Ornithology, Evolution, and Philosophy 123

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A Modern Unified Theory of Evolution 213 (1985) “recognition concept” of species. Evolutionists need to distinguish between genetic isolation, reproductive isolation, and ecological isolation of species (Bock 1979, 1986, 2004d). Genetic isolation must be complete before the neospecies establish secondary contact in order not to hybridize, but reproductive and ecological isolation may be perfected in sympatry through natural selection and mutual selective demands. The “horizontal” biological species concept refers to genetically separated groups of populations which live during a particular time plane like the Present or any time plane of the geological past (Fig. 5.3). Under this concept, species have, strictly speaking, no origin, age or duration. They represent horizontal “cross sections” of vertical phyletic species lineages in the time dimension (Bock 1979, 1986, 1992a; Szalay and Bock 1991). A phyletic lineage is the continuum of a species intimeanddocumentsitshistory;itdoesnotparticipateinthedevelopmentof species (populations do). The “phyletic lineage” and the “species” should be kept separate conceptually. Mayr’s (1931b, 1942e) consistent application of the superspecies concept in his systematic work was of general significance because it permitted the combination of several closely related geographical representatives under one superspecific name without reducing these allo- or paraspecies to subspecies status. Such ‘superlumping’ had been advocated by several workers (e.g., Kleinschmidt and Fig.5.3. Several imaginary phyletic lineages illustrate “species” limits under the cladistic concept (clad.) and the paleontological concept (pal.). Schematic representation. Groups of populations representing the various lineages at particular time levels (t1–t4) aredifferent biological species (oval circles). Vertical scale–geological time; horizontal scale– morphological and other biological changes. A–L represent paleontological “species,” except C–F, which together are one paleontological “species” but represent 2 cladistic “species.” The current time level is t4
214 5 Biological Species and Speciation—Mayr’s First Synthesis Stresemann) during the 1920s, even though many of such allopatric forms actually were too distinct to be treated as subspecies. The units of a regional fauna are zoogeographical species (Bock 1956, Mayr and Short 1970f) composed of two elements, (a) taxonomically isolated species (isospecies; that are not members of superspecies) and (b) superspecies (Amadon 1966; Bock and Farrand 1980; Amadon and Short 1992). Much earlier Mayr had already emphasized that one must count zoogeographical species rather than individual species in a particular region of the world, in order to avoid misleading implications. In his letter to Stresemann dated 29 September 1936 he wrote: “A renewed count of the birds of New Guinea resulted in 513 breeding zoogeographical species (reckoning [the superspecies] Astrapia and Parotia as one species each).” However, it was not until 1970 that Mayr (l.c.) introduced the term “zoogeographical species” formally (actually already used in Bock 1956). He himself credited it later to B. Rensch (Mayr 1989k: 156) without a specific reference. However, Rensch (1934) had used the term species geographicum only in the sense of polytypic species (not in the sense of superspecies plus taxonomically isolated species). In their analysis of zoogeographic species Mayr and Short (1970f) studied the avian species taxa of a well-defined region (North America) to answer the question of how often the application of the biological species concept leads to difficulties, controversies, or ambiguities presenting in full their data on which their conclusions were based. Most of the tabulation and accompanying text was written by L. Short and evaluated in discussions with Mayr. They showed that invariably the application of the biological species concept helped to clarify difficult situations. In a number of peripheral isolates one could not be sure whether or not they had already reached species level, but this created as much difficulty for a morphological as for the biological species concept. Only in Mexican populations of towhees, Pipilo, the same two forms behave in one area as two biospecies and in another area as one because of extensive hybridization. Several other cases of “interspecific hybridization between largely allopatric members of the same superspecies” are not clear-cut either, especially those where there are large zones of geographical range overlap and extensive hybridization. The theoretical concept of biological species is nondimensional and refers most clearly to genetic-reproductively isolated populations at a particular locality (Mayr 1941i, 1942e, 1946l, 1953b, 1963b). Because a fully differentiated biospecies represents a genetic unit, a reproductive unit and an ecological unit, Mayr (1951l: 92, 1982d: 273) specified as one aspect of biological species a specific niche in nature, i.e., ecological isolation permitting sympatry with competitors. This theoretical notion of biological species must be distinguished from the multidimensional species taxon. If several differentiated groups of populations are in contact and intergrade, they belong to the same species taxon (Fig. 5.4). Allopatric, i.e., geographically separated, representative taxa are assigned subspecies or species status on the basis of inference (Mayr 1948c, Mayr et al. 1953a: 104, 1969b: 197, Mayr and Ashlock 1991i: 104–105). Auxiliary criteria used for that purpose include:
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Jürgen Haffer Ornithology, Evoluti
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Preface Ernst Mayr (1904-2005), emi
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Contents Introduction 1 Part I The
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Contents IX 2 Bibliography of Ernst
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2 Introduction and speciation, the
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4 Introduction (1928-1929) to repla
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Ernst Mayr has been an observer all
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The Family 9 court system. He had a
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The Family 11 Fig.1.1. Family genea
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The Family 13 where they learned La
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Birdwatching 15 even finishing befo
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Birdwatching 17 Fig.1.5. ErnstMayr
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Birdwatching 19 Fig.1.6. Ernst Mayr
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Birdwatching 21 he simply smoked ci
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University Student in Greifswald an
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Friendships at School and Universit
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2 The Budding Scientist PhD Thesis
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TheInfluenceofTeachers 37 excellent
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Erwin Stresemann—Teacher and Frie
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Erwin Stresemann—Teacher and Frie
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Assistant Curator at the Museum of
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Expeditions to New Guinea and the S
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Part II Ornithologist and Evolution
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96 3 The New York Years (1931-1953)
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98 3 The New York Years (1931-1953)
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100 3 The New York Years (1931-1953
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4 Ornithologist and Zoogeographer B
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142 4 Ornithologist and Zoogeograph
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Page 167 and 168: 162 4 Ornithologist and Zoogeograph
Page 189 and 190: 184 5 Biological Species and Specia
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Page 261 and 262: 256 7 The Harvard Years (1953-2005)
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264 7 The Harvard Years (1953-2005)
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8 Evolutionary Biology— Mayr’s
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268 8 Evolutionary Biology—Mayr
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284 9 Ernst Mayr—the Man his coll
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286 9 Ernst Mayr—the Man addresse
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288 9 Ernst Mayr—the Man undertak
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290 9 Ernst Mayr—the Man unaware
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292 9 Ernst Mayr—the Man His hand
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294 9 Ernst Mayr—the Man Prize (1
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296 9 Ernst Mayr—the Man the inte
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298 9 Ernst Mayr—the Man I have s
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300 9 Ernst Mayr—the Man Fig.9.4.
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302 9 Ernst Mayr—the Man not mali
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304 9 Ernst Mayr—the Man papers d
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306 9 Ernst Mayr—the Man 2005. Me
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308 9 Ernst Mayr—the Man Fig.9.5b
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310 9 Ernst Mayr—the Man health i
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312 9 Ernst Mayr—the Man person a
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314 9 Ernst Mayr—the Man with a r
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316 9 Ernst Mayr—the Man conceptu
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318 9 Ernst Mayr—the Man the two
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10 Systematics and Classification T
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322 10 Systematics and Classificati
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11 History and Philosophy of Biolog
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340 11 History and Philosophy of Bi
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376 12 Summary: Appreciation of Ern
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Acknowledgments Iamdeeplygratefulto
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List of Abbreviations AAAS American
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386 References Beehler BM, Pratt TK
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388 References Cracraft J (2002) Go
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390 References Haffer J (1997a) Ess
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392 References Keast JA (1996) Avia
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394 References Olson SL (1975) The
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396 References Smith JJ (1936) Eric
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398 References Wiglesworth LW (1898
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Appendix 1 Curriculum Vitae (includ
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Appendix 1 403 1978 Prêmio Jabuti,
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Appendix 1 405 Society Offices Linn
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Appendix 2 407 1926c Das Freibrüte
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Appendix 2 409 1932 1932a Birds col
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Appendix 2 411 1936g Melidectes bel
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Appendix 2 413 1941e Birds collecte
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Appendix 2 415 1944f Chromosomes an
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Appendix 2 417 1947g On the correct
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Appendix 2 419 1952e Six new subspe
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Appendix 2 421 1954g The tail molt
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Appendix 2 423 1958c Concluding rem
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Appendix 2 425 in E. Mayr, Evolutio
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Appendix 2 427 1964l Introduction [
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Appendix 2 429 1967d Subfamily Pach
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Appendix 2 431 in Acta Soc. Zool. B
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Appendix 2 433 1974b The diversity
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Appendix 2 435 1976m Evolution and
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Appendix 2 437 Mankind 13:430; P.W.
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Appendix 2 439 (1984) in Queen’s
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Appendix 2 441 1984c Evolution and
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Appendix 2 443 1987b The status of
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Appendix 2 445 1990 1990a The Myth
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Appendix 2 447 1992f Haldane’s Ca
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Appendix 2 449 1995g The search for
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Appendix 2 451 1999 1999a Thoughts
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List of Books by Ernst Mayr 453 200
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Coauthorship 455 Coauthorship In th
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Appendix 3 457 (4) Systematics 1942
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Index Achievements 379 Adaptation 2
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Index 461 Faunas 164-165 Family 8-1
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Index 463 Realism and education 372
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