Geoffrey Douglas Hale Carpenter
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G.D. Hale Carpenter MBE (26 October 1882 in
Eton, Berkshire Eton ( ) is a town in Berkshire, England, on the opposite bank of the River Thames to Windsor, Berkshire, Windsor, connected to it by Windsor Bridge. The civil parish, which also includes the village of Eton Wick two miles west of the town, had ...
– 30 January 1953 in
Oxford Oxford () is a City status in the United Kingdom, cathedral city and non-metropolitan district in Oxfordshire, England, of which it is the county town. The city is home to the University of Oxford, the List of oldest universities in continuou ...
) was a British
entomologist Entomology (from Ancient Greek ἔντομον (''éntomon''), meaning "insect", and -logy from λόγος (''lógos''), meaning "study") is the branch of zoology that focuses on insects. Those who study entomology are known as entomologists. In ...
and
medical doctor A physician, medical practitioner (British English), medical doctor, or simply doctor is a health professional who practices medicine, which is concerned with promoting, maintaining or restoring health through the study, diagnosis, prognosis ...
. He worked first at the
London School of Hygiene and Tropical Medicine The London School of Hygiene & Tropical Medicine (LSHTM) is a public university, public research university in Bloomsbury, central London, and a constituent college, member institution of the University of London that specialises in public hea ...
, and in
Uganda Uganda, officially the Republic of Uganda, is a landlocked country in East Africa. It is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the ...
, on tse-tse flies and
sleeping sickness African trypanosomiasis is an insect-borne parasitic infection of humans and other animals. Human African trypanosomiasis (HAT), also known as African sleeping sickness or simply sleeping sickness, is caused by the species '' Trypanosoma b ...
. His main work in
zoology Zoology ( , ) is the scientific study of animals. Its studies include the anatomy, structure, embryology, Biological classification, classification, Ethology, habits, and distribution of all animals, both living and extinction, extinct, and ...
was on
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...
in
butterflies Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...
, an interest he developed in Uganda and Tanganyika. He succeeded E.B. Poulton as Hope Professor of Zoology at
Oxford University The University of Oxford is a collegiate research university in Oxford, England. There is evidence of teaching as early as 1096, making it the oldest university in the English-speaking world and the second-oldest continuously operating u ...
from 1933 to 1948.


Biography

Douglas was a son of Philip Herbert Carpenter DSc FRS, a schoolmaster at
Eton College Eton College ( ) is a Public school (United Kingdom), public school providing boarding school, boarding education for boys aged 13–18, in the small town of Eton, Berkshire, Eton, in Berkshire, in the United Kingdom. It has educated Prime Mini ...
; a grandson of the
naturalist Natural history is a domain of inquiry involving organisms, including animals, fungi, and plants, in their natural environment, leaning more towards observational than experimental methods of study. A person who studies natural history is cal ...
and
physiologist Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a subdiscipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out chemical and ...
William Benjamin Carpenter William Benjamin Carpenter CB FRS (29 October 1813 – 19 November 1885) was an English physician, invertebrate zoologist, and physiologist. He was instrumental in the early stages of the unified University of London. Life Carpenter was bor ...
; and a great-grandson of Lant Carpenter, a Unitarian minister. Carpenter attended
St Catherine's College, Oxford St Catherine's College (colloquially called St Catz or Catz) is one of the Colleges of the University of Oxford, constituent colleges of the University of Oxford. In 1974, it was also one of the first men's colleges to admit women. It has 528 un ...
, graduating in 1904. He studied medicine at St George's Hospital, London, graduating as a
Bachelor of Medicine, Bachelor of Surgery A Bachelor of Medicine, Bachelor of Surgery (; MBBS, also abbreviated as BM BS, MB ChB, MB BCh, or MB BChir) is a medical degree granted by medical schools or universities in countries that adhere to the United Kingdom's higher education trad ...
(the standard combined medical degree at the University of London at that time) in 1908. He then joined the London School of Hygiene and Tropical Medicine, and became a
Doctor of Medicine A Doctor of Medicine (abbreviated MD, from the Latin language, Latin ) is a medical degree, the meaning of which varies between different jurisdictions. In the United States, and some other countries, the ''MD'' denotes a professional degree of ph ...
in 1913 with a dissertation on the tsetse fly ('' Glossina palpalis'') and sleeping sickness. In 1919 he married Amy Frances Thomas-Peter from
Cornwall Cornwall (; or ) is a Ceremonial counties of England, ceremonial county in South West England. It is also one of the Celtic nations and the homeland of the Cornish people. The county is bordered by the Atlantic Ocean to the north and west, ...
. The marriage had no issue.


Career

In 1910 he joined the Colonial Medical Service, where he worked in
Uganda Uganda, officially the Republic of Uganda, is a landlocked country in East Africa. It is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the ...
on the north shore of
Lake Victoria Lake Victoria is one of the African Great Lakes. With a surface area of approximately , Lake Victoria is Africa's largest lake by area, the world's largest tropics, tropical lake, and the world's second-largest fresh water lake by surface are ...
. Upon the outbreak of World War I, Carpenter was called to service in the
British Army The British Army is the principal Army, land warfare force of the United Kingdom. the British Army comprises 73,847 regular full-time personnel, 4,127 Brigade of Gurkhas, Gurkhas, 25,742 Army Reserve (United Kingdom), volunteer reserve perso ...
Medical Corps. He was stationed with the troops at the border between Uganda and
German East Africa German East Africa (GEA; ) was a German colonial empire, German colony in the African Great Lakes region, which included present-day Burundi, Rwanda, the Tanzania mainland, and the Kionga Triangle, a small region later incorporated into Portugu ...
. In December 1914 he was appointed Medical Officer at the fort in Kakindu, southern Uganda. As it turned out, he had plenty of spare time, and spent it studying the local butterflies. "The hosts of butterflies at Kakindu passed beyond anything I had ever see; some days are quite unforgettable". From May 1916 to January 1918, he worked in Tanganyika (former German E. Africa), 200 miles south to south-east of Lake Victoria. Here he conducted experiments on
palatability Palatability (or palatableness) is the hedonic reward (which is pleasure of taste in this case) provided by foods or drinks that are agreeable to the "palate", which often varies relative to the homeostatic satisfaction of nutritional and/or wa ...
with young insectivorous monkeys. He tested the edibility of cryptic and
aposematic Aposematism is the Advertising in biology, advertising by an animal, whether terrestrial or marine, to potential predation, predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the pr ...
insects. This was propitious, because later, back at Oxford, both he and E.B. Poulton worked on the role of predators in shaping
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...
. The standard theory was that cryptic forms were palatable, and aposematic forms were distasteful, implying that palatable mimics of distasteful forms could gain protection from
predation Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
. The question at stake was whether the observations, which dated from work by naturalists in the 19th century, could be accounted for by
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
.


Mimicry

In England the geneticist R.C. Punnett, and in America the ornithologist W.L. McAtee, doubted both that birds could distinguish distasteful forms, and that their predation was heavy enough to bring about the colour forms found in butterflies. Punnett's 1915 ''Mimicry in butterflies'' rejected selection as the main cause of mimicry. He noted: :1. The absence of transitional forms and the frequent lack of mimicry in male butterflies were unexplained by selectionist theory. :2. The enigma of polymorphic mimicry. Some species of butterfly mimicked not merely one, but several models. In breeding experiments these polymorphs cleanly segregated according to Mendel’s law of segregation. :3. Evidence of birds as selective agents was slight and little was known of birds' discriminatory powers, and :4. The gradual accumulation of minute variations did not fit with the facts of heredity. For Punnett, none of these observations were explained by gradual selectionism. Instead he thought mimicry had arisen from sudden mutational jumps (saltations). Once a mimic was formed by mutation, natural selection might play a conservative role. However, one by one, each of these objections were shown to be without substance. Evidence from field observations and experiments showed that birds were often the agents of selection in insects.Carpenter, G.D.H.; Ford, E.B. (1933). ''Mimicry''. London: Methuen. Evidence that small-scale mutations were common arrived as soon as breeding experiments were designed to detect them: it was a consequence of experimental methods that early mutations were so noteworthy. Explanations for polymorphism were advanced by E.B. Ford and Dobzhansky and colleagues, who developed experimental methods for populations in the wild.Ford, E.B. (1975). ''Ecological Genetics, Fourth edition''. London: Chapman and Hall. The question of polymorphism is discussed further in
polymorphism (biology) In biology, polymorphism is the occurrence of two or more clearly different morphs or forms, also referred to as alternative '' phenotypes'', in the population of a species. To be classified as such, morphs must occupy the same habitat at the ...
. The gradual coming-together of field observations and experimental genetics is part of the evolutionary synthesis which took place in the middle of the twentieth century. The small 1933 book on mimicry by Carpenter and E.B. Ford was the first book on
ecological genetics Ecological genetics is the study of genetics in natural populations. It combines ecology, evolution, and genetics to understand the processes behind adaptation. It is virtually synonymous with the field of molecular ecology. This contrasts wit ...
, a field which produced a series of classic studies uniting fieldwork with laboratory genetics. The book is a minor masterpiece of evolutionary biology.


Legacy

Carpenter is commemorated in the scientific names of two species of African reptiles: '' Chilorhinophis carpenteri'' and '' Kinyongia carpenteri''. Beolens, Bo; Watkins, Michael; Grayson, Michael (2011). ''The Eponym Dictionary of Reptiles''. Baltimore: Johns Hopkins University Press. xiii + 296 pp. . ("Carpenter, G.D.H.", p. 48).


Obituaries

* Hobby, B.M. (1953). "Geoffrey Douglas Hale Carpenter". ''British Medical Journal'' 1: 406. * Remington, Charles L. (1954). "Geoffrey Douglas Hale Carpenter". ''Lepidopterist's News'' 8: 31–43.


References


Publications

* (1920) ''A Naturalist on Lake Victoria, with an Account of Sleeping Sickness and the Tse–tse Fly''. London: Unwin. * (1921) "Experiments on the relative edibility of insects, with special reference to their coloration". ''Transactions of the Royal Entomological Society, London'' 54: 1–105. * with E.B. Ford (1933). ''Mimicry''. London: Methuen. * (1933) "Gregarious roosting habits of aposematic butterflies". ''Proceedings of the Entomological Society of London'' 8: 110–111. * (1935) "The Rhopalocera of Abyssinia a faunistic study". ''Trans. R. Entomol. Soc. London'' 83 : 313–447. * (1936) "Charles Darwin and entomology". ''Transactions of the South–eastern Union of Scientific Societies''. Papers Contributed to Congress 1936: 1–23. * (1938) "Audible emission of defensive froth by insects with an appendix on the anatomical structures concerned in a moth by H. Eltringham". ''Proceedings of the Zoological Society of London'' 108: 243–252. * (1939) "Birds as enemies of butterflies, with special reference to mimicry". ''Proceedings, VII Internationaler Kongress für Entomologie'', Berlin 1938: 1061–1074. * (1941) "The relative frequency of beakmarks on butterflies of different edibility to birds". ''Proc. Zool. Soc. London (Series A)'' 3: 223–231. * (1947) "The writings of I. Portschinsky on warning colours and eyespots". ''Proc. Entomol. Soc. London. Series A. General Entomology'' 22: 103–113. * (1949) "''Pseudacraea eurytus'' (L.) (Lep. Nymphalidae): a study of a polymorphic mimic in various degrees of speciation". ''Trans. R. Entomol. Soc. London'' 100: 71–133. * (1953) "The genus ''Euploea'' (Lep. Danaidae) in Microneia, Melanesia, Polynesia and Australia. A zoo–geographical study". ''Trans. Zool. Soc. London'' 28: 1–184, plates 1–9.


External links


Carpenter, G.D. Hale (1920). ''A Naturalist on Lake Victoria, with an Account of Sleeping Sickness and the Tse-tse Fly''. London: T.F. Unwin Ltd. Biodiversity Archive
{{DEFAULTSORT:Carpenter, G.D.H. Hope Professors of Zoology Fellows of Jesus College, Oxford 1882 births 1953 deaths British Army personnel of World War I British entomologists British evolutionary biologists 20th-century British zoologists Members_of_the_Order_of_the_British_Empire