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)

Speciation is an aspect of evolutionary biology that has received little philosophical attention apart from articles mainly by biologists such as Mayr (1988). The role of speciation as a terminus a quo for the individuality of species or in the context of punctuated equilibrium theory has been discussed, but not the nature of speciation events themselves. It is the task of this paper to attempt to bring speciation events into some kind of general scheme, based primarily upon the work (...) of Sergey Gavrilets on adaptive landscapes, using migration rate, or gene flow, as the primary scale, and concluding that adaptive and drift explanations are complementary rather than competing. I propose a distinction between intrinsic and extrinsic selection, and the notion of reproductive reach and argue that speciation modes should be discriminated in terms of gene flow, the nature of selection maintaining reproductive reach, and whether the predominant cause is selective or stochastic. I also suggest that the notion of an adaptive “quasispecies” for asexual species is the primitive notion of species, and that members of reproductively coherent sexual species are additionally coadapted to their mating partners. (shrink)

The formal definition of species as explanatory hypotheses presented by Fitzhugh is emended. A species is an explanatory account of the occurrences of the same character among gonochoristic or cross-fertilizing hermaphroditic individuals by way of character origin and subsequent fixation during tokogeny. In addition to species, biological systematics also employs hypotheses that are ontogenetic, tokogenetic, intraspecific, and phylogenetic, each of which provides explanatory hypotheses for distinctly different classes of causal questions. It is suggested that species hypotheses (...) can not be applied to organisms with obligate asexual, parthenogenetic, and self-fertilizing modes of reproduction. Hypotheses explaining shared characters among such organisms are, instead, strictly phylogenetic. Several implications of this emended definition are examined, especially the relations between species, intraspecific, and phylogenetic hypotheses, as well as the limitations of species names to be applied to temporally different characters within populations. (shrink)

Species concepts for bacteria and other microbes are contentious, because they are often asexual. There is a Problem of Homogeneity: every mutation in an asexual lineage forms a new strain, of which all descendents are clones until a new mutation occurs. We should expect that asexual organisms would form a smear or continuum. What causes the internal homogeneity of asexual lineages, if they are in fact homogeneous? Is there a natural “species concept” for “microbes”? (...) Two main concepts devised for metazoans and metaphytes have been applied to bacteria. One is the Recombination Concept, a revised form of the Biological Species Concept in which the homogenizing mechanism is the sharing of genome fragments, somewhat akin to sexual recombination. The other is the Ecological Species Concept, in which the ecological niche is that which maintains lineages as cohesive. In this paper I will discuss these two concepts, and offer an underlying model that conjoins them, and consider the implications for species concepts in general. In short, my argument is that asexual species are instances of the most primitive and underived notion of species, which I will call “quasispecies”, following Eigen, and that sexual species are merely one derived kind of species. Moreover, I will argue that there is a continuum of recombination from simple viral models in which each strain is a clone, through to obligate recombination of 50% of the parents’ genome, and that consequently there is no sharp division between “microbial” and more familiar species. (shrink)

The biological species (biospecies) concept applies only to sexually reproducing species, which means that until sexual reproduction evolved, there were no biospecies. On the universal tree of life, biospecies concepts therefore apply only to a relatively small number of clades, notably plants andanimals. I argue that it is useful to treat the various ways of being a species (species modes) as traits of clades. By extension from biospecies to the other concepts intended to capture the natural (...) realities of what keeps taxa distinct, we can treat other modes as traits also, and so come to understand that theplurality of species concepts reflects the biological realities of monophyletic groups.We should expect that specialists in different organisms will tend to favour those concepts that best represent the intrinsic mechanisms that keep taxa distinct in their clades. I will address the question whether modes ofreproduction such as asexual and sexual reproduction are natural classes, given that they are paraphyletic in most clades. (shrink)

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)

Sexual reproduction is almost ubiquitous among multicellular organisms even though it entails severe fitness costs. To resolve this apparent paradox, an extensive body of research has been devoted to identifying the selective advantages of recombination that counteract these costs. Yet, how easy is it to make the transition to asexual reproduction once sexual reproduction has been established for a long time? The present review approaches this question by considering factors that impede the evolution of parthenogenesis in animals. Most importantly, (...) eggs need a diploid chromosome set in most species in order to develop normally. Next, eggs may need to be activated by sperm, and sperm may also contribute centrioles and other paternal factors to the zygote. Depending on how diploidy is achieved mechanistically, further problems may arise in offspring that stem from ‘inbreeding depression’ or inappropriate sex determination systems. Finally, genomic imprinting is another well-known barrier to the evolution of asexuality in mammals. Studies on species with occasional, deficient parthenogenesis indicate that the relative importance of these constraints may vary widely. The intimate evolutionary relations between haplodiploidy and parthenogenesis as well as implications for the clade selection hypothesis of the maintenance of sexual reproduction are also discussed. (shrink)

The genomes of virtually all sexually reproducing species contain transposable elements. Although active elements generally transpose more rapidly than they are inactivated by mutation or excision, their number can be kept in check by purifying selection if its effectiveness becomes disproportionately greater as their copy number increases. In sexually reproducing species, such synergistic selection can result from ectopic crossing-over or from homologous recombination under negative epistasis. In addition, there may be controls on transposon activity that are associated with (...) meiosis. Because a sexual lineage that abandons sex must lack such mechanisms, it may be driven to extinction by the unchecked proliferation of deleterious transposons inherited from its sexual progenitor. An important component of the evolutionary advantage of sex over asex may therefore lie in the ability of sex, despite facilitating the spread of deleterious elements within interbreeding populations, also to restrain their intragenomic proliferation. BioEssays 27:76–85, 2005. © 2004 Wiley Periodicals, Inc. (shrink)

This article examines the critique of the biological species concept advanced by protozoan geneticist Tracy Sonneborn at the 1955 AAAS symposium on "the species problem," published subsequently in 1957. Although Sonneborn was a strong proponent of a population genetical conception of species, he became critical of the biological species concept for its failure to incorporate asexual and obligatory inbreeding organisms. It is argued that Sonneborn's intimate knowledge of the ciliate protozoan Paramecium aurelia species complex (...) brought him into conflict with a growing pressure in the biological sciences to emphasize universal principles of life. Faced with the need to defend the value of P. aurelia as an investigative tool, Sonneborn argued that the sharp break in nature between sexual and asexual organisms posited by proponents of the biological species concept was not an existential feature of the living world, but rather the misleading consequence of an operational definition of species based only upon sexual organisms. Drawing upon his knowledge of the immense variability of P. aurelia, he proposed instead a continuum of breeding systems from obligatory outbreeding to asexual organisms, and a more broadly unifying definition of species that incorporated asexual as well as sexual organisms. Paradoxically, the push for unification that then characterized the evolutionary synthesis served to debar critical consideration of Sonneborn's more unificatory alternative, and his underlying contention that biological anomaly could serve as an important source of conceptual unification. (shrink)

Recent investigations with non‐model species and whole‐genome approaches have challenged several paradigms in animal epigenetics. They revealed that epigenetic variation in populations is not the mere consequence of genetic variation, but is a semi‐independent or independent source of phenotypic variation, depending on mode of reproduction. DNA methylation is not positively correlated with genome size and phylogenetic position as earlier believed, but has evolved differently between and within higher taxa. Epigenetic marks are usually not completely erased in the zygote and (...) germ cells as generalized from mouse, but often persist and can be transgenerationally inherited, making them evolutionarily relevant. Gene body methylation and promoter methylation are similar in vertebrates and invertebrates with well methylated genomes but transposon silencing through methylation is variable. The new data also suggest that animals use epigenetic mechanisms to cope with rapid environmental changes and to adapt to new environments. The main benefiters are asexual populations, invaders, sessile taxa and long‐lived species. (shrink)

Can an AGI create a more intelligent AGI? Under idealized assumptions, for a certain theoretical type of intelligence, our answer is: “Not without outside help”. This is a paper on the mathematical structure of AGI populations when parent AGIs create child AGIs. We argue that such populations satisfy a certain biological law. Motivated by observations of sexual reproduction in seemingly-asexual species, the Knight-Darwin Law states that it is impossible for one organism to asexually produce another, which asexually produces (...) another, and so on forever: that any sequence of organisms (each one a child of the previous) must contain occasional multi-parent organisms, or must terminate. By proving that a certain measure (arguably an intelligence measure) decreases when an idealized parent AGI single-handedly creates a child AGI, we argue that a similar Law holds for AGIs. (shrink)

Here, I propose a new hypothesis: sex originated from an archaic gene transfer process among prebiotic bubbles without the prerequisite for reproduction. This de‐coupling from reproduction might make the thorny problem of accounting for the evolution of sex, despite the apparent advantages of parthenogenicity, more tractable.

Under the influence of deleterious mutation and selection a population will reach equilibrium and contain individuals with [0, 1, 2 - - mutations.] This deterministic equilibrium distribution is exactly the same for asexual and sexual populations. The size of the optimal class , i.e. the class with the smallest number of mutations, is determined by the genome mutation rate and the average selective disadvantage of the mutations. A large U and small s gives a very small n o. If (...) n o is small in an asexual population it will be lost by drift and this causes a reduction in the mean fitness of the population . It is argued that diploidy increases U and reduces s. Values of U and s observed from Drosophila indicate a diploid would have a vanishingly small n o. The argument suggests Muller's ratchet is more powerful than previously accepted in asexual species derived from diploid sexual ancestors. (shrink)

Contemporary biology has inherited two key assumptions from the Modern Synthesis about the nature of population lineages: sexual reproduction is the exemplar for how individuals in population lineages inherit traits from their parents, and random mating is the exemplar for reproductive interaction. While these assumptions have been extremely fruitful for a number of fields, such as population genetics and phylogenetics, they are increasingly unviable for studying the full diversity and evolution of life. I introduce the “mixture” account of population lineages (...) that escapes these assumptions by dissolving the Modern Synthesis’s sharp line separating reproduction and development and characterizing reproductive integration in population lineages by the ephemerality of isolated subgroups rather than random mating. The mixture account provides a single criterion for reproductive integration that accommodates both sexual and asexual reproduction, unifying their treatment under Kevin de Queiroz’s generalized lineage concept of species. The account also provides a new basis for empirically assessing the effect of random mating as an idealization on the empirical adequacy of population genetic models. (shrink)

All species continuously evolve to adapt to changing environments. The genetic variation that fosters such adaptation is caused by a plethora of mechanisms, including meiotic recombination that generates novel allelic combinations in the progeny of two parental lineages. However, a considerable number of eukaryotic species, including many fungi, do not have an apparent sexual cycle and are consequently thought to be limited in their evolutionary potential. As such organisms are expected to have reduced capability to eliminate deleterious mutations, (...) they are often considered as evolutionary dead ends. However, inspired by recent reports we argue that such organisms can be as persistent as organisms with conventional sexual cycles through the use of other mechanisms, such as genomic rearrangements, to foster adaptation. (shrink)

“Because those who come after us may not be like us, or because those like us may not come after us, or because after a time there may be none to come after us, mankind must now set to work to insure that those who come after us will be more unlike us. In this there is at work the modern intellect’s penchant for species suicide.” With these words Paul Ramsey brings to a conclusion his provocative and surprising study (...) of the problems we are encountering – and will encounter – as scientific advances make the genetic control of man a real possibility. Now, increased knowledge of chromosome structure has taken the alteration of the human race out of the realm of science fiction and placed it in the hands of laboratory scientists. Ramsey sees this advance in science and technology as a mixed blessing. He examines the ramifications of genetic control, not only the now common measures of birth control and artificial insemination, but also the effect on the genetic pool of the increasing numbers of genetically poor individuals. His concern for humanistic values informs each argument as he tackles such issues as asexual reproduction of men, frozen semen banks, and the breeding of human beings for special purposes. No less than the future of man as we know him hangs on these issues. One must pay particular attention to this book. (shrink)

Arguments for or against a trend in the evolution of complexity are weakened by the lack of an unambiguous definition of complexity. Such definitions abound for both dynamical systems and biological organisms, but have drawbacks of either a conceptual or a practical nature. Physical complexity, a measure based on automata theory and information theory, is a simple and intuitive measure of the amount of information that an organism stores, in its genome, about the environment in which it evolves. It is (...) argued that physical complexity must increase in molecular evolution of asexual organisms in a single niche if the environment does not change, due to natural selection. It is possible that complexity decreases in co‐evolving systems as well as at high mutation rates, in sexual populations, and in time‐dependent landscapes. However, it is reasoned that these factors usually help, rather than hinder, the evolution of complexity, and that a theory of physical complexity for co‐evolving species will reveal an overall trend towards higher complexity in biological evolution. BioEssays 24:1085–1094, 2002. © 2002 Wiley‐Periodicals, Inc. (shrink)

The present work is a contribution to the understanding of the sempiternal problem of the “burden of factor two” implied by sexual reproduction versus asexual one, as males are energy consumers not contributing to the production of offspring. We construct a deterministic mathematical model in population dynamics where a species enjoys both sexual and parthenogenetic capabilities of reproduction and lives on a limited resource. We then show how polygamy implies instability of a parthenogenetic population with a small number (...) of sexually born males. This instability implies evolution of the system towards an attractor involving both populations. We also exhibit the analogy with a parasite/host system. (shrink)

Discussions of the implications of sexual reproduction have appeared throughout the history of evolutionary biology, from Darwin to Weismann, Fisher, Muller, Maynard Smith, and Williams. The latest of these appearances highlighted an evolutionary paradox that had previously been overlooked. In many animal and plant species reproduction is obligately sexual and also half the offspring are male, yet the males contribute nothing but genes to reproduction. If asexual mutants of such a species were to produce as many (...) class='Hi'>asexual offspring on average as sexual females in that species produce daughters and sons, the growth rate of their population would be twice as large. Whereas half the offspring of sexual females are noncontributory males, all the offspring of such an asexual mutant would contribute to growth rate. Nevertheless, sexual reproduction prevails in most species and is not lost evolutionarily. (shrink)

It has become increasingly evident that gene content of the sex chromosomes is markedly different from that of the autosomes. Both sex chromosomes appear enriched for genes related to sexual differentiation and reproduction; but curiously, the human X chromosome also seems to bear a preponderance of genes linked to brain and muscle functions. In this review, we will synthesize several evolutionary theories that may account for this nonrandom assortment of genes on the sex chromosomes, including 1) asexual degeneration, 2) (...) sexual antagonism, 3) constant selection, and 4) hemizygous exposure. Additionally, we will speculate on how the evolution of sex‐chromosome gene content might have impacted on the phenotypic evolution of mammals and particularly humans. Our discussion will focus on the mammalian sex chromosomes, but will cross reference other species where appropriate. BioEssays 26:159–169, 2004. © 2004 Wiley Periodicals, Inc. (shrink)

This essay explores how the human biological self is socially constructed, and rejects various truisms that define our character. Rather than being stand-alone entities, the human biological self forms what biologists call “superorganisms” and what I call “poly-super-organisms.” Thus, along with prokaryotes , viruses, and other entities, we are combined in an inseparable menagerie of species that is spread across multiple bodies. Biologists claim that only males and females are organisms. As described here, however, human sperm and eggs are (...) equally entitled to that rank. Much about human reproduction is also socially constructed. For example, contrary to scientific wisdom, humans have always reproduced both sexually and asexually. Moreover, human life does not begin between conception and birth, and neither event creates new life. In addition, some cancer cells are organisms; they are contagious entities formed by asexual human reproduction. Finally, while science universally describes humans as being living entities, accumulating evidence makes clear that the life concept is failing. In fact, much that originates from human bodies does not fall under current biological or cultural classifications.How we construct the human biological self has consequences for understanding our health, how we reproduce, and what we are. These social constructions greatly influence our views on abortion, stem-cell research, human cloning, and our post-human future. A more accurate construction of the human biological self will help us better navigate these contentious issues. (shrink)

Supernumerary B chromosomes are dispensable elements of the genome which can be retained in populations at high frequencies, despite being deleterious, through the ability to undergo non‐Mendelian inheritance. Their mode of origin is, however, obscure. Recent work on gynogenetic fish has demonstrated the incorporation of small, unstable, centromere‐containing microchromosomes, probably of interspecific derivation, into an asexual lineage(1). That these resemble B chromosomes both in structure and behaviour is consistent with the proposal that hybridisation between closely related species may (...) be a significant mode of origin for such selfish genetic elements. Additional work on the B chromosome of a parasitoid wasp and observations on patterns of chromosome breakage from somatic cell hybrids also support this hypothesis. (shrink)

Alternative ways to ensure mate compatibility, such as hermaphroditism and the breakdown of self‐incompatibility, evolved repeatedly when finding a mating partner is difficult. In a variety of microorganisms where compatibility is determined by mating‐types, a highly regulated form of universal compatibility system called mating‐type switching has evolved several times. This sophisticated system allows for the genetic adjustment of the mating type during asexual growth, and it most likely evolved for reproductive assurance of immotile species under low densities. In (...) this review, we compare the switching strategy to other universal compatibility systems such as “unisexual mating” and homothallism. We identify the costs of switching, including genome instability, and mechanistic costs, as well as the benefits, mainly the maintenance of important mating‐type functions. Given the potential benefits of mating‐type switching, we speculate that switching is likely to have evolved many times independently, and may be more common in groups where genetic mating types regulate mate compatibility than assumed so far. (shrink)

The study of alternative genetic systems and mixed modes of reproduction, whereby sexual and asexual reproduction is combined within the same lifecycle, is of fundamental importance as they may shed light on classical evolutionary issues, such as the paradox of sex. Recently, several such cases were discovered in social insects. A closer examination of these systems has revealed many amazing facts, including the mixed use of asexual and sexual reproduction for the production of new queens and workers, males (...) that can clone themselves and the routine use of incest without deleterious genetic consequences. In addition, in several species, remarkable cases of asexually reproducing socially parasitic worker lineages have been discovered. The study of these unusual systems promises to provide insight into many basic evolutionary questions, including the maintenance of sex, the expression of sexual conflict and kin conflict and the evolution of cheating in asexual lineages. (shrink)

Asexuality is overlooked in the philosophical literature and in wider society. Such neglect produces incomplete or inaccurate accounts of romantic life and harms asexual people. We develop an account of asexuality to redress this neglect and enrich discussion of romantic life. Asexual experiences are diverse. Some asexual people have sex; some have romantic relationships in the absence of sex. We accept the common definition of asexuality as the absence of sexual attraction and explain how sexual attraction and (...) sexual desire differ by giving an affordance‐like account of sexual attraction. Armed with that distinction, we show that asexuality is clearly different from celibacy or disorders of desire and that some existing philosophical theories of sexual desire struggle to accommodate asexual sexuality. We then build on asexual testimony about the diversity of non‐sexual attractions to answer two common objections levelled at asexual romance: that romantic relationships require sexual attraction or that sex in the absence of sexual attraction is insufficiently focused on someone as an individual. Finally, we describe some of the ways asexuality has been erased or denigrated in society, and the specific injustices and harms that result. (shrink)

In this essay, we aim to provide an overview of the political and philosophical issues pertaining to asexuality. The first section, “What Is Asexuality?,” offers an account of asexuality. The second section, “Asexuality as a Unique Sexual Orientation,” argues that asexuality should be understood as a unique sexual orientation. The third section, “Asexuality and Oppression,” discusses the various forms of oppression facing asexual persons today. The fourth section, “The Goods of Asexuality,” articulates some goods that asexuality brings to human (...) lives, and we do this with an eye toward conceiving of asexuality as something other than a mere deficit. (shrink)

This paper poses a problem for traditional phylogenetics: The identity of organisms that reproduce through fission can be understood in several different ways. This prompts questions about how to differentiate parent organisms from their offspring, making vertical gene transfer unclear. Differentiating between parents and offspring stems from what I call the identity problem. How the problem is resolved has implications for phylogenetic groupings. If the identity of a particular asexual organism persists through fission, the vertical lineage on a phylogenetic (...) tree will split differently than if the identity of an organism does not survive the fission process. (shrink)

Stamos squarely confronts the problem of determining what a biological species is, whether species are real, and the nature of their reality. He critically considers the evolution of the major contemporary views of species and also offers his own solution to the species problem.

The notion that eukaryotes are ancestrally sexual has been gaining attention. This idea comes in part from the discovery of sets of “meiosis‐specific genes” in the genomes of protists. The existence of these genes has persuaded many that these organisms may be engaging in sex, even though this has gone undetected. The involvement of sex in protists is supported by the view that asexual reproduction results in the accumulation of mutations that would inevitably result in the decline and extinction (...) of such lineages. It is argued that this phenomenon can be obviated by polyploidy and that the “meiosis‐specific genes” are used in other processes, including polyploidy control and homologous recombination, independent of meiosis. These phenomena account for the finding that these genes are expressed in cultures devoid of apparent cell fusion events. Hence, it is also proposed that asexual, and not sexual, reproduction is the ancestral condition. (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 (...) class='Hi'>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)

this requirement for adaptations. Emergent characters are always potential adaptations. Not all selection processes produce adaptations, however. The key issue, in delineating a selection process, is the relationship between a character and fitness. The emergent character approach is more restrictive than alternative schemas that delineate selection..

This collection of original essays--by philosophers of biology, biologists, and cognitive scientists--provides a wide range of perspectives on species. Including contributions from David Hull, John Dupre, David Nanney, Kevin de Queiroz, and Kim Sterelny, amongst others, this book has become especially well-known for the three essays it contains on the homeostatic property cluster view of natural kinds, papers by Richard Boyd, Paul Griffiths, and Robert A. Wilson.

What are species? Are they objective features of the world? If so, what sort of features are they? Do everyday intuitions that species are real stand up to philosophical and scientific scrutiny? Two rival accounts of species' reality have dominated the discussion: that species are natural kinds defined by essential properties and that species are individuals. Unfortunately, neither account fully accommodates biological practice. In Are Species Real?, Slater presents a novel approach to this question (...) aimed at accommodating the attractions to both realism and antirealism about species. (shrink)

“When Species Meet is a breathtaking meditation on the intersection between humankind and dog, philosophy and science, and macro and micro cultures.” —Cameron Woo, Publisher of Bark magazine In 2006, about 69 million U.S. households had pets, giving homes to around 73.9 million dogs, 90.5 million cats, and 16.6 million birds, and spending over $38 billion dollars on companion animals. As never before in history, our pets are truly members of the family. But the notion of “companion species”—knotted (...) from human beings, animals and other organisms, landscapes, and technologies—includes much more than “companion animals.” In When Species Meet, Donna J. Haraway digs into this larger phenomenon to contemplate the interactions of humans with many kinds of critters, especially with those called domestic. At the heart of the book are her experiences in agility training with her dogs Cayenne and Roland, but Haraway’s vision here also encompasses wolves, chickens, cats, baboons, sheep, microorganisms, and whales wearing video cameras. From designer pets to lab animals to trained therapy dogs, she deftly explores philosophical, cultural, and biological aspects of animal-human encounters. In this deeply personal yet intellectually groundbreaking work, Haraway develops the idea of companion species, those who meet and break bread together but not without some indigestion. “A great deal is at stake in such meetings,” she writes, “and outcomes are not guaranteed. There is no assured happy or unhappy ending—socially, ecologically, or scientifically. There is only the chance for getting on together with some grace.” Ultimately, she finds that respect, curiosity, and knowledge spring from animal-human associations and work powerfully against ideas about human exceptionalism. One of the founders of the posthumanities, Donna J. Haraway is professor in the History of Consciousness Department at the University of California, Santa Cruz. Author of many books and widely read essays, including The Companion Species Manifesto: Dogs, People, and Significant Otherness and the now-classic essay “The Cyborg Manifesto,” she received the J. D. Bernal Prize in 2000, a lifetime achievement award from the Society for Social Studies in Science. (shrink)

There is long-standing disagreement among systematists about how to divide biodiversity into species. Over twenty different species concepts are used to group organisms, according to criteria as diverse as morphological or molecular similarity, interbreeding and genealogical relationships. This, combined with the implications of evolutionary biology, raises the worry that either there is no single kind of species, or that species are not real. This book surveys the history of thinking about species from Aristotle to modern (...) systematics in order to understand the origin of the problem, and advocates a solution based on the idea of the division of conceptual labor, whereby species concepts function in different ways - theoretically and operationally. It also considers related topics such as individuality and the metaphysics of evolution, and how scientific terms get their meaning. This important addition to the current debate will be essential for philosophers and historians of science, and for biologists. (shrink)

The so-called ‘‘species problem’’ has plagued evolution- ary biology since before Darwin’s publication of the aptly titled Origin of Species. Many biologists think the problem is just a matter of semantics; others complain that it will not be solved until we have more empirical data. Yet, we don’t seem to be able to escape discussing it and teaching seminars about it. In this paper, I briefly examine the main themes of the biological and philosophical liter- atures on the (...) species problem, focusing on identifying common threads as well as relevant differences. I then argue two fundamental points. First, the species problem is not primarily an empirical one, but it is rather fraught with philosophical questions that require—but cannot be settled by—empirical evidence. Second, the (dis-)solution lies in explicitly adopting Wittgenstein’s idea of ‘‘family resemblance’’ or cluster concepts, and to consider spe- cies as an example of such concepts. This solution has several attractive features, including bringing together apparently diverging themes of discussion among bio- logists and philosophers. The current proposal is con- ceptually independent (though not incompatible) with the pluralist approach to the species problem advocated by Mishler, Donoghue, Kitcher and Dupre ́, which implies that distinct aspects of the species question need to be emphasized depending on the goals of the researcher. From the biological literature, the concept of species that most closely matches the philosophical discussion pre- sented here is Templeton’s cohesion idea. (shrink)

h e Darwinian hypothesis has the merit of being eminently simple and comprehensible in principle, and its essential positions may be stated in a very few words: all species have been produced by the development of varieties from common stocks; by the conversion of these, first into permanent races and then into new species, by the process of natural selection , which process is essentially identical with that artificial selection by which man has originated the races of domestic (...) animals—the struggle for existence taking the place of man, and exerting, in the case of natural selection, that selective action which he performs in artificial selection. (shrink)

This paper aims to offer a new argument in defence bacterial species pluralism. To do so, I shall first present the particular issues derived from the conflict between the non-theoretical understanding of species as units of classification and the theoretical comprehension of them as units of evolution. Secondly, I shall justify the necessity of the concept of species for the bacterial world, and show how medicine and endosymbiotic evolutionary theory make use of different concepts of bacterial (...) class='Hi'>species due to their distinctive purposes. Finally, I shall show how my argument provides a new source of defence for bacterial pluralism. (shrink)

Features Covers the philosophical and historical development of the concept of "species" Documents that variation was recognized by pre-Darwinian scholars Includes a section on the debates since the time of the New Synthesis Better suited to non-philosophers Summary Over time the complex idea of "species" has evolved, yet its meaning is far from resolved. This comprehensive work is a fresh look at an idea central to the field of biology by tracing its history from antiquity to today. (...) class='Hi'>Species is a benchmark exploration and clarification of a concept fundamental to the past, present, and future of the natural sciences. In this edition, a section is added on the debate over species since the time of the New Synthesis, and brings the book up to date. A section on recent philosophical debates over species has also been added. This edition is better suited non-specialists in philosophy, so that it will be of greater use for scientists wishing to understand how the notion came to be that living organisms form species. (shrink)

Auf die im Jahre 1938 entwickelte allgemeine Gleichung derMendel Gesetze bezugnehmend, die die Vererbung als Ausdruck desNewtonschen Binoms hält, sucht der Verfasser dieselbe auf die Geschlechtsvererbung anzuwenden, indem er, sich des transformiertenPascal'schen arithmetischen Dreiecks bedienend, zum Schluss kommt, dass die ungeschlechtliche Vermehrung, vom rein biometrischen Standpunkt aus betrachtet, als Beispiel der Zero-Potenz in dem Kreuzungsrange angesehen werden könnte, d.h. eine komplette Konsolidierung und Homozygotie, bei alledem vielleicht sekundär aus der früher durchgemachten geschlechtlichen Vermehrung entstanden, darstellte.Die geschlechtliche Vermehrung müsste sodann etwa (...) für ein Beispiel des Binoms, wenn auch von nicht reiner, aber deformierter oder verkümmerter Form, von einer Kreuzungspotenzn=1 angesehen werden, und zwar vom Typus .Zudem wird ein weiterer Versuch gemacht, von demselben Standpunkt aus, auch einige kompliziertere Geschlechtserscheinungen zu analysieren, wie die des Geschlechtspolymorphismus , des Hermaphroditismus , so wie auch die oft in den natürlichen Populationen beobachtbaren Verschiebungen in der üblichen Zahlproportion 1 ∶ 1 der Geschlechter .In relation to the general equation of the laws ofMendel, given by the author in 1938, as forming the expression ofNewton's binomial, the author analyses the possibilities of application of this equation to the phenomena of sex inheritance and, while usingPascal's transformed arithmetical triangle, comes to the conclusion that the asexual propagation might be taken, from the purely biometrical point of view, as an instance of Zero-power of the crosses, which would mean a complete consolidation or homozygoty of germ material, acquired, it is true, as a secondary achievement from the previous sexual propagation.The latter, again, ought to be considered as an instance ofNewton's binomial of the cross power ofn=1 of a rather irregular, disfigured type of .Then, the author is making an attempt of analysing, from this point of view, a few more complicated cases from this field of inquiry, such as sexual polymorphism , hermaphroditism and finally the fluctuations in the number proportions of sexes, found in natural populations. (shrink)

For more than fifty years, taxonomists have proposed numerous alternative definitions of species while they searched for a unique, comprehensive, and persuasive definition. This monograph shows that these efforts have been unnecessary, and indeed have provably been a pursuit of a will o’ the wisp because they have failed to recognize the theoretical impossibility of what they seek to accomplish. A clear and rigorous understanding of the logic underlying species definition leads both to a recognition of the inescapable (...) ambiguity that affects the definition of species, and to a framework-relative approach to species definition that is logically compelling, i.e., cannot not be accepted without inconsistency. An appendix reflects upon the conclusions reached, applying them in an intellectually whimsical taxonomic thought experiment that conjectures the possibility of an emerging new human species. (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)

The species category is defined as thesmallest historical individual within which there is a parental pattern of ancestry and descent. The use of historical individual in this definition is consistent with the prevailing notion that speciesper se are not involved in processes — they are effects, not effectors. Reproductive isolation distinguishes biparental historical species from their parts, and it provides a basis for understanding the nature of the evidence used to discover historical individuals.

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

Species pluralism gives us reason to doubt the existence of the species category. The problem is not that species concepts are chosen according to our interests or that pluralism and the desire for hierarchical classifications are incompatible. The problem is that the various taxa we call 'species' lack a common unifying feature.

Some critics of invasion biology have argued the invasion of ecosystems by nonindigenous species can create more valuable ecosystems. They consider invaded communities as more valuable because they potentially produce more ecosystem services. To establish that the introduction of nonindigenous species creates more valuable ecosystems, they defend that value is provisioned by ecosystem services. These services are derived from ecosystem productivity, the production and cycling of resources. Ecosystem productivity is a result of biodiversity, which is understood as local (...) species richness. Invasive species increase local species richness and, therefore, increase the conservation value of local ecosystems. These views are disseminating to the public via a series of popular science books. Conservationists must respond to these views, and I outline a method of rejecting such arguments against controlling invasive species. Ecological systems are valuable for more than local productivity and biodiversity is not accurately described by a local species count. (shrink)

If ecological systems are functionally organised, they can possess functions or malfunctions. Natural function would provide justification for conservationists to act for the protection of current ecological arrangements and control the presence of populations that create ecosystem malfunctions. Invasive species are often thought to be malfunctional for ecosystems, so functional arrangement would provide an objective reason for their control. Unfortunately for this prospect, I argue no theory of function, which can support such normative conclusions, can be applied to large (...) scale ecosystems. Instead ecological systems have causal structure, with small clusters of populations achieving functional arrangement. This, however, does not leave us without reason to control invasive species. We can look at the causal arrangement of ecological systems for populations that support ecological features that we should preserve. Populations that play a causal role in reducing biodiversity should be controlled, because biodiversity is a good all prudent agents should want to preserve. (shrink)

In the Preface of Animal Species and Evolution (1963), I wrote that it was "an attempt to summarize and review critically what we know about the biology and genetics of animal species and their role in evolution." The result was a volume of XIV ...