Academia.eduAcademia.edu

Outline

Classes or Individuals? The Paradox of Systematics Revisited

2004, Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences

https://doi.org/10.1016/J.SHPSC.2004.09.006

Abstract

The circumscription of taxa and classification of organisms are fundamental tasks in the systematization of biological diversity. Their success depends on a unified idea concerning the species concept, evolution, and taxonomy; paradoxically, however, it requires a complete distinction between taxa and evolutionary units. To justify this view, I discuss these three topics of systematics. Species concepts are examined, and I propose a redefinition for the Taxonomic Species Concept based on nomenclatural properties, in which species are classes conventionally represented by a binomial. Speciation is subsequently discussed, to demonstrate that concepts on the evolutionary process are damaged when species are considered as evolutionary units. Speciation should be considered a transition among patterns of a population (anagenesis) and, consequently, always a sympatric process. This view contrasts with the majority of speciation models, which analyze cladogeneses and classify speciation geographically, usually considering allopatry the essential condition for the differentiation process. Finally, taxonomy is considered, to show that the equivalence between evolutionary and taxonomic units may also damage the practice of biological systematization. Principles and rules of nomenclature and classification should offer freedom to accommodate divergent opinions about systematics and to incorporate the evolution of knowledge; therefore, they should remain independent of biological theories. #

References (130)

  1. Andersson, L. (1992). Definitions and models-a reply to Bremer, Eriksson and Grant. Taxon, 41, 313- 314.
  2. Ayala, F. J. (1982). Gradualism versus punctualism in speciation: reproductive isolation, morphology, genetics. In C. Barigozzi (Ed.), Mechanisms of speciation (pp. 51-66). New York: Alan R. Liss.
  3. Baum, D. A. (1992). Phylogenetic species concepts. Trends in Ecology and Evolution, 7, 1-2.
  4. Baum, D. A., & Donoghue, M. J. (1995). Choosing among alternative 'phylogenetic' species concepts. Systematic Botany, 20, 560-573.
  5. Beckner, M. (1959). The biological way of thought. New York: Columbia University Press.
  6. Bernier, R. (1984). The species as an individual: Facing essentialism. Systematic Zoology, 33, 460-469.
  7. Bessey, C. E. (1908). The taxonomic aspect of the species question. American Naturalist, 42, 218-224.
  8. Blackwelder, R. E. (1962). Animal taxonomy and the new systematics. Survey of Biological Progress, 4, 31-35, 53-57. (Reprinted in Slobodchikoff, 1976)
  9. Bremer, K., & Wanntorp, H.-E. (1979). Hierarchy and reticulation in systematics. Systematic Zoology, 28, 624-627.
  10. Bryant, H. N., & Cantino, P. D. (2002). A review of criticisms of phylogenetic nomenclature: Is taxo- nomic freedom the fundamental issue? Biological Review, 77, 39-55.
  11. Brummitt, R.K. (1996). Quite happy with the present code, thank you. In J. L. Reveal (Ed.), Proceed- ings of a mini-symposium on biological nomenclature in the 21st century. (Available at http://www.li- fe.umd.edu/emeritus/reveal/pbio/nomcl/brum.html)
  12. Brummitt, R. K. (1997). Taxonomy versus cladonomy, a fundamental controversy in biological systema- tics. Taxon, 46, 723-734.
  13. Brummitt, R. K. (2002). How to chop up a tree. Taxon, 51, 31-41.
  14. Brummitt, R. K., & Sosef, M. S. M. (1998). Paraphyletic taxa are inherent in Linnaean classification-a reply to Freudenstein. Taxon, 47, 411-412.
  15. Burma, B. H. (1954). Reality, existence, and classification: A discussion of the species problem. Madron ˜o, 127, 193-209.
  16. Camp, W. H. (1951). Biosystematy. Brittonia, 7, 113-127.
  17. Camp, W. H., & Gilly, C. L. (1943). The structure and origin of species. Brittonia, 4, 323-385.
  18. Cantino, P. D. (2000). Phylogenetic nomenclature: Addressing some concerns. Taxon, 49, 85-93.
  19. Cantino, P. D., Olmstead, R. G., & Wagstaff, S. J. (1997). A comparison of phylogenetic nomenclature with the current system: A botanical case study. Systematic Biology, 46, 313-331.
  20. Caplan, A. L. (1981). Back to class: A note on the ontology of species. Philosophy of Science, 48, 130- 140.
  21. Chappill, J. A. (1989). Quantitative character in phylogenetic analysis. Cladistics, 5, 217-234.
  22. Claridge, M. F., & Dawah, H. A. (Eds.). (1997). Species: The units of biodiversity. London: Chapman and Hall.
  23. Coyne, J. A. (1992). Genetics and speciation. Nature, 355, 511-515.
  24. Coyne, J. A., & Orr, H. A. (1989). Two rules of speciation. In D. Otte, & J. A. Endler (Eds.), Speciation and its consequences (pp. 180-207). Sunderland: Sinauer Associates.
  25. Cracraft, J. (1983). Species concepts and speciation analysis. Current Ornithology, 1, 159-187.
  26. Cracraft, J. (1989). Speciation and its ontology: The empirical consequences of alternative species con- cepts for understanding patterns and processes of differentiation. In D. Otte, & J. A. Endler (Eds.), Speciation and its consequences (pp. 28-59). Sunderland: Sinauer Associates.
  27. Cronquist, A. (1978). Once again, what is a species? In L. V. Knutson (Ed.), Biosystematics in agri- culture (pp. 3-20). Montclair: Allanheld and Osmun.
  28. Cronquist, A. (1987). A botanical critique of cladism. Botanical Review, 53, 1-52.
  29. Crowson, R. A. (1970). Classification and biology. New York: Atherton Press.
  30. Davis, J. I., & Nixon, K. (1992). Populations, genetic variation, and the delimitation of phylogenetic species. Systematic Biology, 41, 421-435.
  31. Davis, P. H., & Heywood, V. H. (1963). Principles of angiosperm taxonomy. Edinburgh: Oliver and Boyd.
  32. De Pinna, M. C. C. (1999). Species concepts and phylogenetics. Review of Fish Biology and Fisheries, 9, 353-373.
  33. De Queiroz, K. (1996). A phylogenetic approach to biological nomenclature as an alternative to the Lin- naean systems in current use. In J. L. Reveal (Ed.), Proceedings of a mini-symposium on biological nomenclature in the 21st century. (Available at http://www.life.umd.edu/emeritus/reveal/pbio/ nomcl/dequ.html)
  34. De Queiroz, K. (1997). The Linnaean hierarchy and the evolutionization of taxonomy, with emphasis on the problem of nomenclature. Aliso, 15, 125-144.
  35. De Queiroz, K. (1998). The general lineage concept of species, species criteria, and the process of specia- tion.
  36. In D. J. Howard, & S. H. Berlocher (Eds.), Endless forms: Species and speciation (pp. 57-75). Oxford: Oxford University Press.
  37. De Queiroz, K. (1999). The general lineage concept of species and the defining properties of the species category. In R. A. Wilson (Ed.), Species: New interdisciplinary essays (pp. 49-89). Cambridge: MIT Press.
  38. De Queiroz, K., & Donoghue, M. J. (1988). Phylogenetic systematics and the species problem. Cladis- tics, 4, 317-338.
  39. De Queiroz, K., & Gauthier, J. (1990). Phylogenetic as a central principle in taxonomy: Phylogenetic definition of taxon names. Systematic Zoology, 39, 307-322.
  40. De Queiroz, K., & Gauthier, J. (1992). Phylogenetic taxonomy. Annual Review of Ecology and Systema- tics, 23, 449-480.
  41. De Queiroz, K., & Gauthier, J. (1994). Toward a phylogenetic system of biological nomenclature. Trends in Ecology and Evolution, 9, 27-31.
  42. Dobzhansky, T. (1937). Genetics and the origin of species. New York: Columbia University Press.
  43. Dobzhansky, T. (1951). Genetics and the origin of species (3rd ed). New York: Columbia University Press.
  44. Dobzhansky, T. (1970). Genetics of the evolutionary process. New York: Columbia University Press.
  45. Dominguez, E., & Wheeler, Q. D. (1997). Taxonomic stability is ignorance. Cladistics, 13, 367-372.
  46. Dupre ´, J. (1999). On the impossibility of monistic account of species. In R. A. Wilson (Ed.), Spe- cies:New interdisciplinary essays (pp. 3-22). Cambridge: MIT Press.
  47. Dupre ´, J. (2001). In defense of classification. Studies in History and Philosophy of Biological and Biome- dical Sciences, 32, 203-219.
  48. Ehrlich, P. R., & Raven, P. H. (1969). Differentiation of populations. Science, 165, 1228-1232.
  49. Eldredge, N., & Cracraft, J. (1980). Phylogenetic patterns and the evolutionary process. New York: Columbia University Press.
  50. Ereshefsky, M. (Ed.) (1992), The units of evolution: Essays on the nature of species. Cambridge: MIT Press.
  51. Ereshefsky, M. (1997). The evolution of the Linnaean hierarchy. Biology and Philosophy, 12, 493-519.
  52. Ereshefsky, M. (2001). Names, numbers and indentations: a guide to post-Linnaean taxonomy. Studies in History and Philosophy of Biological and Biomedical Sciences, 32, 361-383.
  53. Forey, P. L. (2001). The PhyloCode: Description and commentary. Bulletin of Zoological Nomenclature, 58, 81-96.
  54. Forey, P. L. (2002). PhyloCode-pain, no gain. Taxon, 51, 43-54.
  55. Frost, D. R., & Kluge, A. G. (1994). A consideration of epistemology in systematic biology, with special reference to species. Cladistics, 10, 259-294.
  56. Ghiselin, M. T. (1974). A radical solution to the species problem. Systematic Zoology, 23, 536-544.
  57. Ghiselin, M. T. (1987). Species concepts, individuality, and objectivity. Biology and Philosophy, 2, 127- 143. Grant, V. (1992). Comments on the Ecological Species Concept. Taxon, 41, 310-312.
  58. Graybeal, A. (1995). Naming species. Systematic Biology, 44, 237-250.
  59. Gregg, J. R. (1950). Taxonomy, language and reality. American Naturalist, 84, 419-435.
  60. Griffths, G. C. D. (1974). On the foundations of biological classification. Systematic Zoology, 22, 338- 343.
  61. Griffths, G. C. D. (1976). The future of Linnean nomenclature. Systematic Zoology, 25, 168-173.
  62. Ha ¨rlin, M. (1998). Taxonomic names and phylogenetic trees. Zoologica Scripta, 27, 390-391.
  63. Hedberg, O. (1995). Cladistics in taxonomic botany-master or slave? Taxon, 44, 3-11.
  64. Hennig, W. (1966). Phylogenetic systematics. Urbana: University Illinois Press.
  65. Hey, J. (2001). The mind of the species problem. Trends in Ecology and Evolution, 16, 326-339.
  66. Huey, R. B., Gilchrist, G. W., Carlson, M. L., Berrigan, D., & Serra, L. (2000). Rapid evolution of geo- graphic cline size in an introduced fly. Science, 287, 308-309.
  67. Hull, D. L. (1965). The effects of essentialism on taxonomy-two thousand years of stasis (II). British Journal for the Philosophy of Science, 16, 1-18.
  68. Hull, D. L. (1976). Are species really individuals? Systematic Zoology, 25, 174-191.
  69. Hull, D. L. (1979). The limits of cladism. Systematic Zoology, 28, 416-440.
  70. Hull, D. L. (1981). Discussion: Kitts and Kitts and Caplan on species. Philosophy of Science, 48, 141- 152.
  71. Hull, D. L. (1997). The ideal species concept and why we can't get it. In M. F. Claridge, H. A. Dawah, & M. R. Wilson (Eds.), Species: the units of biodiversity (pp. 357-380). London: Chapman and Hall.
  72. Hull, D. L. (1999). On the plurality of species: Questioning the party line. In R. A. Wilson (Ed.), Spe- cies: New interdisciplinary essays (pp. 23-48). Cambridge: MIT Press.
  73. Jeanmonod, D. (1984). La spe ´ciation: Aspects divers et mode `les re ´cents. Candollea, 39, 151-194.
  74. Jonsell, B. (1984). The biological species concept reexamined. In W. F. Grant (Ed.), Plant biosystematics (pp. 159-168). Ontario: Academic Press Canada.
  75. Jorgensen, P. M. (2000). Names are defined, but not as taxa. Taxon, 49, 779.
  76. Judd, W. S., Sanders, R. W., & Donoghue, M. J. (1994). Angiosperms family pairs: Preliminary phylo- genetic analyses. Harvard Papers in Botany, 5, 1-51.
  77. Kitcher, P. (1984). Species. Philosophy of Science, 51, 308-333.
  78. Kluge, G., & Wolf, A. J. (1993). Cladistics: What's in a word? Cladistics, 9, 183-199.
  79. Knox, E. B. (1998). The use of hierarchies as organizational models in systematics. Biological Journal of the Linnean Society, 63, 1-49.
  80. Kondrashov, A. S., & Mina, M. V. (1986). Sympatric speciation: When is it possible? Biological Journal of the Linnean Society, 27, 201-223.
  81. Levin, D. A. (1979). The nature of plant species. Science, 204, 381-384.
  82. Lide ´n, M. (1992). Species-where's the problem? Taxon, 41, 315-317.
  83. Lide ´n, M. (1997). Process and pattern, again. In response to Sosef. Taxon, 46, 527-534.
  84. Lide ´n, M., & Oxelman, B. (1989). Species-pattern or process? Taxon, 38, 228-232.
  85. Lide ´n, M., & Oxelman, B. (1996). Do we need 'Phylogenetic Taxonomy'. Zoologica Scripta, 25, 183- 185. Lide ´n, M., Oxelman, B., Backlund, A., Andersson, L., Bremer, B., Eriksson, R., Moberg, R., Nordal, I., Persson, K., Thulin, M., & Zimmer, B. (1997). Charlie is our darling. Taxon, 46, 735-738.
  86. Loevtrup, S. (1987). On species and other taxa. Cladistics, 3, 157-177.
  87. Luckow, M. (1995). Species concepts: Assumptions, methods, and applications. Systematic Botany, 20, 589-605.
  88. Lynch, J. D. (1989). The gauge of speciation: On the frequencies of modes of speciation. In D. Otte, & J. A. Endler (Eds.), Speciation and its consequences (pp. 527-553). Sunderland: Sinauer Associates.
  89. McDade, L. A. (1995). Species concepts and problems in practice: Insight from botanical monographs. Systematic Botany, 20, 22-60.
  90. Mallet, J. (1995). A species definition for the modern synthesis. Trends in Ecology and Evololution, 10, 294-299.
  91. Mayden, R. L. (1997). A hierarchy of species concepts: the denouement in the saga of the species problem. In M. F. Claridge, H. A. Dawah, & M. R. Wilson (Eds.), Species: The units of biodiversity (pp. 381-424). London: Chapman and Hall.
  92. Maynard Smith, J. (1966). Sympatric speciation. American Naturalist, 100, 637-650.
  93. Mayr, E. (1942). Systematics and the origin of species. New York: Columbia University Press.
  94. Mayr, E. (1958). Species concepts and definitions. In E. Mayr (Ed.), The species problem (2nd ed.) (pp. 1-22). Publication, no. 50 Washington, DC: American Association for the Advancement of Science.
  95. Mayr, E. (1970). Population, species, and evolution: An abridgment of animal species and evolution. Cam- bridge: Belknap Press of Harvard University Press.
  96. Minelli, A. (1993). Biological systematics: The state of the art. London: Chapman and Hall.
  97. Mishler, B. D. (1999). Getting rid of species? In R. A. Wilson (Ed.), Species: New interdisciplinary essays (pp. 307-315). Cambridge: MIT Press.
  98. Mishler, B. D., & Donoghue, M. J. (1982). Species concepts: A case for pluralism. Systematic Zoology, 31, 491-503.
  99. Moore, G. (1998). A comparison of traditional and phylogenetic nomenclature. Taxon, 47, 561-579.
  100. Nelson, G. J., & Platnick, N. I. (1981). Systematics and biogeography: Cladistics and vicariance. New York: Columbia University Press.
  101. Nixon, K. C., & Carpenter, J. M. (2000). On the other 'Phylogenetic Systematics'. Cladistics, 16, 269- 318.
  102. Nixon, K. C., & Wheeler, Q. D. (1990). An amplification of the phylogenetic species concept. Cladistics, 6, 211-223.
  103. Paterson, H. E. H. (1985). The recognition concept of species. In E. S. Vrba (Ed.), Species and speciation (pp. 21-29). Pretoria: Transvaal Museum.
  104. Platnick, N. I. (1979). Philosophy and the transformation of cladistics. Systematic Zoology, 28, 537-546.
  105. Rieseberg, L. H., & Brouillet, L. (1994). Are many plant species paraphyletic? Taxon, 43, 21-32.
  106. Rundle, H. D., Nagel, L., Boughman, J. W., & Schluter, D. (2000). Natural selection and parallel spe- ciation in sympatric sticklebacks. Science, 287, 306-308.
  107. Schander, C., & Thollesson, M. (1995). Phylogenetic taxonomy-some comments. Zoologica Scripta, 24, 263-268.
  108. Schneider, C. J., Smith, T. B., Larison, B., & Moritz, C. (1999). A test of alternative models of diversifi- cation in tropical rainforest: Ecological gradients vs. rainforest refugia. Proceedings of the National Academy of Science of the United States of America, 96, 13869-13873.
  109. Sennblad, B., & Bremer, B. (1996). The familial and subfamilial relationships of Apocynaceae and Asclepiadaceae evaluated with rbcL data. Plant Systematics and Evolution, 202, 153-175.
  110. Simpson, G. G. (1961). Principles of animal taxonomy. New York: Columbia University Press.
  111. Slobodchikoff, C. N. (Ed.) (1976), Concepts of species. Benchmark Papers in Systematics and Evolution- ary Biology, Vol. 3. Stroudsburg: Dowden, Hutchinson and Ross.
  112. Sober, E. (1980). Evolution, population thinking, and essentialism. Philosophy of Science, 47, 350-383.
  113. Sokal, R. R., & Crovello, T. J. (1970). The biological species concept: A critical evaluation. American Naturalist, 104, 127-153.
  114. Sosef, M. S. M. (1997). Hierarchical models, reticulate evolution and inevitability of paraphyletic supraspecific taxa. Taxon, 46, 75-85.
  115. Stevens, P. F. (1991). Character states, morphological variation, and phylogenetic analysis: A review. Systematic Botany, 16, 553-583.
  116. Stevens, P. F. (1998). What kind of classification should the practising taxonomist use to be saved? In J. Dransfield, M. J. E. Coode, & D. A. Simpson (Eds.), Plant Diversity in Malesia III (pp. 295-319). Kew: Royal Botanic Gardens.
  117. Struwe, L., Albert, V. A., & Bremer, B. (1994). Cladistics and family level classification of the Gentia- nales. Cladistics, 10, 175-205.
  118. Stuessy, T. F. (2001). Taxon names are still not defined. Taxon, 50, 185-186.
  119. Tauber, C. A., & Tauber, M. J. (1989). Sympatric speciation in insects: Perception and perspective. In D. Otte, & J. A. Endler (Eds.), Speciation and its consequences (pp. 307-344). Sunderland: Sinauer Associates.
  120. Templeton, A. R. (1989). The meaning of species and speciation: a genetic perspective. In D. Otte, & J. A. Endler (Eds.), Speciation and its consequences (pp. 3-27). Sunderland: Sinauer Associates.
  121. Thoday, J. M., & Boam, T. B. (1959). Effects of disruptive selection II. Polymorphism and divergence without isolation. Heredity, 13, 205-218.
  122. Tregenza, T., & Butlin, R. K. (1999). Speciation without isolation. Nature, 400, 311-312.
  123. Van Valen, L. (1976). Ecological species, multispecies, and oaks. Taxon, 25, 233-239.
  124. Welzen, P. C. (1997). Paraphyletic groups or what should a classification entail. Taxon, 46, 99-103.
  125. Welzen, P. C. (1998). Phylogenetic versus Linnaean taxonomy, the continuing story. Taxon, 47, 413- 423.
  126. Wheeler, Q. D., & Meir, R. (Eds.) (2000), Species concepts and phylogenetic theory: A debate. New York: Columbia University Press.
  127. Wheeler, Q. D., & Nixon, K. C. (1990). Another way of looking at the species problem: A reply to de Queiroz and Donoghue. Cladistics, 6, 77-81.
  128. Wiley, E. O. (1978). The evolutionary species concept reconsidered. Systematic Zoology, 27, 17-26.
  129. Wilson, R. A. (Ed.) (1999), Species: New interdisciplinary essays. Cambridge: MIT Press.
  130. Woodger, J. H. (1952). From biology to mathematics. The British Journal for the Philosophy of Science, 3, 1-21.