Here is a working list of species concepts presently in play. I quote “Concepts” above because, for philosophical reasons, I think there is only one concept – “species”, and all the rest are conceptions, or definitions, of that concept. I have christened this the Synapormorphic Concept of Species in (Wilkins 2003). More under the fold:
A Summary of 26 species concepts
There are numerous species “concepts” (i.e., conceptions of “species”) at the research and practical level in the scientific literature. (Mayden 1997) has listed 22 distinct species concepts along with synonyms, which provides a useful starting point for a review. I have added authors where I can locate them in addition to Mayden’s references, and instead of his abbreviations I have tried to give the concepts names, such as biospecies for Biological Species, etc. (following George 1956), except where nothing natural suggests itself. There have also been several additional concepts since Mayden’s review, which I have added the views of Pleijel and Wu, and several new revisions presented in Wheeler and Meier (2000). I also add some “partial” species concepts – the compilospecies concept and the nothospecies concept. I distinguish between two phylospecies concepts that go by various names, mostly the names of the authors presenting at the time (as in the Wheeler and Meier volume). To remedy this terminological inflation, I have christened them the Autapomorphic species concept and the Phylogenetic Taxon species concept.
Synonyms: Microspecies, paraspecies, pseudospecies, semispecies, quasispecies, genomospecies (for prokaryotes Euzéby 2006) Principal authors: Cain (1954), Eigen (1993, for quasispecies) Specifications: Asexual lineages, uniparental organisms (parthenogens and apomicts), that cluster together in terms of their genome. May be secondarily uniparental from biparental ancestors.
2. Autapomorphic species
See: Phylospecies Principal authors: Nelson and Platnick (1981); Rosen (1979) Specification: A geographically constrained group of individuals with some unique apomorphous characters, the unit of evolutionary significance (Rosen); simply the smallest detected samples of self-perpetuating organisms that have unique sets of characters (Nelson and Platnick); the smallest aggregation of (sexual) populations or (asexual) lineages diagnosable by a unique combination of character traits (Wheeler and Platnick 2000).
Synonyms: Syngen, speciationist species concept Related concepts: Biological species concept, Genetic species, isolation species Principal authors: John Ray, Buffon, Dobzhansky (1935); Mayr (1942) Specifications: Inclusive Mendelian population of sexually reproducing organisms (Dobzhansky 1935, 1937, 1970), interbreeding natural population isolated from other such groups (Mayr 1942, 1963, 1970; Mayr and Ashlock 1991). Depends upon endogenous reproductive isolating mechanisms (RIMs).
Synonyms: Internodal species concept, Hennigian species concept, Hennigian convention Principal authors: Hennig (1966; 1950); Kornet (1993) Specifications: Set of organisms between speciation events or between speciation event and extinction (Ridley 1989), a segment of a phylogenetic lineage between nodes. Upon speciation the ancestral species is extinguished and two new species are named.
5. Cohesion species
Synonyms: Cohesive individual (in part) (Ghiselin and Hull) Principal authors: Templeton (1989) Specifications: Evolutionary lineages bounded by cohesion mechanisms that cause reproductive communities, particularly genetic exchange, and ecological interchangeability.
Synonyms: None Related concepts: Introgressive taxa Principal authors: Harlan (1963), Aguilar (1999) Specifications: A species pair where one species “plunders” the genetic resources of another via introgressive interbreeding.
7. Composite Species
Synonyms: Phylospecies (in part), Internodal species (in part), cladospecies (in part) Principal authors: Kornet (1993) Specifications: All organisms belonging to an internodon and its descendents until any subsequent internodon. An internodon is defined as a set of organisms whose parent-child relations are not split (have the INT relation).
Synonyms: Ecotypes Related concepts: Evolutionary species sensu Simpson, Ecological mosaics Principal authors: Simpson (1961); Sterelny (1999); Turesson (1922); Van Valen (1976) Specifications: A lineage (or closely related set of lineages) which occupies an adaptive zone minimally different from that of any other lineage in its range and which evolves separately from all lineages outside its range.
9. Evolutionary species
Synonyms: Unit of evolution, evolutionary group Related concepts: Evolutionary significant unit Principal authors: Simpson (1961); Wiley (1978); (1981) Specifications: A lineage (an ancestral-descendent sequence of populations) evolving separately from others and with its own unitary evolutionary role and tendencies (Simpson).
10. Evolutionary significant unit
Synonyms: Biospecies (in part) and evolutionary species (in part) Principal authors: Waples (1991) Specifications: A population (or group of populations) that (1) is substantially reproductively isolated from other conspecific population units, and (2) represents an important component in the evolutionary legacy of the species.
11. Genealogical concordance species
Synonyms: Biospecies (in part), cladospecies (in part), phylospecies (in part) Principal authors: Avise and Ball (1990) Specifications: Population subdivisions concordantly identified by multiple independent genetic traits constitute the population units worthy of recognition as phylogenetic taxa
12. Genic species
Synonyms: none Related concepts: Genealogical concordance species, genetic species (in part), biospecies (in part), autapomorphic species (in part) Principal author: Wu (2001b; 2001a) Specifications: A species formed by the fixation of all isolating genetic traits in the common genome of the entire population.
13. Genetic species
Synonyms: Gentes (sing. Gens) Related concepts: Biospecies, phenospecies, morphospecies, genomospecies Principal authors: Dobzhansky (1950); Mayr (1969); Simpson (1943) Specifications: Group of organisms that may inherit characters from each other, common gene pool, reproductive community that forms a genetic unit
14. Genotypic cluster
Synonyms: Polythetic species Related concepts: Agamospecies, biospecies, genetic species, Hennigian species, morphospecies, non-dimensional species, phenospecies, autapomorphic phylospecies, successional species, taxonomic species , genomospecies Principal author: Mallet (1995) Specifications: Clusters of monotypic or polytypic biological entities, identified using morphology or genetics, forming groups that have few or no intermediates when in contact.
15. Hennigian species
Synonyms: Biospecies (in part), cladospecies (in part), phylospecies (in part), internodal species Principal authors: Hennig (1966; 1950); Meier and Willman (1997) Specifications: A tokogenetic community that arises when a stem species is dissolved into two new species and ends when it goes extinct or speciates.
16. Internodal species
Synonyms: Cladospecies and Hennigian species (in part), phylospecies Principal author: Kornet (1993) Specifications: Organisms are conspecific in virtue of their common membership of a part of a genealogical network between two permanent splitting events or a splitting event and extinction
17. Least Inclusive Taxonomic Unit (LITUs)
Synonyms: evolutionary group (in part), phylospecies Principal authors: Pleijel (Pleijel 1999; Pleijel and Rouse 2000) Specifications: A taxonomic group that is diagnosable in terms of its autapomorphies, but has no fixed rank or binomial.
Synonyms: Classical species, Linnaean species. Related concepts: Linnean species, binoms, phenospecies, monothetic species, monotypes, types, Taxonomic species Principal authors: Aristotle and Linnaeus, and too many others to name, but including Owen, Agassiz, and recently, Cronquist (1978) Specifications: Species are the smallest groups that are consistently and persistently distinct, and distinguishable by ordinary means (Cronquist). Contrary to the received view, this was never anything more than a diagnostic account of species.
19. Non-dimensional species
Synonyms: Folk taxonomical kinds (Atran 1990) Related concepts: Biospecies, genetic species, morphospecies, paleospecies, successional species, taxonomic species Principal authors: Mayr (1942; 1963) Specifications: Species delimitation in a non-dimensional system (a system without the dimensions of space and time, Mayr 1963)
Synonyms: hybrid species, reticulate species Related concepts: Compilospecies, horizontal or lateral genetic transfer Principal author: Wagner (1983) Specifications: Species formed from the hybridization of two distinct parental species, often by polyploidy.
Synonyms: Autapomorphic phylospecies, monophyletic phylospecies, minimal monophyletic units, monophyletic species, lineages Related concepts: Similar to internodal species cladospecies, composite species, least inclusive taxonomic units. Principal authors: Cracraft (1983); Eldredge and Cracraft (1980); Nelson and Platnick (1981); Rosen (1979) Specifications: The smallest unit appropriate for phylogenetic analysis, the smallest biological entities that are diagnosable and monophyletic, unit product of natural selection and descent. A geographically constrained group with one or more unique apomorphies (autapomorphies). There are two versions of this and they are not identical. One derives from Rosen and is what I call the Autapomorphic species concept. It is primarily a concept of diagnosis and tends to be favoured by the tradition known as pattern cladism. The other is what I call the Phylogenetic Taxon species concept, and tends to be favoured by process cladists.
21. Phylogenetic Taxon species
See: Phylospecies Principal authors: Cracraft (1983); Eldredge and Cracraft (1980); Nixon and Wheeler(1990) Specifications: A species is the smallest diagnosable cluster of individual organisms within which there is a parental pattern of ancestry and descent (Cracraft); the least inclusive taxon recognized in a classification, into which organisms are grouped because of evidence of monophyly (usually, but not restricted to, the presence of synapomorphies), that is ranked as a species because it is the smallest ‘important’ lineage deemed worthy of formal recognition, where ‘important’ refers to the action of those processes that are dominant in producing and maintaining lineages in a particular case (Mishler and Brandon 1987).
Synonyms: Phena (sing. phenon) (Smith 1994), operational taxonomic unit (OTU) Related concepts: Biospecies, genetic concordance species, morphospecies, non-dimensional species, phylospecies (in part), phenospecies, successional species, taxonomic species, quasispecies, viral species, genomospecies (bacteria) Principal authors: Beckner (1959); Sokal and Sneath (1963) Specifications: A cluster of characters that statistically covary, a family resemblance concept in which possession of most characters is required for inclusion in a species, but not all. A class of organisms that share most of a set of characters.
23. Recognition species
Synonyms: Specific mate recognition system (SMRS) Related concepts: Biospecies Principal author: Paterson (1985) Specifications: A species is that most inclusive population of individual, biparental organisms which share a common fertilization system
24. Reproductive competition species
Synonyms: Hypermodern species concept , Biospecies (in part) Principal author: Ghiselin (1974) Specifications: The most extensive units in the natural economy such that reproductive competition occurs among their parts.
25. Successional species
Synonyms: Paleospecies, evolutionary species (in part), chronospecies Principal authors: George (1956); Simpson (1961) Specifications: Arbitrary anagenetic stages in morphological forms, mainly in the paleontological record.
26. Taxonomic species
Synonyms: Cynical species concept (Kitcher 1984) Related concepts: Agamospecies, genealogical concordance species, morphospecies, phenospecies, phylospecies Principal author: Blackwelder (1967), but see Regan (1926) and Strickland et al. (1843) Specifications: Specimens considered by a taxonomist to be members of a kind on the evidence or on the assumption they are as alike as their offspring of hereditary relatives within a few generations. Whatever a competent taxonomist chooses to call a species.
Aguilar, Javier Fuertes, Josep Antoni Roselló, and Gonzalo Nieto Feliner (1999), “Molecular evidence for the compilospecies model of reticulate evolution in >Armeria> (Plumbaginaceae)”, Systematic Biology 48 (4):735-754.
Atran, Scott (1990), The cognitive foundations of natural history. New York: Cambridge University Press.
Avise, J. C., and R. M. Ball Jr (1990), “Principles of genealogical concordance in species concepts and biological taxonomy”, in D. Futuyma and J. Atonovics (eds.), Oxford Surveys in Evolutionary Biology, Oxford: Oxford University Press, 45-67.
Beckner, M (1959), The biological way of thought. New York: Columbia University Press.
Blackwelder, Richard E. (1967), Taxonomy: a text and reference book. New York: Wiley.
Cain, Arthur J. (1954), Animal species and their evolution. London: Hutchinson University Library.
Cracraft, Joel (1983), “Species concepts and speciation analysis”, in R. F. Johnston (ed.), Current Ornithology, New York: Plenum Press, 159-187.
Cronquist, A (1978), “Once again, what is a species?” in LV Knutson (ed.), BioSystematics in Agriculture, Montclair, NJ: Alleheld Osmun, 3-20.
Dobzhansky, Theodosius (1935), “A critique of the species concept in biology”, Philosophy of Science 2:344-355.
—— (1937), Genetics and the origin of species. New York: Columbia University Press.
—— (1950), “Mendelian populations and their evolution”, American Naturalist 74:312-321.
—— (1970), Genetics of the evolutionary process. New York: Columbia University Press.
Eigen, Manfred (1993), “Viral quasispecies”, Scientific American July 1993 (32-39).
Eldredge, Niles, and Joel Cracraft (1980), Phylogenetic patterns and the evolutionary process: method and theory in comparative biology. New York: Columbia University Press.
Euzéby, J.P. (2006), List of Prokaryotic Names with Standing in Nomenclature 2006 [cited 17/2/2006 2006]. Available from http://www.bacterio.cict.fr/.
George, T. N. (1956), “Biospecies, chronospecies and morphospecies”, in P. C. Sylvester-Bradley (ed.), The species concept in paleontology, London: Systematics Association, 123-137.
Ghiselin, Michael T. (1974), The economy of nature and the evolution of sex. Berkeley: University of California Press.
Harlan, J. R., and J. M. J. De Wet (1963), “The compilospecies concept”, Evolution 17:497-501.
Hennig, Willi (1950), Grundzeuge einer Theorie der Phylogenetischen Systematik. Berlin: Aufbau Verlag.
—— (1966), Phylogenetic systematics. Translated by D. Dwight Davis and Rainer Zangerl. Urbana: University of Illinois Press.
Kitcher, Philip (1984), “Species”, Philosophy of Science 51:308-333. Kornet, D (1993), “Internodal species concept”, J Theor Biol 104:407-435.
Kornet, D, and JW McAllister (1993), “The composite species concept”, in, Reconstructing species: Demarcations in genealogical networks, Rijksherbarium, Leiden: Unpublished phD dissertation, Institute for Theoretical Biology.
Mallet, J (1995), “The species definition for the modern synthesis”, Trends in Ecology and Evolution 10 (7):294-299.
Mayden, R. L. (1997), “A hierarchy of species concepts: the denoument in the saga of the species problem”, in M. F. Claridge, H. A. Dawah and M. R. Wilson (eds.), Species: The units of diversity, London: Chapman and Hall, 381-423.
Mayr, Ernst (1942), Systematics and the origin of species from the viewpoint of a zoologist. New York: Columbia University Press.
—— (1963), Animal species and evolution. Cambridge MA: The Belknap Press of Harvard University Press.
—— (1969), Principles of systematic zoology. New York: McGraw-Hill.
—— (1970), Populations, species, and evolution: an abridgment of Animal species and evolution. Cambridge, Mass.: Belknap Press of Harvard University Press.
Mayr, Ernst, and Peter D. Ashlock (1991), Principles of systematic zoology. 2nd ed. New York: McGraw-Hill,.
Meier, Rudolf, and Rainer Willmann (1997), “The Hennigian species concept”, in QD Wheeler and R Meier (eds.), Species concepts and phylogenetic theory: A debate, New York: Columbia University Press.
Mishler, Brent D., and Robert N. Brandon (1987), “Individuality, pluralism, and the Phylogenetic Species Concept”, Biology and Philosophy 2:397-414.
Nelson, Gareth J., and Norman I. Platnick (1981), Systematics and biogeography: cladistics and vicariance. New York: Columbia University Press.
Nixon, K. C., and Q. D. Wheeler (1990), “An amplification of the phylogenetic species concept”, Cladistics 6:211-223.
Paterson, Hugh E. H. (1985), “The recognition concept of species”, in E. Vrba (ed.), Species and speciation, Pretoria: Transvaal Museum, 21-29.
Pleijel, Frederik (1999), “Phylogenetic taxonomy, a farewell to species, and a revision of Heteropodarke (Hesionidae, Polychaeta, Annelida)”, Systematic Biology 48 (4):755-789.
Pleijel, Frederik, and G. W. Rouse (2000), “Least-inclusive taxonomic unit: a new taxonomic concept for biology”, Proceedings of the Royal Society of London – Series B: Biological Sciences 267 (1443):627-630.
Regan, C. Tate (1926), “Organic evolution”, Report of the British Association for the Advancement of Science, 1925:75-86.
Ridley, M (1989), “The cladistic solution to the species problem”, Biology and Philosophy 4:1-16.
Rosen, Donn E. (1979), “Fishes from the uplands and intermontane basins of Guatemala: revisionary studies and comparative biogeography”, Bulletin of the American Museum of Natural History 162:267-376.
Simpson, George Gaylord (1943), “Criteria for genera, species and subspecies in zoology and paleontology”, Annals New York Academy of Science 44:145-178.
—— (1961), Principles of animal taxonomy. New York: Columbia University Press.
Smith, Andrew B. (1994), Systematics and the fossil record: documenting evolutionary patterns. Oxford, OX; Cambridge, Mass., USA: Blackwell Science.
Sokal, Robert R., and P. H. A. Sneath (1963), Principles of numerical taxonomy, A Series of books in biology. San Francisco,: W. H. Freeman.
Sterelny, Kim (1999), “Species as evolutionary mosaics”, in R. A. Wilson (ed.), Species, New interdisciplinary essays, Cambridge, MA: Bradford/MIT Press, 119-138.
Strickland, Hugh. E., John Phillips, John Richardson, Richard Owen, Leonard Jenyns, William J. Broderip, John S. Henslow, William E. Shuckard, George R. Waterhouse, William Yarrell, Charles R. Darwin, and John O. Westwood (1843), “Report of a committee appointed “to consider of the rules by which the nomenclature of zoology may be established on a uniform and permanent basis”", Report of the British Association for the Advancement of Science for 1842:105-121.
Templeton, Alan R. (1989), “The meaning of species and speciation: A genetic perspective”, in D Otte and JA Endler (eds.), Speciation and its consequences, Sunderland, MA: Sinauer, 3-27.
Turesson, Göte (1922), “The species and variety as ecological units”, Hereditas 3:10-113.
Van Valen, L (1976), “Ecological species, multispecies, and oaks”, Taxon 25:233-239.
Wagner, Warren H. (1983), “Reticulistics: The recognition of hybrids and their role in cladistics and classification”, in N. I. Platnick and V. A. Funk (eds.), Advances in cladistics, New York: Columbia Univ. Press, 63-79.
Waples, R S (1991), “Pacific salmon, Oncorhynchus spp., and the definition of ‘species’ under the Endangered Species Act”, Marine Fisheries Review 53:11-22.
Wheeler, Quentin D., and Rudolf Meier, eds. (2000), Species concepts and phylogenetic theory: a debate. New York: Columbia University Press.
Wheeler, Quentin D., and Norman I. Platnick (2000), “The phylogenetic species concept (sensu Wheeler and Platnick)”, in Quentin D. Wheeler and Rudolf Meier (eds.), Species concepts and phylogenetic theory: A debate, New York: Columbia University Press, 55-69.
Wiley, E. O. (1978), “The evolutionary species concept reconsidered”, Systematic Zoology 27:17-26.
—— (1981), “Remarks on Willis’ species concept”, Systematic Zoology 30:86-87.
Wilkins, John S. (2003), “How to be a chaste species pluralist-realist: The origins of species modes and the Synapomorphic Species Concept”, Biology and Philosophy 18:621-638.
Wu, Chung-I (2001a), “Genes and speciation”, Journal of Evolutionary Biology 14 (6):889-891. —— (2001b), “The genic view of the process of speciation”, Journal of Evolutionary Biology 14:851-865.