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3 The New York Years (1931–1953)

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202 5 Biological Species and Speciation—Mayr’s First Synthesis<br />

for the Chermock circle Mayr’s Systematics and the Origin of Species, following<br />

upon Dobzhansky’s book, was the hunter’s vade mecum. From Mayr we learned<br />

how to define species as biological units. With the help of his written word we<br />

pondered the exceptions to be expected and the processes by which races evolved<br />

into species. We acquired a clearer, more logical way to think about classification<br />

by using the phylogenetic method” (Wilson 1994: 110–112; see also Wilson<br />

1998: 4).<br />

Working on a revision of his successful 1942 book, Mayr soon decided to separate<br />

methodological aspects of systematic zoology from the science of species and<br />

evolutionary biology. In this way originated his textbook, Methods and Principles of<br />

Systematic Zoology (1953a, with Linsley and Usinger) which covered methodological<br />

aspects (p. 321) and his masterly monograph on Animal Species and Evolution<br />

(1963b) treating the species problem and aspects of evolutionary biology.<br />

In his article, “Trends in avian systematics” Mayr (1959e) emphasized two<br />

main topics: (1) Population systematics and (2) macrotaxonomy. Subspecies are<br />

unsuitable for describing the population structure of a species. This can be done in<br />

terms of three major population phenomena: (a) Geographical isolates which are<br />

particularly common near the periphery of the species range and which represent<br />

important units of evolution; (b) <strong>The</strong> population continuum, a continuous series<br />

of populations which make up the main body of the species, and (3) Zones of<br />

secondary intergradation between a former geographical isolate (which did not<br />

yet attain full species rank) and the main body of the species. To make such an<br />

analysis of all species of a family, and to record the relative frequency of the<br />

three elements, is one of the steps leading to “comparative systematics.” Recording<br />

the position of belts of secondary hybridization permits the reconstruction of<br />

formerly existing barriers and of the location of “refuges” caused by drought<br />

conditions in low latitudes during the Pleistocene. With respect to macrotaxonomy,<br />

Mayr proposed to utilize various new sources of information like promising new<br />

character complexes of behavior and biochemistry to determine the phylogenetic<br />

relationships between genera, families and orders.<br />

Behavioral characters are completely equivalent to morphological characters<br />

(Mayr 1958g). If there is a conflict between the evidence provided by morphological<br />

characters and that of behavior, taxonomists are increasingly inclined to<br />

give greater weight to the ethological evidence, e.g., in the case of grasshoppers,<br />

in certain swallows (p. 327), ducks (p. 327), and finches (p. 329). Based on behavioral<br />

characteristics the weaver finches (Estrildidae) and weaverbirds (Ploceidae)<br />

are not as closely related as previously thought. Movements often precede special<br />

morphological features that make them particularly conspicuous. Comparative<br />

studies of behavior permit statements about trends in the evolution of behavior,<br />

as shown, e.g., by the work of Whitman on pigeons, Tinbergen on gulls, Heinroth<br />

and Lorenz on ducks, Meyerriecks on herons, Hinde on finches, Morris on<br />

weaver finches. Other groups studied in this way include grasshoppers, Drosophila,<br />

hymenopterans, and cichlid fishes.<br />

In 1964(n), Mayr again summarized the then “new systematics” mentioning that<br />

certain aspects can be traced back to some authors of the mid-19th century who

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