Species concepts and the endangered species act: How a valid biological definition of species enhances the legal protection of biodiversity

Natural Resources Journal

Volumn 45, Issue 2, 2005, Pages 369-407

  George, Anna L ^a   Mayden, Richard L ^b  

^a FRANKLIN AND MARSHALL COLLEGE   (United States)

^b SAINT LOUIS UNIVERSITY   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

BIODIVERSITY; ENDANGERED SPECIES; ENVIRONMENTAL LEGISLATION; HIERARCHICAL SYSTEM;

EID: 28044457420     PISSN: 00280739     EISSN: None     Source Type: Journal    
DOI: None     Document Type: Review

References (310)

1. Jody Hey, Genes, Categories, and Species: The Evolutionary and Cognitive Causes of the Species Problems 32 (2001).
2. See Richard L. Mayden, On Biological Species, Species Concepts and Individuation in the Natural World, 3 Fish & Fisheries 171, 192 (2002).
3. Endangered Species Act of 1973,16 U.S.C. §§ 1531-1544 (2000).
4. See Daniel J. Rohlf, There's Something Fishy Going on Here: A Critique of the National Marine Fisheries Service's Definition of Species Under the Endangered Species Act, 24 Envtl. L. 617, 618 (1994).
5. See Richard L. Madyen, Consilience and a Hierarchy of Species Concepts: Advances Toward Closure on the Species Puzzle, 31 J. Nematology 95, 95 (1999).
6. The phrase "species concept" is used to avoid the limitations that are inherent within a set definition. As defined in Merriam-Webster's Collegiate Dictionary (1998), a concept is "1: something conceived in the mind: THOUGHT, NOTION 2: an abstract or generic idea generalized from particular instances." When a new species is discovered, the published document that formally recognizes and names this new species is referred to as "species description." Species can only be described, not defined, as a definition would set limits on something that is constantly changing. Species are identified in nature by various methods employed by scientists that are closely tied to the concept used. In this article, the term "species concepts" will be used during the discussions of biological conceptualizations of species, while "species definition" will be used during discussions of the legal definition of species.
7. See, e.g., Species Concepts and Phylogenetic Theory: A Debate (Quentin D. Wheeler & Rudolf Meier eds., 2000) (prominent scientists were invited to present a series of papers documenting their position on favored species concepts);
8. see also Edward O. Wiley, The Evolutionary Species Concept Reconsidered, 27 Systematic Zoology 17, 18 (1978);
9. Richard L. Mayden & Robert M. Wood, Systematics, Species Concepts, and the Evolutionary Significant Unit in Biodiversity and Conservation Biology, 17 Am. Fisheries Soc'y Symp. 58, 84 (1995). These examples are only a small selection of the wide-ranging literature available on these concepts.
10. See Ray Vaughan, State of Extinction: The Case of the Alabama Sturgeon and Ways Opponents of the Endangered Species Act Thwart Protection for Rare Species, 46 Ala. L. Rev. 569, 586 (1995).
11. 16 U.S.C. § 1533.
12. Id. § 1536.
13. Id. §§ 1538, 1540.
14. Id. § 1531(a)(3).
15. Id. § 1532.
16. Id. § 1533(a)(3).
17. Id. § 1533(b)(1)(A).
18. Id. § 1533(c).
19. Defined in § 1532(5)(A) to include the specific areas both within and outside the geographic range of the species, so long as it is "essential to the conservation of the species." The purchase of this land is covered by funds designated in § 1534(a); this critical habitat purchase has been the basis for the creation of many of the lands in the National Wildlife Refuge system.
20. Id. § 1533(f). Recovery plans contain descriptions of management actions, measurable criteria to determine when species may be removed from the list, and estimates of the time required and costs involved in reaching these goals.
21. Id. § 1536(a)(2).
22. Id. § 1536(c).
23. See Conservation Law Found. of New England, Inc. v. Andrus, 623 F.2d 712, 715 (Mass. App. Ct. 1979) (holding that ESA applies to applicants for leases for offshore drilling as well as the government).
24. 16 U.S.C. § 1538(e) (colloquially known as the "God Squad" due to the Committee's power to determine the fate of a species).
25. Id. § 1538(a)(1).
26. Id. § 1539(a)(2) (one such procedure may be the adoption of an approved Habitat Conservation Plan to mitigate the harm to the species and its critical habitat).
27. Id. § 1532.
28. Id. § 1532(19).
29. Endangered and Threatened Wildlife and Plants, 50 C.F.R. § 17.3 (2002).
30. Id.
31. 16 U.S.C § 1532(16).
32. See, e.g., John J. Wiens & Maria R. Servedio, Species Delimitation in Systematics: Inferring Diagnotic Differences Between Species, 267 Proc. Royal Soc'y London Series B 631, 632 (2000);
33. Jack W. Sites, Jr. & Jonathan C. Marshall, Delimiting Species: A Renaissance Issue in Systematic Biology, 18 Trends Ecology & Evolution 462, 462 (2003).
34. 471 F. Supp. 985 (D. Haw. 1979), aff'd, 639 F.2d 495 (9th Cir. 1981).
35. U.S. Fish & Wildlife Serv., Palila Recovery Plan 6 (1986).
36. As defined in 16 U.S.C. § 1532(5)(A)(i), "critical habitat" includes those geographic areas in the species' range "on which are found those physical or biological features (I) essential to the conservation of the species and (II) which may require special management consideration or protection." The critical habitat may not be established throughout the entire species range except in circumstances determined by the Secretary of the Interior. Thus, critical habitat is used to designate areas vital to the species' behavioral patterns, such as breeding or feeding.
37. Palila, 471 F. Supp. at 990.
38. The palila primarily fed upon the seeds and pods, so there was no direct competition for food in the mamane, but, eventually, heavy grazing by goats upon the forest would inhibit regrowth of the mamane to an extent that there were no seeds and pods produced for palila consumption. U.S. Fish & Wildlife Serv., supra note 32, at 10 (listing food sources for the palila);
39. Palila, 471 F. Supp. at 990 (listing mamane as a food source for goats).
40. At the time Palila was first brought to court, "harm," as used in the ESA's definition of "take," was defined as an act or omission which actually injures or kills wildlife, including acts which annoy it to such an extent as to significantly disrupt essential behavioral patterns, which include, but are not limited to, breeding, feeding, or sheltering; significant environmental modification or degradation which has such effects is included within the meaning of "harm"....
41. Palila, 471 F. Supp. at 989.
42. Id. at 990-91.
43. Id. at 995.
44. Palila, 639 F.2d 495, 495 (9th Cir. 1981).
45. Id. at 497.
46. Id. at 497-98.
47. Palila, 471 F. Supp. at 989;
48. Palila v. Haw. Dep't of Land & Natural Res., 649 F. Supp. 1070, 1073 (D. Haw. 1986), aff'd, 852 F.2d 1106 (9th Cir. 1988).
49. Endangered and Threatened Wildlife and Plants, Final Redefinition of "Harm," 46 Fed. Reg. 54,748 (Nov. 4, 1981).
50. Id.
51. Palila, 471 F. Supp. at 989;
52. Palila v. Haw. Dep't of Land & Natural Res., 649 F. Supp. 1070, 1073 (D. Haw. 1986), aff'd, 852 F.2d 1106 (9th Cir. 1988).
53. U.S. Fish & Wildlife Serv., supra note 32, at 6.
54. Palila, 649 F. Supp. at 1075.
55. Id. at 1077.
56. Id.
57. Palila, 852 F.2d at 1111.
58. Id. at 1110-11.
59. Id. at 1108.
60. Sweet Home Chapter of Cmtys. for a Great Or. v. Lujan, 806 F. Supp. 279, 282 (D.D.C. 1992), aff'd sub nom.
61. Sweet Home Chapter of Cmtys. for a Great Or. v. Babbitt, 1 F.3d 1 (D.C. Cir. 1993), rev'd,
62. 17 F.3d 1463 (D.C. Cir. 1994), rev'd sub nom.
63. Babbitt v. Sweet Home Chapter of Cmtys. for a Great Or., 515 U.S. 687 (1995).
64. Id.
65. Id.
66. Id.
67. Id. at 284.
68. Id.
69. Id.
70. Id. at 286.
71. Sweet Home, 1 F.3d at 1.
72. Id. at 5.
73. Id. at 11.
74. Sweet Home Chapter of Comtys. for a Greet Or., 17 F.3d at 1464.
75. Id.
76. Id. at 1472.
77. Id. at 1464
78. Id.
79. 1464-1465
80. Id. at 1465.
81. Sweet Home Chapter of Cmtys. for a Great Or. v. Babbitt, 30 F.3d 190, 191 (D.C. Cir. 1994).
82. Nat'l Wildlife Fed'n v. Burlington N. R.R., 23 F.3d 1508, 1513 (9th Cir. 1994).
83. Id. at 1512-13.
84. Seattle Audubon Soc'y v. Lyons, 871 F. Supp. 1291, 1313 (W.D. Wash. 1994).
85. See Sierra Club v. Lyng, 694 F. Supp. 1260 (E.D. Tex. 1988);
86. Swan View Coalition, Inc. v. Turner, 824 F. Supp. 923 (D. Mont. 1992).
87. Babbitt v. Sweet Home Chapter of Cmtys. for a Great Or., 515 U.S. 687 (1995).
88. Id. at 697.
89. Id. at 717-18 (Scalia, J., dissenting).
90. See id. at 721 (Scalia, J. dissenting).
91. Id. at 687.
92. Vaughan, supra note 8, at 605-06.
93. Endangered and Threatened wildlife and Plants; Final Rule to List the Alabama Sturgeon as Endangered, 65 Fed. Reg. 26,438, 26,443 (May 5, 2000) (list of endangered and threatened wildlife codified at 50 C.F.R. § 17.11 (2004)).
94. Jack E. Williams et al., Fishes of North America Endangered, Threatened, or of Special Concern: 1989, Fisheries, Nov.-Dec. 1989, at 2, 3.
95. "Described" is the scientific term for a formal recognition and naming of a new species in order to "present a general picture of the described taxon." Ernst Mayr, Principles of Systematic Zoology 265 (1969).
96. James E. Williams & Glenn H. Clemmer, Scaphirhynchus Suttkusi, a New Sturgeon (Pisces: Acipenseridae) from the Mobile Basin of Alabama and Mississippi, Ala. Museum Nat. Hist. Bull., June 1, 1991, at 17, 20.
97. Proposed Endangered Status and Designation of Critical Habitat for the Alabama Sturgeon, 58 Fed. Reg. 33,148 (June 15, 1993).
98. Alabama-Tombigbee Rivers Coalition v. Dep't of Interior, 26 F.3d 1103, 1105 (11th Cir. 1994).
99. See also Joe Nabbefeld, Controversial Report Backs Argument for Sturgeon, Birmingham News, Nov. 23, 1993.
100. Vaughan, supra note 8, at 608-09. Although the listing process itself does not require any economic consideration, the designation of critical habitat is made "on the basis of the best scientific data available and after taking into consideration the economic impact...of specifying any particular area as critical habitat." 16 U.S.C. § 1533(b)(2) (2000). Although this allows for some economic considerations, the Secretary must list areas that have economic importance if "the failure to designate such area as critical habitat will result in the extinction of the species concerned."
101. Although the listing process itself does not require any economic consideration, the designation of critical habitat is made "on the basis of the best scientific data available and after taking into consideration the economic impact...of specifying any particular area as critical habitat." 16 U.S.C. § 1533(b)(2) (2000). Although this allows for some economic considerations, the Secretary must list areas that have economic importance if "the failure to designate such area as critical habitat will result in the extinction of the species concerned."
102. Id.
103. Although the listing process itself does not require any economic consideration, the designation of critical habitat is made "on the basis of the best scientific data available and after taking into consideration the economic impact...of specifying any particular area as critical habitat." 16 U.S.C. § 1533(b)(2) (2000). Although this allows for some economic considerations, the Secretary must list areas that have economic importance if "the failure to designate such area as critical habitat will result in the extinction of the species concerned."
104. David Pace, Scientists Conclude Sturgeon Not Genetically Unique, Tuscaloosa News, Apr. 13, 1994, at 7B (implying that because this study was released only in a draft report through the FWS, it was not subjected to the unbiased peer review process required for publication in scientific journals).
105. Vaughan, supra note 8, at 606.
106. Id. For any animal, a lack of sighting for some years is not a clear sign of extinction. For a list of fish species that were presumed extinct and were then later rediscovered after periods longer than eight years,
107. see Richard L. Mayden & Bernard R. Kuhajda, Systematics, Taxonomy, and Conservation Status of the Endangered Alabama Sturgeon, Scaphirhynchus Suttkusi Williams and Clemmer (Actinopterygii, Acipenseridae), 1996 Copeia 241, 267.
108. Reopening of Comment Period, Public Hearing, and Availability of Panel Report on Proposed Endangered Status and Designation of Critical Habitat for the Alabama Sturgeon, 58 Fed. Reg. 55,036 (Oct. 25, 1993).
109. Alabama-Tombigbee Rivers Coalition v. Dep't of Interior, 26 F.3d 1103, 1105 (11th Cir. 1994).
110. Id. at 1105.
111. Federal Advisory Committee Act §§ 1-15, 5 U.S.C. app. 2 (2000).
112. See id. § 2.
113. Alabama-Tombigbee Rivers Coalition, 26 F.3d at 1105.
114. Joe Nabbefeld, Net Turns Up Elusive Sturgeon, but DNA Test Urged on Pedigree, Birmingham News, Dec. 4, 1993, at 1A.
115. See also Vaughan, supra note 8, at 624.
116. See Alabama-Tombigbee Rivers Coalition, 26 F.3d at 1107.
117. Vaughan, supra note 8, at 627.
118. Endangered and Threatened Wildlife and Plants, Extension of the Final Decision to List the Mobile River System Population of the Alabama Sturgeon as an Endangered Species, 59 Fed. Reg. 31,970, 31,973-74 (proposed June 21, 1994) (to be codified at 50 C.F.R. pt. 17).
119. Id.;
120. see also Pace, supra no 90.
121. Endangered and Threatened Wildlife and Plants; Withdrawal of Proposed Rule for Endangered Status and Critical Habitat for the Alabama Sturgeon, 59 Fed. Reg. 64,794 (proposed Dec. 15, 1994) (to be codified at 50 C.F.R. pt. 17).
122. Richard Shelby, Sturgeon Used in Political Game, Tuscaloosa News, June 6, 1999, at 6D.
123. Ala. Dep't of Conservation of Nat. Resources et al., Conservation Agreement & Strategy for the Alabama Sturgeon 4 (2000).
124. Mayden & Kuhajda, supra note 92, at 241-73.
125. Endangered and Threatened Wildlife and Plants; Final Rule to List the Alabama Sturgeon as Endangerd. 65 Fed. Reg. 26,440 (May 5, 2000) (list of endangered and threatened wildlife codified at 50 C.F.R. § 17.11).
126. Endangered and Threatened Wildlife and Plants; Proposed Rule to List the Alabama Sturgeon as Endangered, 64 Fed. Reg. 14,676 (proposed Mar. 26, 1999) (to be codified at 50 C.F.R. pt. 17).
127. Michael Sznajderman, Big Fuss Over an Alabama Fish, Birmingham News, Aug. 11, 1999, at 1A 6A;
128. see also Shelby, supra note 106.
129. Endangered and Threatened Wildlife and Plants; Final Rule to List the Alabama Sturgeon as Endangered. 65 Fed. reg. 26,448 (May 5, 2000) (list of endangered and threatened wildlife codified at 50 C.F.R. § 17.11).
130. Id. at 26,438.
131. Id. at 26.448.
132. 16 U.S.C. § 1533 (b)(1)(A) (2000).
133. See Tenn. Valley Auth. v. Hill, 437 U.S. 153 (1978).
134. Public support for more charismatic organisms helps create a bias in conservation projects even more severe than that seen in taxonomic work. See J. Alan Clark & Robert M. May, Taxonomic Bias in Conservation Research, 297 Science 191, 191 (2002).
135. Kevin D. Hill, The Endangered Species Act: What Do We Mean by Species?, 20 B.C. Envtl. Aff. L. Rev. 239, 263 (1993) ("If the Act is meant to do more than just protect charismatic megafauna who are valued for their sentimental appeal, the definition of species must reflect the need to protect biodiversity.").
136. 16 U.S.C. § 1532(16) (2000).
137. See generally Joel Cracraft, Species Concepts in Theoretical and Applied Biology: A Systematic Debate with Consequences, in Species Concepts and Phylogenetic Theory: A Debate, supra note 7, at 3;
138. Species: New Interdisciplinary Essays (Robert A. Wilson ed., 1999).
139. Mayden, supra note 5, at 103.
140. See id.;
141. R.L. Mayden, A Hierarchy of Species Concepts: The Denouement in the Saga of the Species Problem, in Species: The Unit's of Biodiversity 381, 388 (M.F. Claridge et al. eds., 1997).
142. Mayr, supra note 85, at 56.
143. Id. at 66-67.
144. Id. at 57-58.
145. For example, the Scaphirhynchus suttkusi is the Alabama sturgeon.
146. Mayr, supra note 85, at 58.
147. "Essentialism" refers to a classification where kinds are grouped together based upon possession of a defined attribute or property. Such a property is both necessary and sufficient for being a member of the kind. See, e.g., David L. Hull, Are Species Really Individuals?, 25 Systematic Zoology, 176 (1976);
148. Robert A. Wilson, Realism, Essence, and Kind: Resuscitating Species Essentialism?, in Species: New Interdisciplinary Essays, supra note 120, at 187, 188.
149. See Marc Ereshefsky, Species and the Linnean Hierarchy, in Species: New Interdisciplinary Essays 285, 287 (Robert A. Wilson ed., 1999).
150. Mayden & Wood, supra note 7, at 84.
151. Hull, supra note 128, at 180.
152. E.O. Wiley, Phylogenetics: The Theory and Practice of Phylogenetic Systematic 8 (1981).
153. Id.
154. Id. at 7.
155. See Mayden, supra note 2, at 181.
156. See John Dupré, On the Impossibility of a Monistic Account of Species, in Species: New Interdisciplinary Essays, supra note 120, at 3. The pre-Darwinian understanding of species was of an essentialist concept (i.e., species are immutable over time). However, adoption of evolutionary theory meant that there were no sharp demarcations between species (i.e., they could change during time, and at different rates). Thus, some argue, our current concepts are unable to account for species because they have not fully incorporated the change associated with evolution.
157. See Mayden, supra note 2, at 176.
158. See, e.g., Species: The Units of Biodiversity, supra note 122;
159. Endless Forms: Species and Speciation (Daniel J. Howard & Stewart H. Berlocher eds., 1998);
160. Speciation and the Recognation Concept: Theory and Application (David M. Lambert & Hamish G. Spencer eds., 1995);
161. Hey, supra note 1;
162. Species Concepts and Phylogenetic Theory: A Debate, supra note 7.
163. Mayden, supra note 122, at 389.
164. Id. at 387.
165. Id. at 388.
166. Hull, supra note 128, at 177.
167. See also Mayden, supra note 2, at 179.
168. This occurs both in biology and law; the majority of debates surrounding how a species is defined focus on the empirical methods used for a research study, rather than question the conceptualization of species. See, e.g., Kevin W. Grierson, The Concept of Species and the Endangered Species Act, 11 Va. Envtl. L.J. 463, 486 (1992);
169. Alan R. Templeton, The Meaning of Species and Speciation: A Genetic Perspective, in Speciation and Its Consequences 3, 3 (Daniel Otte & John A. Endler eds., 1989) ("[A] species concept can be evaluated only in terms of a particular goal or purpose," indicating a use of species concepts only to further the study of a subdiscipline in biology, not to understand philosophically what a species is.").
170. A character is "a feature (attribute, observable part) of an organism." Wiley, supra note 132, at 8.
171. Mayden, supra note 5, at
172. See Mayden, supra note 2, at 184. Species descriptions should be treated as hypotheses of lineage independence, to be empirically tested. Without operational criteria for testing, biologists may learn little from studying arbitrarily defined taxa with no uniting theoretical basis.
173. Mayden & Wood, supra note 7, at 94.
174. Id. at 95. For example, a hypothetical species concept that only accepts as valid species differences in consistent patterns of female coloration would ignore the diversity that becomes apparent seasonally when breeding males exhibit different colors within the recognized species. The operation used is strictly based upon the conception of reality of the user of that operation-a conception that may be artificial or real for that species (e.g., colors perceived by Homo sapiens may not be comparable to colors perceived by the species we study).
175. See Wiley, supra note 7, at 18. For example, a hypothetical species concept that only accepts differences in females as valid species would ignore diversity that becomes apparent when breeding males take on different colors.
176. This is often referred to as cryptic biodiversity. See Robert S. Butler & Richard L. Mayden, Cryptic Biodiversity, 28 Endangered Species Bull. 24, 24 (2003).
177. Ernst Mayr, A Critique from the Biological Species Concept Perspective: What Is a Species, and What Is Not?, in Species Concepts and Phylogenetic Theory: A Debate, supra note 7, at 93, 97.
178. Id.
179. See Theodosius Dobzhansky, Mendelian Populations and Their Evolution, 84 Am. Naturalist 401, 405 (1950) (discussing the genetic species concept);
180. Ernst Mayr, Speciation Phenomena in Birds, 74 Am. Naturalist 249, 256 (1940) (discussing the biological species concept);
181. C.T. Regan, Organic Evolution, 1926 British Association for Advancement of Science Report of Ninety-Third Meeting 75 (discussing the morphological species concept).
182. Mayden, supra note 5, at 110.
183. Id. at 111;
184. see also Peter C.H. Pritchard, Status of the Black Turtle, 13 Conservation Biology 1000, 1003 (1999). Pritchard states that it is "vital that systematists avoid dependence upon a single 'favored technique' for answering questions relating to the rank of related taxa and instead utilize all the tools at their disposal."
185. Id. This conclusion stems from the debate over the status of the black turtle, which is known to be distinct from the green turtle based upon consistent morphological differences, despite few genetic differences.
186. The BSC is widely used among ornithologists and mammalogists, partly due to the influence that the author of this concept has among specialists in these groups. See Cracraft, supra note 120, at 4. This is also the concept commonly introduced in introductory biological textbooks.
187. See, e.g., Sylvia Mader, Biology 1, 332 (4th ed. 1993);
188. Cecie Starr & Ralph Taggart, Biology: The Unity and Diversity of Life 2, 298 (8th ed. 1998).
189. The BSC contains both a genetic and ecological component and is defined as: a group of populations which replace each other geographically or ecologically and of which the neighboring ones intergrade or hybridize wherever they are in contact or which are potentially capable of doing so (with one or more of the populations) in those cases where contact is prevented by geographical or ecological barriers. Mayr, supra note 151.
190. Mayden & Wood, supra note 7, at 96.
191. Marty Bergoffen, Endangered Species Act Reauthorization: A Biocentric Approach 11 (1995).
192. Hill, supra note 118, at 262.
193. The reproductive isolation test is particularly troubling when used on species that do not even occur in overlapping regions. Interbreeding in nature will not occur naturally without outside intervention on the part of the researcher, and it makes little sense for biologists to test for possibility of hybridization by forcing individuals from disparate areas together in a laboratory setting.
194. See Niles Lehman et al., Introgression of Coyote Mitochondrial DNA into Sympatric North American Gray Wolf Populations, 45 Evolution 104, 108 (1991).
195. Mayden & Wood, supra note 7, at 97;
196. Paul-Michael Agapow et al., The Impact of Species Concept on Biodiversity Studies, 79 Q. Rev. Biology 161, 163 (2004).
197. Unfortunately, microbiology is still largely uncharted territory in biology; current estimates of diversity range from 105 to 107 species. Part of this is due to the ability for bacteria to rapidly accumulate genetic mutations that lead to new species.
198. See William B. Whitman et al., Prokaryotes: The Unseen Majority, 95 Proc. Nat'l Acad. Sci. 6578, 6582 (1998).
199. See Mayden & Wood, supra note 7, at 97.
200. Mayden, supra note 122, at 396.
201. Wiley, supra note 7, at 18.
202. Id. at 21.
203. Edward O. Wiley & Richard L. Mayden, The Evolutionary Species Concept, in Species Concepts and Phylogenetic Theory: A Debate, supra note 7, at 70, 73.
204. Jack W. Sites, Jr. & Keith A. Crandall, Testing Species Boundaries in Biodiversity Studies, 11 Conservation Biology 1289, 1292 (1997).
205. Mayr, supra note 151, at 97.
206. Id.
207. Id. at 97-98.
208. Id. at 98.
209. Edward O. Wiley & Richard L. Mayden, A Defense of the Evolutionary Species Concept, in Species Concepts and Phylogenetic Theory: A Debate, supra note 7, at 198, 200.
210. Mayden, supra note 122, at 405. Phylogenetic systematics is a specific approach to systematics that studies the history of speciation; the recovery of evolutionary relationships is then used to guide classification schemes.
211. See Wiley, supra note 132, at 6.
212. "Monophyly" is a term denoting a group that shares a common ancestor, thus, when used in a species concept, all members of that species should share a most recent common ancestor, with no members excluded.
213. See Donn E. Rosen, Vicariant Patterns and Historical Explanation in Biogeography, 27 Systematic Zoology 159, 176 (1978).
214. K.C. Nixon & Quentin D. Wheeler, An Amplification of the Phylogenetic Species Concept, 6 Cladistics 211, 218 (1990).
215. See Mary C. McKitrick & Robert M. Zink, Species Concepts in Ornithology, 90 Condor 1, 2 (1988).
216. Mayden & Wood, supra note 7, at 102.
217. Brent D. Mishler & Edward C. Theriot, The Phylogenetic Species Concept (sensu Mishler and Theriot): Monophyly, Apomorphy, and Phylogenetic Species Concepts, in Species Concepts and Phylogenetic Theory: A Debate, supra note 7, at 44, 45.
218. Mayr, supra note 151, 99.
219. Id.
220. Id. 100.
221. Stephen A. Karl& Brian W. Bowen, Evolutionary Significant Units Versus Geopolitical Taxonomy: Molecular Systematics of an Endangered Sea Turtle (Genus Chelonia), 13 Conservation Biology 990, 96 (1999).
222. Id.
223. Id.
224. Id. at 994,
225. Black turtles have darker pigmentation patterns, a smaller size, and a different shell shape than green turtles.
226. See Pritchard, supra note 155, at 1001-02.
227. Karl& Bowen, supra note 186, at 996. Karl and Bowen "propose the label geopolitical species (GS) for taxonomic designations that persist, but for which there are essentially no supporting data."
228. Id.
229. H.R. Rep. No. 93-412, at 10 (1973).
230. S. Rep. No. 96-151, at 7 (1979).
231. See Kristin Shrader-Frechette & Earl D. McCoy, Molecular Systematics, Ethics, and Biological Decision Making Under Uncertainty, 13 Conservation Biology 1008, 1011 (1999).
232. 16 U.S.C. § 1532(16) (2000).
233. Determination of certain bald eagle populations as endangered or threatened, 43 Fed. Reg. 6230 (Feb. 14,1978).
234. See Robin S. Waples, Evolutionary Significant Units and the Conservation of Biological Diversity Under the Endangered Species Act, 17 Am. Fisheries Soc'y Symp. 8, 20 (1995).
235. S. Rep. No. 96-151, at 7 (1979).
236. Policy Regarding the Recognition of Distinct Vertebrate Population Segments Under the Endangered Species Act, 61 Fed. Reg. 4722 (Feb. 7, 1996).
237. Id. at 4722.
238. Robin S. Waples, Pacific Salmon, Oncorhynchus spp., and the Definition of "Species" Under the Endangered Species Act, 53 U.S. Nat'l Marine Fisheries Serv, Marine Fisheries Rev. 11, 15 (1991).
239. See, e.g., Rohlf, supra note 4, at 637;
240. David S. Pennock & Walter W. Dimmick, Critique of the Evolutionary Significant Unit as a Definition for "Distinct Population Segments" Under the U.S. Endangered Species Act, 11 Conservation Biology 611, 615 (1997).
241. Waples, supra note 202, at 12.
242. Mayden & Wood, supra note 7, at 109.
243. See Waples, supra note 202, at 13.
244. Id. at 14.
245. Robin S. Waples, Evolutionary Significant Units, Distinct Population Segments, and the Endangered Species Act: Reply to Pennock and Dimmick, 12 Conservation Biology 718, 719 (1998).
246. See Mayden & Wood, supra note 7, at 109.
247. Martha Rojas, The Species Problem and Conservation: What Are We Protecting?, 6 Conservation Biology 170, 174 (1992). By viewing a species as a "work in progress," management decisions can be made by looking at a species' ability to change and adaptabilities of greater importance in the natural world than the ability to maintain status quo. For example, listing decisions could be based upon the variability in a population, an indication of greater capacity to adapt than size of a population alone.
248. See Waples, supra note 198, at 9.
249. See id. at 14.
250. See Grierson, supra note 143, at 489.
251. See Mayden & Wood, supra note 7, at 110.
252. Prior to its extinction in 1987.
253. See Endangered and Threatened Wildlife and Plants; Final Rule to Delist the Dusky Seaside Sparrow and Remove Its Critical Habitat Designation, 55 Fed. Reg. 51,002, 51,113 (Dec. 12, 1990) (codified at 50 C.F.R. pt. 17).
254. Id. at 51,112.
255. John C. Avise & William S. Nelson, Molecular Genetic Relationships of the Extinct Dusky Seaside Sparrow, 243 Science 646, 648 (1989).
256. For another example of pigmentation differences and species status, see Pritchard, supra note 155, at 1001, regarding the black turtle.
257. Mayden, supra note 122, at 418.
258. Id. at 419.
259. Id.
260. Mayden, supra note 5, at 144.
261. This debate is not limited to either the biological or legal professions; both sides have advocates who feel that this problem would disappear if a single genetic criterion were used to establish management units, subspecies, or species status.
262. See Grierson, supra note 143, at 486, 487;
263. C. Moritz, Applications of Mitochondrial DNA Analysis in Conservation: A Critical Review, 3 Molecular Ecology 401, 405 (1994).
264. T.M. Caro & Gillian O'Doherty, On the Use of Surrogate Species in Conservation Biology, 13 Conservation Biology 805,810 (1999).
265. See Shelby, supra note 106, at 6D;
266. see also Nick J.B. Isaac et al., Taxonomic Inflation: Its Influence on Macroecology and Conservation, 19 Trends Ecology & Evolution 464, 466 (2004).
267. Janice S. Golding & Jonathan Timberlake, How Taxonomists Can Bridge the Gap Between Taxonomy and Conservation Science, 17 Conservation Biology 1177, 1178 (2003).
268. Mayden, supra note 122, at 419.
269. Ernst Mayr, Species Concepts and Definitions, in The Species Problem, 1, 10 (1957).
270. Mayden, supra note 122, at 418.
271. Id. at 419.
272. Id.
273. See id. at 420.
274. Id. at 421.
275. Id.
276. The largest problem related to many species concepts is their inability to correctly diagnose asexual species. The Biological Species Concept is unable to identify diversity within asexual groups as it uses the criterion of interbreeding, an activity that is not found in these species. Thus, the primary species concept must allow for the existence of these species rather than ignoring their existence because they do not fit the "normal" expectations of natural groups.
277. Mayden, supra note 122, at 419.
278. Id.
279. As mentioned above, the Biological Species Concept does not recognize asexual species, so it should not be used in any study examining asexual organisms.
280. Sites & Marshall, supra note 30, at 462.
281. Mayden, supra note 2, at 190.
282. A. Townsend Peterson & Adolfo G. Navarro-Sigüenza, Alternate Species Concepts as Bases for Determining Priority Conservation Areas, 13 Conservation Biology 427, 429 (1999);
283. see also Helen M. Regan et al., A Taxonomy and Treatment of Uncertainty for Ecology and Conservation Biology, 12 Ecological Applications 618, 619 (2002).
284. See, e.g., Mayr, supra note 149, at 96.
285. Agapow et al., supra note 161, at 169.
286. Mayden, supra note 2, at 187.
287. Id.;
288. see, e.g., Alan R. Templeton, Using Phylogeographic Analyses of Gene Trees to Test Species Status and Processes, 10 Molecular Ecology 779, 780 (2001);
289. John J. Wiens & Tonya A. Penkrot, Delimiting Species Using DNA and Morphological Variation and Discordant Species Limits in Spiny Lizards (Sceloporus), 51 Systematic Biology 69, 86 (2002).
290. Morphological characters, such as color or body shape, though easiest for non-scientists to use because they do not require sophisticated techniques, tend to be disdained because they may change over an individual's lifetime or are environmentally controlled through diet or temperature.
291. Grierson, supra note 143, at 489.
292. See, e.g., Amy R. McCune & Nathan R. Lovejoy, The Relative Rate of Sympatric and Allopatric Speciation in Fishes: Tests Using DNA Sequence Divergence Between Sister Species and Among Clades, in Endless Forms: Species and Speciation, supra note 138, at 172, 176.
293. Mitochondrial DNA is found in the mitochondria, the cellular organelle that is responsible for producing much of the energy used for cellular activities. It is inherited through the maternal line.
294. Picking the most rapidly evolving portion of DNA is useful because differences in composition will appear quickly, ideally replicating differences in the lineages of the two species.
295. Jeannette Krieger & Paul A. Fuerst, Evidence for a Slowed Rate of Molecular Evolution in the Order Acipenseriformes, 19 Molecular Biology & Evolution 891, 896 (2002);
296. see also Mayden, supra note 2, at 188-89.
297. See Krieger & Fuerst, supra note 251, at 891, 895-96,
298. see also Andrew M. Simons et al., Phylogenetics of Scaphirhynchus Based on Mitochondrial DNA Sequences, 130 Transactions Am. Fisheries Soc'y 359, 364 (2001). Sturgeons, a controversial endangered group, offer a prime example of how genetic tests should not be the sole method to evaluate species status. Alabama sturgeon, found only in the Mobile Basin, and shovelnose sturgeon, found only in the Mississippi Basin, have been found to have low genetic divergence as seen in two mitochondrial genes, despite obvious morphological differences. However, the two river basins have been isolated from each other for one to five million years, indicating that in this instance mitochondrial DNA does not necessarily evolve rapidly and, thus, should not be used as the sole benchmark for species divergence.
299. See Walter R. Hoeh et al., Multiple Origins of Gender-Associated Mitochondrial DNA Lineages in Bivalves (Mollusca: Bivalvia), 50 Evolution 2276, 2278 (1996). Within the past decade, evolutionary studies of mussels have uncovered this phenomenon, known as gender-associated mitotypes. Because mitochondrial DNA is typically inherited through the maternal line, there is usually only one form of it found in any individual. However, within these mussels, an unusual type of mitochondrial DNA inheritance leads to certain individuals in a population having both forms of the DNA. These two forms may be quite genetically distinct; there may be up to 33% divergence between the two genomes. In this instance, using a single criterion as a means to evaluate species status would result in an overestimate of actual biodiversity.
300. Using one gene to distinguish species can be similar to using one sentence to distinguish papers. In other words, just because two fairy tales begin with the sentence "Once upon a time..." does not mean that they will end the same way.
301. Manuel Lujan, Secretary of the Interior for the first Bush administration, stated, "Nobody's told me the difference between a red squirrel, a black one or a brown one." John Lancaster, Lujan: Endangered Species Act "Too Tough," Needs Changes, Wash. Post, May 12, 1990, at A1.
302. Of the 1.4 million currently recognized species, see Edward O. Wilson, The Diversity of Life 13-33 (1992), 4450 of these are mammals.
303. Calculated as a percentage of the currently listed animal species out of 1078 listed animal species, 349 are mammals. Mammals still account for 19% of the listed species when combined animal and plant species (1827 listed) are used. See U.S. Fish & Wildlife Serv., Species Information: Threatened and Endangered Animals and Plants, available at http://endangered.fws.gov/wildlife.html#Species (last visited May 23, 2005).
304. Wilson, supra note 256, at 132 (Wilson estimates that between 10 and 100 million species may be present on Earth.).
305. See, e.g., Karl & Bowen, supra note 186, at 997;
306. Stephen J. O'Brien & Ernst Mayr, Bureaucratic Mischief: Recognizing Endangered Species and Subspecies, 251 Science 1187, 1187 (1991);
307. Vaughan, supra note 8, at 586.
308. See Shrader-Frechette & McCoy, supra note 195, at 1009-10.
309. Robert J. Taylor, Biological Uncertainty in the Endangered Species Act, 8 Nat. Resources & Env't 6, 6 (1993).
310. 16 U.S.C. § 1532(16) (2000).