylogenetic measures of distance. They identify problems in implementing quantitative phylogenetic notions of difference for non-prokaryotic species. I suggest that if we focus on microbiological life forms that engage in frequent, promiscuous lateral gene (...) transfer (LGT), and their associated reticulated phylogenies, we need to rethink the notion of species as the natural unit, and we discover additional problems with phylogenetic notions of distance. These problems suggest that a phenetic approach based on morphospaces has just as much appeal, if not more, for microbes as they do for multi-cellular life. Facts about LGT, however, offer no new insight into the additional challenge of reconciling units and differences into a single measure of biodiversity. (shrink)

Philosophical discussions of species have focused on multicellular, sexual animals and have often neglected to consider unicellular organisms like bacteria. This article begins to fill this gap by considering what species concepts, if any, apply neatly to the bacterial world. First, I argue that the biological species concept cannot be applied to bacteria because of the variable rates of genetic transfer between populations, depending in part on which gene type is prioritized. Second, I present a critique (...) of phylogenetic bacterial species, arguing that phylogenetic bacterial classification requires a questionable metaphysical commitment to the existence of essential genes. I conclude by considering how microbiologists have dealt with these biological complexities by using more pragmatic and not exclusively evolutionary accounts of species. I argue that this pragmatism is not borne of laziness but rather of the substantial conceptual problems in classifying bacteria based on any evolutionary standard. (shrink)

Biodiversity is a key concept in the biological sciences. While it has its origin in conservation biology, it has become useful across multiple biological disciplines as a means to describe biological variation. It remains, however, unclear what particular biological units the concept refers to. There are currently multiple accounts of which biological features constitute biodiversity and how these are to be measured. In this paper, I draw from the species concept debate to argue for a set of desiderata for (...) the concept of “biodiversity” that is both principled and coheres with the concept’s use. Given these desiderata, this concept should be understood as referring to difference quantified in terms of the phylogenetic structure of lineages, also known as the ‘tree of life’. (shrink)

Cognitive biases can lead to misinterpretations of human and non-human biology and behavior. The concept of the Umwelt describes phylogenetic contrasts in the sensory realms of different species and has important implications for evolutionary studies of cognition (including biases) and social behavior. It has recently been suggested that the microbiome (the diverse network of microorganisms in a given environment, including those within a host organism such as humans) has an influential role in host behavior and health. In this (...) paper, we discuss the host’s microbiome in relation to cognitive biases and the concept of the Umwelt. Failing to consider the role of host–microbiome (collectively termed a “holobiont”) interactions in a given behavior, may underpin a potentially important cognitive bias – which we refer to as the Holobiont Blindspot. We also suggest that microbially mediated behavioral responses could augment our understanding of the Umwelt. For example, the potential role of the microbiome in perception and action could be an important component of the system that gives rise to the Umwelt. We also discuss whether microbial symbionts could be considered in System 1 thinking – that is, decisions driven by perception, intuition and associative memory. Recognizing Holobiont Blindspots and considering the microbiome as a key factor in the Umwelt and System 1 thinking has the potential to advance studies of cognition. Furthermore, investigating Holobiont Blindspots could have important implications for our understanding of social behaviors and mental health. Indeed, the way we think about how we think may need to be revisited. (shrink)

This chapter defends a pluralist understanding of species on which a normative species concept is viable and can support natural goodness evaluations. The central question here is thus: Since organisms are to be evaluated as members of their species, how does a proper understanding of species affect the feasibility of natural goodness evaluations? Philippa Foot has argued for a form of natural goodness evaluation in which living things are evaluated by how well fitted they are for (...) flourishing as members of their species, in ways characteristic of their species. She has further argued that assessments of moral goodness in humans are of the same evaluative form. However, natural goodness evaluations and, by extension, the natural goodness approach, do not garner justification in virtue of employing a scientifically privileged conception of species. The natural goodness approach is only justified given particular metaethical and normative commitments that are independent of naturalism, since the approach does not depend upon naturalism alone. (shrink)

Species are the basis of the taxonomic scheme. They are the lowest taxonomic category that are used as units for describing biodiversity and evolution. In this contribution we discuss the current species concept for prokaryotes. Such organisms are considered to represent the widest diversity among living organisms. Species is currently circumscribed as follows: A prokaryotic species is a category that circumscribes a (preferably) genomically coherent group of individual isolates/strains sharing a high degree of similarity in (many) (...) independent features, comparatively tested under highly standardized conditions. Although the number of described prokaryotic species is underrepresented in the living world, this phylo-phenetic or polythetic species concept currently in use is considered to be pragmatic, operational and universally applicable and successfully used for identification processes. (shrink)

The species concept is one of the oldest and most fundamental in biology. And yet it is almost universally conceded that no satisfactory definition of what constitutes a species has ever been proposed. The present article is devoted to an attempt to review the status of the problem from a methodological point of view. Since the species is one of the many taxonomic categories, the question of the nature of these categories in general needs to be entered (...) into. (shrink)

The concept that renders morphology a tool for phylogeny reconstruction is homology. The concept of homology is rooted in pre-evolutionary idealistic morphology. The claim that the goal of idealistic morphology was the seriability of form may sound paradoxical given that this discipline proceeded within a framework of strictly delimited types. But the types only demarcate where seriability starts and where it comes to an end. Carl Gegenbaur’s was recognized as a milestone in idealistic morphology. A comparison with the second edition (...) of 1870 illustrates Gegenbaur’s turn to evolutionary morphology. The methodology remained the same–seriability of form–but the series was no longer merely descriptive or conceptual but now a historical, evolutionary one. Gegenbaur emphasized that seriability of form was possible not only between species of the same type, but also between parts of organisms of the same type. Pursuing this project, he found that different parts of organisms evolve at different rates, resulting in an incongruence between the series of parts relative to the series of species under comparison. This incongrence was called chevauchement des spécialisations by Louis Dollo, Spezialisationskreuzungen by Othenio Abel, and heterobathmy of characters by Armen Takhtajan. Willi Hennig, the founder of modern methods in phylogenetic systematics, discovered that the heterobathmy of characters was a precondition for the establishment of the phylogenetic relationships based on shared derived characters. The result was a replacement of the search for ancestors by a search for relative degrees of phylogenetic relationships. (shrink)