An Annotated List of Species Concepts

  • Frank E. Zachos


There are about 30 species concepts in the literature. The exact number depends on what one considers a full or acknowledged concept (sometimes a comment or definition in the literature may not be called a concept and/or may not have made it into the canon of “official” species concepts), and it also depends on where one draws the line between similar or nearly identical concepts. The list below of 31 species concepts is therefore neither exhaustive nor the only way of listing species concepts. I mainly follow Mayden (1997, 1999), the list in Wilkins (2009b) which is based on Wilkins (2006a) and Wilkins’ website (Wilkins 2006b) but have included the General Lineage Species Concept and the Unified Species Concept by de Queiroz (1998, 1999, 2005b, 2007), added Stamos’ (2003) Biosimilarity Species Concept, Baum and Shaw’s (1995) Genealogical Species Concept and finally the Differential Fitness Species Concepts which was published more recently (Hausdorf 2011). I also sometimes use slightly different names than those given in other lists.


Reproductive Isolation Species Concept Phylogenetic Species Lineage Species Phylogenetic Species Concept 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Ackery PR, Vane-Wright RI (1984) Milkweed butterflies: their cladistics and biology. Cornell University Press, Ithaca, NYGoogle Scholar
  2. Avise JC, Ball RM Jr (1990) Principles of genealogical concordance in species concepts and biological taxonomy. In: Futuyma D, Antonovics J (eds) Oxford surveys in evolutionary biology. Oxford University Press, Oxford, pp 45–67Google Scholar
  3. Baker RJ, Bradley RD (2006) Speciation in mammals and the genetic species concept. J Mammal 87:643–662CrossRefPubMedPubMedCentralGoogle Scholar
  4. Baum D (1992) Phylogenetic species concepts. Trends Ecol Evol 7:1–2CrossRefPubMedGoogle Scholar
  5. Baum DA (1998) Individuality and the existence of species through time. Syst Biol 47:641–653CrossRefPubMedGoogle Scholar
  6. Baum DA, Shaw KL (1995) Genealogical perspectives on the species problem. In: Hoch PC, Stephenson AG (eds) Experimental and molecular approaches to plant biosystematics. Monographs in Systematic Botany from the Missouri Botanical Garden, vol 53. Missouri Botanical Graden, St. Louis, pp 289–303Google Scholar
  7. Blackwelder RE (1967) Taxonomy: a text and reference book. Wiley, New YorkGoogle Scholar
  8. Cain AJ (1954) Animal species and their evolution. Hutchinson University Library, LondonGoogle Scholar
  9. Cantino PD, de Queiroz K (2010) PhyloCode. International Code of Phylogenetic Nomenclature. Version 4c.
  10. Cantino PD, Bryant HN, de Queiroz K, Donoghue MJ, Eriksson T, Hillis DM, Lee MSY (1999) Species names in phylogenetic nomenclature. Syst Biol 48:790–807CrossRefPubMedGoogle Scholar
  11. Cracraft J (1983) Species concepts and speciation analysis. In: Johnston RF (ed) Current Ornithology, vol 1. Plenum, New York, pp 159–187 (reprinted in Ereshefsky 1992)Google Scholar
  12. Cracraft J (1997) Species concepts in systematics and conservation biology – an ornithological viewpoint. In: Claridge MF, Dawah HA, Wilson MR (eds) Species: the units of biodiversity. Chapman & Hall, London, pp 325–339Google Scholar
  13. Crandall KA, Bininda-Emonds ORP, Mace GM, Wayne RK (2000) Considering evolutionary processes in conservation biology. Trends Ecol Evol 15:290–295CrossRefPubMedGoogle Scholar
  14. Crisp MD, Chandler GT (1996) Paraphyletic species. Telopea 6:813–844CrossRefGoogle Scholar
  15. Cronquist A (1978) Once again, what is a species? In: Knutson LV (ed) Biosystematics in agriculture. Alleheld Osmun, Montclair, NJ, pp 3–20Google Scholar
  16. Dayrat B, Cantino PD, Clarke JA, de Queiroz K (2008) Species names in the PhyloCode: the approach adopted by the International Society for Phylogenetic Nomenclature. Syst Biol 57:507–514CrossRefPubMedGoogle Scholar
  17. de Queiroz K (1998) The general lineage concept of species, species criteria, and the process of speciation. In: Howard D, Berlocher SH (eds) Endless forms: species and speciation. Oxford University Press, Oxford, pp 57–75Google Scholar
  18. de Queiroz K (1999) The general lineage concept of species and the defining properties of the species category. In: Wilson RA (ed) Species. New interdisciplinary essays. MIT Press, Cambridge, MA, pp 49–89Google Scholar
  19. de Queiroz K (2005b) A unified concept of species and its consequences for the future of taxonomy. Proc Calif Acad Sci 56:196–215Google Scholar
  20. de Queiroz K (2007) Species concepts and species delimitation. Syst Biol 56:879–886CrossRefPubMedGoogle Scholar
  21. de Queiroz K, Gauthier J (1992) Phylogenetic taxonomy. Annu Rev Evol Syst 23:449–480CrossRefGoogle Scholar
  22. de Queiroz K, Gauthier J (1994) Toward a phylogenetic system of biological nomenclature. Trends Ecol Evol 9:27–31CrossRefPubMedGoogle Scholar
  23. Dobzhansky T (1935) A critique of the species concept in biology. Philos Sci 2:344–355CrossRefGoogle Scholar
  24. Dobzhansky T (1937) Genetics and the origin of species. Columbia University Press, New YorkGoogle Scholar
  25. Dobzhansky T (1940) Speciation as a stage in evolutionary divergence. Am Nat 74:312–321CrossRefGoogle Scholar
  26. Dobzhansky T (1950) Mendelian populations and their evolution. Am Nat 84:401–418CrossRefGoogle Scholar
  27. Donoghue MJ (1985) A critique of the biological species concept and recommendations for a phylogenetic alternative. Bryologist 88:172–181CrossRefGoogle Scholar
  28. Eigen M (1993) Viral quasispecies. Sci Am July:32–39Google Scholar
  29. Eldredge N, Cracraft J (1980) Phylogenetic patterns and the evolutionary process. Method and theory in comparative biology. Columbia University Press, New YorkGoogle Scholar
  30. Ereshefsky M (2001) The poverty of the Linnaean hierarchy. A philosophical study of biological taxonomy. Cambridge University Press, Cambridge, UKCrossRefGoogle Scholar
  31. George TN (1956) Biospecies, chronospecies and morphospecies. In: Sylvester-Bradley PC (ed) The species concept in paleontology. Systematics Association, London, pp 123–137Google Scholar
  32. Ghiselin MT (1974a) A radical solution to the species problem. Syst Zool 23:536–544 (reprinted in Ereshefsky 1992)Google Scholar
  33. Ghiselin MT (1974b) The economy of nature and the evolution of sex. University of California Press, Berkeley, CAGoogle Scholar
  34. Ghiselin MT (1997) Metaphysics and the origin of species. State University of New York Press, Albany, NYGoogle Scholar
  35. Graybeal A (1995) Naming species. Syst Biol 44:237–250CrossRefGoogle Scholar
  36. Groves C, Grubb P (2011) Ungulate taxonomy. The Johns Hopkins University Press, Baltimore, MDGoogle Scholar
  37. Harlan JR, De Wet JMJ (1963) The compilospecies concept. Evolution 17:497–501CrossRefGoogle Scholar
  38. Hausdorf B (2011) Progress toward a general species concept. Evolution 65:923–931CrossRefPubMedGoogle Scholar
  39. Hennig W (1950) Grundzüge einer Theorie der Phylogenetischen Systematik. Aufbau Verlag, BerlinGoogle Scholar
  40. Hennig W (1966) Phylogenetic systematics. University of Illinois Press, Urbana, ILGoogle Scholar
  41. Kitcher P (1984) Species. Philos Sci 51:308–333 (reprinted in Ereshefsky 1992)Google Scholar
  42. Kornet D (1993) Permanent splits as speciation events: a formal reconstruction of the internodal species concept. J Theor Biol 164:407–435CrossRefGoogle Scholar
  43. Kornet D, McAllister JW (1993) The composite species concept. In: Reconstructing species: demarcations in genealogical networks. Unpublished PhD dissertation, Institute for Theoretical Biology, Rijksherbarium, LeidenGoogle Scholar
  44. Mallet J (1995) A species definition for the modern synthesis. Trends Ecol Evol 10:294–299CrossRefPubMedGoogle Scholar
  45. Mayden RL (1997) A hierarchy of species concepts: the denouement in the saga of the species problem. In: Claridge MF, Dawah HA, Wilson MR (eds) Species: the units of biodiversity. Chapman & Hall, London, pp 381–424Google Scholar
  46. Mayden RL (1999) Consilience and a hierarchy of species concepts: advances towards closure on the species puzzle. J Nematol 31:95–116PubMedPubMedCentralGoogle Scholar
  47. Mayr E (1940) Speciation phenomena in birds. Am Nat 74:249–278CrossRefGoogle Scholar
  48. Mayr E (1942) Systematics and the origin of species. Columbia University Press, New YorkGoogle Scholar
  49. Mayr E (1963) Animal species and evolution. The Belknap Press of Harvard University Press, Cambridge, MACrossRefGoogle Scholar
  50. Mayr E (1969) Principles of systematic zoology. McGraw-Hill, New YorkGoogle Scholar
  51. Mayr E (1982) The growth of biological thought. Diversity, evolution, and inheritance. The Belknap Press of Harvard University Press, Cambridge, MAGoogle Scholar
  52. McKitrick MC, Zink RM (1988) Species concepts in ornithology. Condor 90:1–14CrossRefGoogle Scholar
  53. Meier R, Willmann R (2000a) The Hennigian species concept. In: Wheeler QD, Meier R (eds) Species concepts and phylogenetic theory – a debate. Columbia University Press, New York, pp 30–43Google Scholar
  54. Minelli A (2015) Taxonomy faces speciation: the origin of species or the fading out of the species? Biodivers J 6:123–138Google Scholar
  55. Mishler BD (1985) The morphological, developmental, and phylogenetic basis of species concepts in bryophytes. Bryologist 88:207–214CrossRefGoogle Scholar
  56. Mishler BD (1999) Getting rid of species? In: Wilson RA (ed) Species. New interdisciplinary essays. MIT Press, Cambridge, MA, pp 307–315Google Scholar
  57. Mishler BD, Brandon RN (1987) Individuality, pluralism, and the phylogenetic species concept. Biol Philos 2:397–414CrossRefGoogle Scholar
  58. Mishler BD, Donoghue MJ (1982) Species concepts: a case for pluralism. Syst Zool 31:491–503 (reprinted in Ereshefsky 1992)Google Scholar
  59. Mishler BD, Theriot EC (2000a) The phylogenetic species concept (sensu Mishler and Theriot): monophyly, apomorphy, and phylogenetic species concepts. In: Wheeler QD, Meier R (eds) Species concepts and phylogenetic theory – a debate. Columbia University Press, New York, pp 44–55Google Scholar
  60. Moritz C (1994) Defining ‘evolutionary significant units’ for conservation. Trends Ecol Evol 9:373–375CrossRefPubMedGoogle Scholar
  61. Nelson G, Platnick NI (1981) Systematics and biogeography: cladistics and vicariance. Columbia University Press, New YorkGoogle Scholar
  62. Nixon KC, Wheeler QD (1990) An amplification of the phylogenetic species concept. Cladistics 6:211–223CrossRefGoogle Scholar
  63. O’Hara RJ (1993) Systematic generalization, historical fate, and the species problem. Syst Biol 42:231–246CrossRefGoogle Scholar
  64. Orr HA (2001) Some doubts about (yet another) view of species. J Evol Biol 14:870–871CrossRefGoogle Scholar
  65. Paterson H (1985) The recognition concept of species. In: Vrba E (ed) Species and speciation. Transvaal Museum, Pretoria, pp 21–29 (reprinted in Ereshefsky 1992)Google Scholar
  66. Paterson H (1988) On defining species in terms of sterility: problems and alternatives. Pac Sci 42:65–71Google Scholar
  67. Platnick NI, Wheeler QD (2000) A defense of the phylogenetic species concept (sensu Wheeler and Platnick). In: Wheeler QD, Meier R (eds) Species concepts and phylogenetic theory – a debate. Columbia University Press, New York, pp 185–197Google Scholar
  68. Pleijel F (1999) Phylogenetic taxonomy, a farewell to species, and a revision of Heteropodarke (Hesionidae, Polychaeta, Annelida). Syst Biol 48:755–789CrossRefGoogle Scholar
  69. Pleijel F, Rouse GW (2000) Least-inclusive taxonomic unit: a new taxonomic concept for biology. Proc R Soc Lond B 267:627–630CrossRefGoogle Scholar
  70. Poelstra JW, Vijay N, Bossu CM, Lantz H, Ryll B, Müller I, Baglione V, Unneberg P, Wikelski M, Grabherr MG, Wolf JBW (2014) The genomic landscape underlying phenotypic integrity in the face of gene flow in crows. Science 344:1410–1414CrossRefPubMedGoogle Scholar
  71. Regan CT (1926) Organic evolution. Report of the British Association for the Advancement of Science 1925:75–86Google Scholar
  72. Ridley M (1989) The cladistic solution to the species problem. Biol Philos 4:1–16CrossRefGoogle Scholar
  73. Rosen DE (1978) Vicariant patterns and historical explanation in biogeography. Syst Zool 27:159–188CrossRefGoogle Scholar
  74. Rosen DE (1979) Fishes from the uplands and intermontane basins of Guatemala: revisionary studies and comparative biogeography. Bull Am Mus Nat Hist 162:267–376Google Scholar
  75. Rosselló-Mora R, Amann R (2001) The species concept for prokaryotes. FEMS Microbiol Rev 25:39–67CrossRefPubMedGoogle Scholar
  76. Rundle HD, Breden F, Griswold C, Mooers AØ, Vos RA, Whitton J (2001) Hybridization without guilt: gene flow and the biological species concept. J Evol Biol 14:868–869CrossRefGoogle Scholar
  77. Ryder OA (1986) Species conservation and systematics: the dilemma of subspecies. Trends Ecol Evol 1:9–10CrossRefGoogle Scholar
  78. Simpson GG (1943) Criteria for genera, species, and subspecies in zoology and paleontology. Ann NY Acad Sci 44:145–178CrossRefGoogle Scholar
  79. Simpson GG (1951) The species concept. Evolution 5:285–298CrossRefGoogle Scholar
  80. Simpson GG (1961) Principles of animal taxonomy. Columbia University Press, New YorkGoogle Scholar
  81. Sneath PHA (1976) Phenetic taxonomy at the species level and above. Taxon 25:437–450CrossRefGoogle Scholar
  82. Sober E (1984) Sets, species, and evolution: comments on Philip Kitcher’s “species”. Philos Sci 51:334–341CrossRefGoogle Scholar
  83. Sokal RR (1973) The species problem reconsidered. Syst Zool 22:360–374CrossRefGoogle Scholar
  84. Sokal RR, Sneath PHA (1963) Principles of numerical taxonomy. W. H. Freeman, San Francisco, CAGoogle Scholar
  85. Stamos DN (2003) The species problem. Biological species, ontology, and the metaphysics of biology. Lexington Books, LanhamGoogle Scholar
  86. Templeton AR (1989) The meaning of species and speciation: a genetic perspective. In: Otte D, Endler JA (eds) Speciation and its consequences. Sinauer Associates, Sunderland, MA, pp 3–27 (reprinted in Ereshefsky 1992)Google Scholar
  87. Templeton AR (1994) The role of molecular genetics in speciation studies. In: Schierwater B, Streit B, Wagner GP, DeSalle R (eds) Molecular ecology and evolution: approaches and applications. Birkhäuser, BaselGoogle Scholar
  88. Turesson G (1929) Zur Natur und Begrenzung der Arteinheiten. Hereditas 12:323–334 (in German with an English summary)CrossRefGoogle Scholar
  89. Van Valen L (1976) Ecological species, multispecies, and oaks. Taxon 25:233–239 (reprinted in Ereshefsky 1992)Google Scholar
  90. Wagner WH (1983) Reticulistics: the recognition of hybrids and their role in cladistics and classification. In: Platnick NI, Funk VA (eds) Advances in cladistics, vol 2, Proceedings of the second meeting of the Willi Hennig Society. Columbia University Press, New York, pp 63–79Google Scholar
  91. Waples RS (1991) Pacific salmon, Oncorhynchus spp., and the definition of ‘species’ under the Endangered Species Act. Mar Fish Rev 53:11–22Google Scholar
  92. Wheeler WC (2012) Systematics. A course of lectures. Wiley-Blackwell, OxfordCrossRefGoogle Scholar
  93. Wheeler WC (2014) Phyletic groups on networks. Cladistics 30:447–451CrossRefGoogle Scholar
  94. Wheeler QD, Platnick NI (2000a) The phylogenetic species concept (sensu Wheeler and Platnick). In: Wheeler QD, Meier R (eds) Species concepts and phylogenetic theory – a debate. Columbia University Press, New York, pp 55–69Google Scholar
  95. Wiley EO (1978) The evolutionary species concept reconsidered. Syst Zool 27:17–26 (reprinted in Ereshefsky 1992)Google Scholar
  96. Wiley EO, Mayden RL (2000a) The evolutionary species concept. In: Wheeler QD, Meier R (eds) Species concepts and phylogenetic theory – a debate. Columbia University Press, New York, pp 70–89Google Scholar
  97. Wilkins JS (2006a) Species, kinds, and evolution. Rep Cent Natl Sci Educ 26:36–45Google Scholar
  98. Wilkins JS (2009a) Species. A history of the idea. University of California Press, Berkeley, CAGoogle Scholar
  99. Wilkins JS (2009b) Defining species. A sourcebook from antiquity to today. Peter Lang Publishing, New YorkGoogle Scholar
  100. Wilkins JS (2011) Philosophically speaking, how many species concepts are there? Zootaxa 2765:58–60Google Scholar
  101. Willmann R (1985) Die Art in Raum und Zeit. Das Artkonzept in der Biologie und Paläontologie. Verlag Paul Parey, BerlinGoogle Scholar
  102. Willmann R (1986) Reproductive isolation and the limits of the species in time. Cladistics 2:356–358CrossRefGoogle Scholar
  103. Wu C-I (2001a) The genic view of the process of speciation. J Evol Biol 14:851–865CrossRefGoogle Scholar
  104. Wu C-I (2001b) Genes and speciation. J Evol Biol 14:889–891CrossRefGoogle Scholar
  105. Zachos FE (2009) Gene trees and species trees – mutual influences and interdependences of population genetics and systematics. J Zool Syst Evol Res 47:209–218CrossRefGoogle Scholar

Copyright information

© Springer International Publishing Switzerland 2016

Authors and Affiliations

  • Frank E. Zachos
    • 1
  1. 1.Mammal CollectionNatural History Museum ViennaViennaAustria

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