Ornithology, Evolution, and Philosophy 123

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A Modern Unified Theory of Evolution 221 (B) Without a speciation event (parental species transformed) (c) Dichopatric speciation (split by a geographic barrier with gradual divergence) (d) Gradual phyletic transformation of a single lineage. The crucial process in allopatric speciation is geographic isolation (not selection). This statement only refers to the geographic position and separation of the speciating populations. It still remains completely unknown as to what happens genetically during speciation. Possibly rather different genetic mechanisms are involved in the speciation of different kinds of organisms and under different circumstances.Ofcourse,thespeciatingpopulationsremainwelladaptedthrough naturalselectionwhichisalwayspresent. Some authors have suggested that Sewall Wright’s writings have influenced Mayr’s thinking and that in particular, his model of the “adaptive landscape” had given Mayr the idea of geographic speciation. 13 Nothing could be further from the truth. Geographic speciation was discussed among naturalists since the 19th century (p. 230) and was standard thinking in Erwin Stresemann’s department in Berlin; Mayr described geographical speciation already in 1927(f) in his paper on the Rosy-Finches (Leucosticte). Moreover, Wright’s model of the “adaptive landscape” was an abstract model, not one of a three-dimensional landscape; it hasverylittletodowithspeciation,butrefersmainlyto“theevolutionofthe species as a whole” (quote in Mayr 1992i: 11). As Mayr (1999k: XXIV) pointed out, S. Wright was never particularly interested in problems of speciation and, although he published reviews of all the major works on evolution, the only one he omitted was Mayr’s Systematics and the Origin of Species (1942e). Species originate through “splitting” and “budding” (Fig. 5.6). “Budding” occurs when, e.g., a derivative population of a widespread mainland species reached species status on a nearby island. This speciation event had no effect on the parental biospecies (no. 3, Fig. 5.6) on the mainland from which neospecies 4 has budded off. The mainland species is real in the sense that it represents a biological unit characterized by close genetic-reproductive and ecological relations among its component subspecies taxa. The cladistic analyses schematically illustrated in Fig. 5.6 (if feasible at that intraspecific level) yield relevant phylogenetic (“vertical”) and biogeographical data on the origin and relationships of the various groups of taxa. Mayr (e.g., 2000f: 164) feels that most new species originate by budding, splitting is less common. One may ask the question why Mayr, in 1942, preferred the dichopatric model of speciation (the “dumbbell” model), although the island populations of the Polynesian and Melanesian birds he studied in detail certainly had originated through jump dispersal (and peripatric speciation). When he studied a particular aberrant peripheral island population, the question foremost in his mind at that 13 Thus M. Ruse (1999: 118) wrote: “Mayr, who in later years put distance between himself and Wright, based his original picture of the evolutionary process on the hypothesis” (the shifting balance hypothesis of S. Wright).
222 5 Biological Species and Speciation—Mayr’s First Synthesis Fig.5.6. Speciation through splitting (A) and budding (B) resulting in monophyletic biospecies 1 and 2 (consisting of 3 and 2 subspecies, respectively) and paraphyletic biospecies 3 (three subspecies). Species 4 which budded off from species 3 is monotypic and may demonstrate its species status by reinvading the ranges of some or all subspecies of species 3. Shading indicates genetic cohesion and intergradation of subspecies along contact zones. From Haffer (1992) time was a taxonomic one, whether this form should be ranked as a subspecies, species or as a monotypic genus. The evolutionary questions why and how these aberrant forms originated on isolated islands were not yet foremost in his mind. Another question is to what extent his fieldwork in New Guinea and the Solomon Islands influenced Mayr’s ideas as developed and promoted in his 1942 volume and in later publications. He gave the following answer in an interview: “The thing is that the influence [of my fieldwork] was only limited because I already had most of the basic ideas. They had been developed further in recent years by Stresemann and several other workers in this area including in Russia. Naturally, everything I had done and observed in New Guinea and the Solomon Islands fitted exactly on these basic ideas of the European workers, but they never had developed all the ideas in these areas: What are the possibilities? What are the methods to be used? What are the mistakes you might make?” (Bock and Lein 2005, video CD-ROM). Mayr’s 1942 book had an enormous influence on the development of evolutionary studies worldwide, but various general questions still remain open: What is the percentage of species of major taxonomic units (e.g., birds or mammals) that originated through peripatric (“budding”) and dichopatric speciation (“splitting”), respectively? To what extent did peripatric speciation take place on continents? How many of the taxonomic species currently recognized are, in a cladistic sense, monophyletic and how many are paraphyletic? 14 And how important is sympatric speciation? 14 Monophyletic species contain all descendants of a common ancestor and paraphyletic species contain some, but not all descendants of a common ancestor. Coyne and Orr (2004: 472) conclude: “Because there is no unitary genetic history at the population level, it is almost impossible to recognize true paraphyly among closely related taxa using genetically based phylogenies.”
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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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272 8 Evolutionary Biology—Mayr
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284 9 Ernst Mayr—the Man his coll
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300 9 Ernst Mayr—the Man Fig.9.4.
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308 9 Ernst Mayr—the Man Fig.9.5b
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314 9 Ernst Mayr—the Man with a r
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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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