Editing David Lack (section)

==Career and research==
After Cambridge, Lack, on the recommendation of [[Julian Huxley]] took up a position as a science mentor at [[Dartington Hall]] School,<ref>Anderson (2013):27-28.</ref> [[Devon]]shire from 1934 until Summer 1938 when he took a year off to study bird behaviour on the [[Galapagos Islands]]. In 1935 he made his first trip to the United States as a chaperone for a Dartington Hall student returning to California. Here he met [[Joseph Grinnell]] and [[Robert McCabe]] and gave a talk at the [[Cooper Ornithological Society|Cooper Ornithological Club]]. In New York, he met [[Ernst Mayr]] at the American Museum of Natural History. He returned via the ''[[SS Bremen (1928)|SS Bremen]]'', only one of about four English speakers on the German ship.<ref>Anderson (2013):43-45.</ref> He was only in the Galapagos for part of that year, starting August 1938. With the data that he collected in the Galapagos, especially on the finches he went to the United States. April to August 1939 was spent at the [[California Academy of Sciences]] which held a large collection of the finches of Galapagos that had been studied earlier by [[Harry Swarth]] and at [[Ernst Mayr]]'s home in [[New Jersey]]. While in the US he made a study of the tricoloured blackbird with John T. Emlen.<ref>Anderson (2013):58-59.</ref>  He returned home in September 1939, after the outbreak of war.<ref>Provine, William B (1986) ''Sewall Wright and evolutionary biology''. Chicago. {{ISBN|0226684733}} p. 406</ref> Lack published ''The Galapagos Finches (Geospizinae), A Study in Variation'' in which he examined variations within species across islands and considered that many of them were non-adaptive and due to founder effect and genetic drift.<ref>Anderson (2013):59-60.</ref> Lack's first major work was ''The Life of the Robin'', which was based on four years of field work that he conducted while teaching at [[Dartington Hall|Dartington Hall School]]. He examined robin behaviour, song, territory, pairing and breeding using ringing to mark and track individual birds.<ref>Anderson (2013):25-27.</ref> The manuscript was completed in 1942 and it went through five editions from 1943 to 1970. One of Lack's students at Dartington Hall was [[Eva Ibbotson]]. A colleague who helped in filming some of the robins for Lack was the geography teacher Bill Hunter. In 1934 Lack went to Tanganyika on an invitation from [[Reginald Ernest Moreau|R.E. Moreau]].<ref>Anderson (2013):34-35.</ref>
[[File:Apus apus 01.jpg|thumb|upright|The Common Swift, one of many subjects studied by Lack.]]

Lack was committed to pacificism and debated the philosophy even during his Dartington days with the founder of the college, [[Leonard Knight Elmhirst]]. During [[World War II]] Lack however served with a [[British Army]] unit called the Army Operational Research Group on the Orkney Islands working on [[radar]] use. During this work he met other biologists who had been inducted into the war including [[George C. Varley|George Varley]], an entomologist who introduced him to the idea of density-dependent regulation of animal populations.<ref>Anderson (2013):47.</ref> Lack's observations on spurious echoes produced by birds would later allow him to establish the field of [[radar ornithology]] to study [[bird migration]].<ref>Anderson (2013):48-49.</ref><ref>{{cite journal|author1=Fox, A.D.|author2=Beasley, Patrick D.L.|year=2010|title=David Lack and the birth of radar ornithology|journal=Archives of Natural History|volume=37|issue=2|pages=325–332|doi=10.3366/anh.2010.0013}}</ref> Lack was released from wartime duty in August 1945 so as to take a position to as Director (succeeding [[Wilfred Backhouse Alexander|W.B. Alexander]]) of the [[Edward Grey Institute of Field Ornithology]] at [[Oxford University]], a position that he held until his death in 1973.

Lack's work in ornithology was almost entirely based on studies of the ''living bird''. He was one of the pioneers of life-history studies in Britain, especially those based on quantitative approaches, when some traditional ornithologists of the time were focussing their studies on morphology and geographic distribution.<ref>{{Cite journal | last1 = Johnson | first1 = K. | title = The Ibis: Transformations in a Twentieth Century British Natural History Journal | doi = 10.1007/s10739-004-1499-3 | journal = Journal of the History of Biology | volume = 37 | issue = 3 | pages = 515–555 | year = 2004 | s2cid = 83849594 }}</ref> Lack's major scientific research included work on [[population biology]] and density dependent regulation. His work suggested that natural selection favoured [[avian clutch size|clutch size]]s that ensured the greatest number of surviving young. This interpretation was however debated by [[V.C. Wynne-Edwards]] who suggested that clutch size was density-independent. This was one of the earliest debates on [[group selection]]. Lack's studies were based on [[nidicolous]] birds and some recent studies have suggested that his findings may not hold for other groups such as seabirds.<ref>{{cite journal|author1=Ydenberg, R.C.  |author2=Bertram, D.F. |year=1989|title= Lack's clutch size hypothesis and brood enlargement studies on colonial seabirds|journal=Colonial Waterbirds|volume=12|issue=1|pages= 134–137|doi=10.2307/1521328|jstor=1521328 }}</ref>

As a mentor for numerous doctoral students, Lack followed a hands-off method, letting students decide their own research topics. He encouraged students to sort out their ideas and find the "simplest explanation, which was probably best." He would make students work on their papers and give only one reading to their thesis asking them to choose either a draft or a final version to submit.<ref>Anderson (2013):159-160.</ref>

He wrote numerous papers in ornithological journals, and had a knack of choosing memorable titles: he once claimed to have single-handedly caused the renaming of a group of birds through the submission of a scientific paper with the title "Territory and Polygamy in a Bishop". This 1935 publication was subsequently titled "Territory and polygamy in a bishop bird, ''[[Euplectes hordeaceus|Euplectes hordeacea]] hordeacea'' (Linn.)" in the journal ''Ibis'' as the journal editor felt that the title might cause misunderstanding.<ref>Anderson (2013):39.</ref>

=== Darwin's finches ===
Lack's most famous work is ''Darwin's Finches,'' a landmark study whose title linked [[Charles Darwin|Darwin]]'s name with the [[Galapagos]] group of species and popularised the term "[[Darwin's finches]]" in 1947, though the term had been introduced by [[Percy Lowe]] in 1936.<ref>{{Cite journal | last1 = Steinheimer | first1 = F. D. | title = Charles Darwin's bird collection and ornithological knowledge during the voyage of H.M.S. Beagle, 1831–1836 | doi = 10.1007/s10336-004-0043-8 | journal = Journal of Ornithology | volume = 145 | issue = 4 | pages = 300–320 | year = 2004 | s2cid = 24957761 }}</ref> There are two versions of this work, differing significantly in their conclusions. The first is a book-length monograph, written after his visit to the Galapagos, but not published until 1945.<ref name=geospizinae>{{cite journal|title=The Galapagos Finches (Geospizinae) A Study in Variation|first=David|last= Lack|journal=The Auk|volume=62|issue=4|year=1945|pages=644–645|issn=0004-8038|doi=10.2307/4079846|jstor=4079846|doi-access=free}}</ref> In it Lack interprets the differences in bill size as species recognition signals, that is, as [[isolating mechanisms]].

The second is the later book in which the differences in bill size are interpreted as adaptations to specific food niches, an interpretation that has since been abundantly confirmed.<ref name=darwinsfinches>Lack, David 1947. ''Darwin's Finches''. [[Cambridge University Press]] reissued in 1961 by Harper, New York, with a new preface by Lack; reissued in 1983 by Cambridge University Press with an introduction and notes by Laurene M. Ratcliffe and Peter T. Boag). {{ISBN|0-521-25243-1}}</ref> This change of mind, according to Lack's Preface, came about as a result of his reflections on his own data whilst he was doing war work. The effect of this change in interpretation is to put the emphasis for speciation onto natural selection for appropriate food handling instead of seeing it primarily as a by-product of an isolating mechanism. In this way his work contributed to the [[modern synthesis (20th century)|modern evolutionary synthesis]], in which [[natural selection]] came to be seen as the prime mover in evolution, and not random or mutational events. Lack's work laid the foundations for the much more extensive work of [[Peter and Rosemary Grant]] and their colleagues.<ref>Grant, Peter R. 1999. ''Ecology and evolution of Darwin's finches''. Princeton NJ.</ref> Lack's work feeds into studies of island biogeography which continue the same range of issues presented by the Galapagos fauna on a more varied canvas.<ref>MacArthur R. and Wilson E.O. 1967. ''The theory of island biogeography''. Princeton 1967.</ref> According to [[Ernst Mayr]],

:"The person who more than anyone else deserves credit for reviving an interest in the ecological significance of species was David Lack... It is now quite clear that the process of speciation is not completed by the acquisition of isolating mechanisms but requires also the acquisition of adaptations that permit co-existence with potential competitors."<ref name=Mayr274>Mayr, Ernst (1985). [https://books.google.com/books?id=pHThtE2R0UQC ''The growth of biological thought: diversity, evolution, and inheritance.''] Harvard University Press. {{ISBN|0-674-36446-5}}, pp. 274–5.</ref>

===Lack's Principle===
In 1943 Lack took an interest in clutch size after reading Moreau's manuscript sent to the ''Ibis''. Lack was then an assistant to the editor of the ''Ibis''.<ref>Anderson (2013):62.</ref> Lack postulated what is now known as Lack's Principle, which states that "the clutch size of each species of bird has been adapted by natural selection to correspond with the largest number of young for which the parents can, on average, provide enough food".<ref>Lack D (1954) The regulation of animal numbers. Clarendon Press, Oxford</ref><ref>{{cite journal |last1=Pasztor |first1=E |last2=Loeschcke |first2=V |title=The Coherence of Cole's Result and Williams' Refinement of Lack's Principle |journal=Oikos |date=November 1989 |volume=56 |issue=3 |pages=416–420 |doi=10.2307/3565627 |jstor=3565627 |bibcode=1989Oikos..56..416P |url=https://www.jstor.org/stable/3565627}}</ref>
<ref>{{cite journal |last1=Williams |first1=George C. |title=Natural Selection, the Costs of Reproduction, and a Refinement of Lack's Principle |journal=The American Naturalist |date=December 1966 |volume=100 |issue=916 |pages=687–690 |doi=10.1086/282461 |jstor=2459305 |bibcode=1966ANat..100..687W |url=https://www.jstor.org/stable/2459305}}</ref>

=== Population regulation ===
Lack took a keen interest in the mechanisms involved in regulating populations in nature. ''The Natural Regulation of Animal Numbers'' is one of Lack's most frequently cited works. Here he gave primacy to natural selection in determining the rate of reproduction and he especially countered the idea that it was adjusted with mortality rates so that constant populations are maintained. It was critiqued by [[J. B. S. Haldane|J.B.S. Haldane]] who found it lacking mathematical precision and biased to bird studies. The other major critic was [[V. C. Wynne-Edwards|V.C. Wynne-Edwards]] with whom he clashed for nearly a decade.<ref>Anderson (2013):78-81.</ref> Lack followed up on the criticisms in his later books including Population Studies of Birds (1966).

===Published books===
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*Lack, David. 1943. ''The Life of the Robin''. Witherby, London. {{ISBN|978-1843681304}} (4th edition, 1965, illustrated by [[Robert Gillmor]])
*Lack, David. 1947. ''Darwin's Finches''.<ref name=darwinsfinches/>
*Lack, David. 1950. ''Robin Redbreast''. Oxford. (A new edition of this book, revised and expanded by Lack's son [[Andrew Lack (author)|Andrew]], was published under the title ''Redbreast: the Robin in life and literature'' by SMH Books in 2008.) {{ISBN|9780955382727}}
*Lack, David. 1954. ''The Natural Regulation of Animal Numbers''. Oxford University Press, Oxford. {{ISBN|978-1299428287}}
*Lack, David. 1956. ''Swifts in a Tower.'' Methuen, London. {{ISBN|0412121700}}
** 2018 Updated edition, illustrated by Colin Wilkinson. Unicorn. {{ISBN|978-1911604365}}
*Lack, David. 1957. ''Evolutionary Theory and Christian Belief: The Unresolved Conflict.'' Methuen, London. {{ISBN|978-0415474900}}
*Lack, David. 1965. ''Enjoying Ornithology''. Methuen, London. {{OCLC|691693809}} (illustrated by Robert Gillmor)
*Lack, David. 1966. ''Population Studies of Birds''. Oxford University Press, Oxford. {{ISBN |9780198573418}} (illustrated by Robert Gillmor)
*Lack, David. 1968. ''Ecological Adaptations for Breeding in Birds''. Methuen, London. {{ISBN|978-0412112201}} (illustrated by Robert Gillmor)
*Lack, David. 1971. ''Ecological Isolation in Birds''. Harvard University Press, Cambridge, Mass. and Blackwell, Oxford. {{ISBN |978-0674224421}} (illustrated by Robert Gillmor)
*Lack, David. 1974. ''Evolution Illustrated by Waterfowl''. Harper & Row, London.{{ISBN|978-0061361692}}
*Lack, David. 1976. ''Island Biology Illustrated by the Land Birds of Jamaica''. University of California Press, Berkeley. {{ISBN|0-520-03007-9}} (posthumously).
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===Published journal articles===
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*{{Cite journal | last1 = Lack | first1 = D. | title = Evolution of the Galapagos Finches | doi = 10.1038/146324a0 | journal = Nature | volume = 146 | issue = 3697 | pages = 324–327 | year = 1940 | bibcode = 1940Natur.146..324L | s2cid = 43465549 }}
*{{cite journal | last1 = Lack | first1 = David | year = 1942 | title = Ecological features of the bird faunas of British small islands | journal = Journal of Animal Ecology | volume = 11 | issue = 1| pages = 9–36 | doi = 10.2307/1298 | jstor = 1298 | bibcode = 1942JAnEc..11....9L }}
*Lack, David. 1945. The Galapagos finches (Geospizinae): a study in variation.<ref name=geospizinae/>
*{{cite journal | doi = 10.1111/j.1474-919x.1947.tb04155.x | volume=89 | title=The Significance of Clutch-size | year=1947 | journal=Ibis | pages=302–352 | last1 = Lack | first1 = David| issue=2 }}; '''90''', 25–45.  
*Lack, David 1949. The significance of reproductive isolation. In Jepsen G, Mayr E and Simpson GG (eds) ''Genetics, palaeontology and evolution''. Princeton.
*Lack, David. 1954. The evolution of reproductive rates. In Huxley J, Hardy AC and Ford EB (eds). ''Evolution as a process''. [[Allen & Unwin]], London. {{ISBN missing}}
*{{cite journal | last1 = Lack | first1 = David | year = 1967 | title = Interrelationship in breeding adaptations as shown by marine birds | journal = Proc. XIVth Int. Orn. Congr. Oxford | volume = 1966 | pages = 3–42 }}
*{{cite journal | last1 = Lack | first1 = David | year = 1973 | title = The numbers of species of hummingbirds in the West Indies | journal = [[Evolution (journal)|Evolution]] | volume = 27 | issue = 2| pages = 326–337 | doi = 10.2307/2406972 | jstor = 2406972 | pmid = 28564781 }}
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