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Before discussing the reaction of the Entomological Society of France to Darwin’s theory of evolution through natural selection, it is necessary to provide a short summary of the position of French biologists in the period just after the

publication of The Origin of Species in 1859. Between 1859 and 1940, some of the most influential French biologists were Lamarckians, whose inclinations motivated them to adopt an experimental ap-proach to scientific investigations under the influence of the famous French physiologist, Claude Bernard (1813-1878). When addressing unexplained natural situations, however, they preferred to engage in discussions (which they considered to be a philosophical approach to investigation) rather than implement a rational, scientific approach.

Laurent Loison, a French specialist in the history of science, defines neo-Lamarckism “by the way scientists understood the nature of individual variation. If individual variation is understood as an effect, then the global conception in which the explanation takes place can be called Lamarckism…When individual variation is an effect, it is then necessary and legitimate to try to understand its causes (its ‘factors’), because by extension, the cause of the indi-vidual variation is the cause of the entire phyletic evolution” (Loison 2011). Loison effectively highlighted the significance of natural variability under the two main evolutionary theories (Loison 2010). He noted that neo-Lamarckism considers species variation as an effect (with environmental conditions as the main factor inducing variability) whereas neo-Darwinism considers species variation as a cause (without necessarily an attempt to search for the origin of variation), with natural selection driving species evolution by acting on this variability.

The most influential French biologists at the end of the 19th cen-

tury and in the first half of the 20th century held to neo-Lamarckism, and totally ignored the neo-Darwinist approach to evolutionary theory. Thomas H. Huxley (1825-1895), known as “Darwin’s bull-dog,” perfectly described this situation in France with the following words: “…to say nothing of the ill will of other powerful members of the Institute, produced for a long time the effect of a conspiracy of silence; and many years passed before the Academy redeemed itself from the reproach that the name of Darwin was not to be found on the list of its members” (Huxley 1888).

The Founding of the Entomological Society of FranceThe Entomological Society of France was founded on 31 January

1832; it is the oldest entomological society in the world. Thirty-five members gathered at its constituent assembly on 14 February of that year. Most of the society’s inaugural members were non-pro-fessionals, a fact that highlights the popularity of amateur entomol-ogy at that time. Among the honorary fellows were six professors of the Museum, including the well-known Baron Georges Cuvier (1769-1832) and Etienne Geoffroy Saint Hilaire (1772-1844). The assembly elected Pierre André Latreille (1762-1833) (Fig. 1) head of the Department of Natural History, Crustaceans, Arachnids, and Insects at the Museum, as Honorary President (Piguet, 2007). He had first been a clergyman and entomologist, and later a student, friend, and successor of Jean Baptiste Lamarck (1744-1829), who had died three years earlier. Were he living, would Lamarck have been chosen as honorary President?

Paul de Peyerimhoff, the President for year 1932, wrote of this question in the book published for the Society’s centennial celebration: “To be sure, Lamarck had been completely unknown

The Entomological Society of France with regard to Darwinism, after 1859

Yves Carton

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in his lifetime. Cuvier, like most officials, did not consider him…under these cir-cumstances, would the 35 fellows have thought of Lamarck? Yet, it is reasonable to argue that Latreille, when consulted, had recommended that his old supervi-sor be placed on the honor roll of the Corporation…” (Peyerimhoff 1932). However, a pamphlet written in 1847 about the professors of the Museum (Gosse 1847) said that Latreille “neither understood Lamarck nor Geffroy and was on the best possible terms with Cuvier!”

Was the Society able to open up to new ideas on the evolution of species? How did these entomologists react when The Origin of Species was published in 1859?

The Entomological Society of France’s Attitude towards Darwinism

It is very difficult to find evidence of any reaction, collectively or individually, to the ideas of Darwin among French en-tomologists. Well before 1859, Latreille, though connected to Lamarck, makes no reference to Lamarck or his ideas in his opening speech given on 29 February 1832. In his speech, Latreille said, “I don’t speak to you, gentlemen, of philosophical principles that should guide you in your studies; we know them, we adopt them generally, but these precepts as well as others are forgotten in practice. I would be afraid to report these abuses, to convert any kind of session, however friendly, in a controversial arena and one in which passions could take hold” (Latreille 1832). Did Latreille have the presentiment that Lamarck-ian theory would be controversial?

One can imagine that Darwinism should not have been expected to enter the consciousness of French entomologists until 1862, when the French version of The Origin of Species was first published. However, no references to this new conception of the evolution of species and its implications for taxonomy are found in speeches of the successive Presidents after 1862 or discussed in the Annals of the Society. While variability within species is one of the foundations of Darwin’s theory, President Auguste Chevrolat (1799-1884) addresses the problem in his speech of 12 February 1862: “It is deplorable to see in our Annals, admitted descriptions of supposed new species based on minor varieties, often on almost nothing” (Chevrolat 1862).

Even if he was right to castigate the species-naming frenzy, Chevrolat could have seized the opportunity to address how these varieties could be usefully studied. This point was not made until the presidency of Alfred Giard (1846-1908), a famous professor and head of the Department (Laboratory of Organized Beings) that had recently been created by the Paris town-hall on year 1888, and not by the University. In his presidential speech of 8 January 1896, President Giard proclaimed: “But by choosing the chair of the department of Evolution at the Sorbonne as flagship, you had another purpose: you wanted to highlight the considerable evidence that is returning to entomology in the discussion of major issues of General biology

that so strongly influence naturalists, and whose solution can only be expected by the convergent efforts of all workers. Who does not know...the interesting dis-coveries of A. R. Wallace, Bates or Trimen on natural selection, sexual selection, geography, zoology, etc.?” (Giard 1896).

However, the silence with respect to Darwinian theory continued: the subse-quent Presidents, all of whom had great responsibilities in University educa-tion, all had the opportunity to publicly embrace the new doctrine during their presidency: E.L. Bouvier (1856-1944) in 1898, P. Marchal (1862-1942) in 1906 and 1918, E. Rabaud (1868-1956) in 1915, R. G. Jeannel (1879-1965) in 1932, and P.-P. Grassé (1895-1985) in 1941. Each of them occupied the highest posi-tions in French biology and zoology, but none considered Darwinian evolution as the main point in biology.

The Election of Darwin as an Honorary Member of the Society

On 11 February 1874, a committee of five members formed with the goal of proposing honorary members for election to the Society, with two seats

reserved for foreign entomologists. After deliberation among the members that were available to vote at the meeting of the Entomo-logical Society on 25 February 1874 (as described in the Bulletin: “to the foreign members, the issue was more complex, because of the many famous names who have appeared during the discussion”1), Mr. Charles Darwin and Jørgen Matthias Christian Schiödte (1815-1884), Professor at the Museum of Copenhagen and entomologist, were both elected. The following letter was sent to Darwin by the Secretary, E. Desmarest2:

Société Entomologique de France. 3, Rue Linné, Paris, Le 27 Février 1874.

Dear Sir,The Entomological Society of France wishes to ally your judge-

ment with that of the men of science in whom it glories and who head its list: Westwood, Zetterstedt, Schiödte, Milne-Edwards, Boisduval, Perris, Guénée, etc. Accordingly, at last Wednesday’s meeting (25 January 1874), it elected you an honorary member. In according you the highest distinction it can bestow, it sought to recognise a learned zoologist, the author of highly appreciated work on the insects and Crustaceans.

The Society is counting on your acceptance; and, in accordance with the custom it has faithfully followed since its foundation in 1832, it seeks to publish your letter of acceptance in its Annals. Please send it to me as soon as possible, so that I can present it at our next meeting (11 March 1874).

The Society hardly dares hope that you might be as good as to participate in its work by means of an entomological memoir,

Fig. 1. Portrait of Pierre-André Latreille (1762-1833), the first President of the Entomological Society of France, in 1832 (reproduced with permission of Société entomologique de France).

11874, meeting on 25 february 1874, Bulletin de la Société entomologique de France, pp. xxiv – xxxv.

2Darwin Correspondence Project Database. http://www.darwinproject.ac.uk/entry-9320 (letter no. 9320, 27 février 1874)

210 American Entomologist  • Winter 2012

which it would be delighted to publish in its Annals. But it is counting on your devotion and hopes that you might be so good as to participate in its work by means of entomological memoirs, which it would be delighted to publish in its Annals. But it is counting on your devotion and hopes that you might be so good as to send some of your works for its library, as well as your photograph, which it would like to add to its albums.

Dear Sir and honoured colleague, please believe me, yours truly,

E Desmarest, Secretary since 1840.

P.S. Please let me know the name and address of the person in Paris to whom we should send the Society’s publica-tions, to which you are entitled from 1874 onwards.

We note that Desmarest only refer-ences “publications highly held in repute on Insects and Crustaceans,” with no al-lusion to Darwin’s crucial contribution to understanding the evolution of species, even 15 years after the publication of The Origin of Species!

Darwin responded on March 4, 18743 with these words:

Down Beckenham, Kent, 4 March 1874

Sir, I beg leave to thank you for your very obliging letter of the 27th

ult., in which you announce to me that the Entomological Society of France has conferred on me the distinguished honour of electing me, one of their Honorary members.

I hope that you will express to the Society of my part, my most sincere acknowledgements.

This honour is particularly gratifying to me, as during former years I collected insects with much ardour both in England and South America.

Although not a special student in Entomology, I have always felt the keenest interest in the subject; believing that it is admirably filled for throwing light on many general problems in Biology.

I have the pleasure, according to your request, of enclosing my photograph; and I have directed Williams and Norgate to send the French translations of my Origin of Species, and Descent of Man, to the Society, thinking that these would be the most acceptable

I have the honour to remain with high respect, Sir, your obliged and obedient servant,

Charles Darwin

In this letter, Darwin recognized the importance of entomological studies for understanding the major issues involved in the biologi-cal sciences. Evidence of the shipment of books that Darwin men-tions in his letter are still present in the archives and library of the Society: the photo of Darwin (Fig. 2), and French editons of two of his books, The Origin of Species and The Descent of Man, which had been autographed to the Society. It would seem that Darwin believed these two books to be his most important, since he does not offer On

the Various Contrivances by which British and Foreign Orchids are Fertilized by In-sects, though it had been translated into French by 1870. It is instructive to note that the library of the Society had, at that time, acquired all of Darwin’s work by purchasing seven more of his books that had been translated into French between 1878 and 1882.

The Members’ Concepts of Species Variability

Despite extensive research in the An-nals and the Bulletin, and in particular the minutes of monthly meetings of the Society, there was no reference to discussions, either positive or contra-dictory, of Darwin’s new approach to the evolution of species. This atmo-sphere cannot compare to the one that prevailed at the Royal Entomological Society of London, which hosted fre-quent, controversial discussions about Darwinism (Kritsky 2008).

However, a paper was published in the Annals of the Society that is par-ticularly instructive. In 1867, an English entomologist, Robert MacLachlan (1837-1904), submitted an article entitled “General Notes on Variations of Lepi-

doptera.” MacLachlan was a fellow of the Entomological Society of London; he joined it in 1858 and was President from 1885-1886. The submission of this work appears to have been a problem in France, because two members of the French Society were designated to lead the French translation of this work: Maurice Girard (1822-1886), a teacher and President of the Company, and Jules Fallou (1812-1895), the librarian of the Company since 1872. The article was published, but what is rather exceptional is that it was accompanied by more than 15 pages of comments from the two translators.

The article discusses the intra-species variability of butterflies according to English localities. Achille Guenée (1809-1880), a French lepidopterist, already considered England the “land of varieties”—as if variation were a feature specific to English wildlife! The comments of the translators were not favorable towards MacLachlan’s ideas: “What facts, will say, will be in favor of Darwin’s theory! But this author, like all naturalists, accepts all facts favorable to his ideas, often with little control, …he neglects or ignores those who are in opposition. Many species or races are without any connection to the colors surrounding them. For example, the writings of Standfuss [see below] suggest that ‘...it is not difficult to imagine that if this district was suddenly isolated, these forms would necessarily soon become what we call species’ or ‘... I would add that what goes for exotic Rhopalocera convinced me of the truth of the theory of devel-opment [or progressive development according to the author, this is in fact the evolution of species]; for it is where we found the greatest difficulty in distinguishing what should be considered as a species from what must be considered as a variety...There is no doubt we do well by designating species; yet we cannot question their common origin.’” The translators’ criticism became increasingly acrid: “A lot

Fig. 2. Charles Darwin (1809-1882) sent his own portrait to the French Entomological Society after his election as an honorary member at the meeting on 25 January 1874 (repro-duced with permission of Société entomologique de France).

31874. Bulletin de la Société entomologique de France, meeting on March 3, 1874, pp. lvi - lvii.

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of difficult discussions on these subjects of natural philosophy [be-come] logomachies...by venturing as little as possible into specula-tive theories.”

Both critics then develop an argument to show their opinion that these transformist ideas, apparently prom-ising new approaches, did not contradict a factual analysis: “Mr. Darwin’s theories are modifications, with sig-nificant and new ad-ditions, of the French doctrines of Etienne Geoffroy Saint Hilaire and Lamarck, especially on the progressive de-velopment of species; Lamarck was the one who moved furthest in this direction, to the point of hitting those who were well disposed towards his premises by the absurdity of his conclusions...In all, Lamarck remains in the right, and we find a very accurate application of his ideas in the way Mr. Grenier explains the varying development of eyes in certain types of cave beetles...We’ll see how easy it is to slip gradually from correct principles to deep errors when we are carried away by our imagina-tion in these sensitive issues, and that we do not bring extreme and careful control of the facts for every new idea that comes to mind...These are the ideas that led Lamarck’s doctrine of the transmutation of species (M. Darwin, Hooker) although you can see continually, and in entomology more than elsewhere, the filiated species that coex-ist under equal conditions in the environment, in food, in customs, and even in mating, without being able to switch from one species to another.”

One might think, from what has been transcribed, that some concepts of Darwinian theory found favor among some, but in reality this was not the case: “But what appears a seductive theory, I believe that it is impossible to admit with all the extension given to it by Mr. Darwin: his ideas about variability and transmutation of species were very serious failures, and in the form of objections that have not yet been answered in the most recent scientific discoveries. If we take groups of insects, highly diversified species, we find in their study one of the best pieces of evidence for the absence of transmutation of species.” We confirm here, with this critical analysis by the future president of the Society, the general thinking and consensus that certainly existed among the members; what we would today call a “unique thought” that would ultimately divide the Society.

Society Members’ Attitudes towards Soft InheritanceIn contrast with the particularly well-developed research that was

conducted on the adaptations of plants, especially in the laboratory of Professor Gaston Bonnier (1853-1922) in Fontainebleau near Paris,

French entomologists were reluctant to ex-perimentally research the heredity of acquired characters (soft inheri-tance). Insects were nevertheless suitable subjects for such re-search. Non-profes-sional entomologists, often involved in devel-oping insect breeding, could have participated in this research, but their pockets were not deep. They turned their attention to for-eign entomologists, especially Swiss ones, such as Max Standfuss (1854-1917), Director of the Zurich Museum, or Arnold Pictet (1869-1948), Professor at the

University of Geneva. In the research conducted by these two scientists,

we see a great deal of evidence of the neo-Lamarckian influence that prevailed at the time.

Standfuss’s experiments involved submitting butterfly pupae to extreme temperatures (+40 °C or -5 °C) : “I arrived at experimentally produced local forms...So I managed to use the cold procedure to directly transform Vanessa urticae L., from Zurich, into the polaris Stgr variety, from Lapland.” It is worth quoting Camille Jourdheuille (1830-1909), President of the Entomological Society of France in 1892, from his presentation of the results of Standfuss’s research at the 1895 annual Congress: “What was sought by Dr. Standfuss is the influence of temperature changes at the pupal instar, and he has succeeded beyond expectations...Different varieties of Vanessa io L. were represented on a plate, some of which show striking ressemblance to Vanessa urticae L.” Dr. Standfuss’s work was accepted without debate, two years later, in the Annals of the Entomological Society of France (Standfuss 1900). In the paper’s conclusion, Standfuss stated that “Only the more aberrant female [issued from breeding at high temperature]…transmitted more or less its characteristics, with an innovative aspect, to a small part of its offspring…”, the best proof that there was no actual inheritance of acquired characters.

Similarly, for over 20 years, Pictet investigated the influence of nutrition and temperature on larval stages, wing morphology, and biology (diapause) of adult moths. Initially, the Professor Pictet appears to observe that these changes are transmitted from one generation to another (Pictet 1911). However, after repeating these experiments over four generations (Pictet 1918-1921, I and II), he is obliged to note that “The first result of our research is that the character is acquired in this way, it is true, and begins to persist through generations, sometimes with increasing intensity, as long as the causal factor is maintained.” He concludes that “the findings of the present study again confirm that acquired characters are not

Fig. 3. Plasticity in wing-pattern development, illustrated by seasonal forms in two butterfly species: Araschnia levana (on the left) and Precis octavia (on the right). Nineteenth-century entomologists were intrigued by such variations, believing that climate conditions can definitively change the morphology of species (reproduced with permission of F. Nijhout, Duke University, NC).

212 American Entomologist  • Winter 2012

inherited...It is recognized, however, as unlikely that such action on the cyto-plasm can change the constitution of the chromosomes; i.e., become hereditary.”

These historical studies explored ideas that we now develop under the model of “phenotypic plasticity.” Certain-ly, French entomologists were particu-larly impressed by the seasonal variation of coloration in butterflies (Fig. 3). Un-fortunately, at that time, the question of how this variation was transmitted had rarely been addressed with suitable rigor. This question—the analysis of the source of reaction norms—is relevant for showing that it is the ability to change, not the change itself, that is genetically determined and subject to natural selec-tion. Thus, populations of insects living in different latitudes, and therefore exposed to different climatic conditions, have different reaction norms, each revealing itself as an adaptation to local conditions (David et al. 2004).

ConclusionWith this short review, we illustrate how members of the French

Entomological Society, after the publication of The Origin of Species in 1859, neither understood nor made an effort to understand Dar-winism. No attempt to deliberate over Darwin’s theory is evident in the presidential addresses that are published in the journals of the Society. One of the best-known French entomologists, Jean-Henri Fabre (1823-1915), even proclaimed himself to be an anti-Darwinian scientist!

One fact is particularly revealing. At the meeting of the Ento-mological Society on 26 April 1882, under the chairmanship of M. L. Reiche (1799-1890), the Secretary dedicated a brief portion of the assembly to the death of Darwin, with an honorary that stated “Mr. Kunckel Herculais [1843-1918, assistant at the Laboratory of Entomology of the Museum and tutor at the Agricultural Institute] is responsible for giving a notice to the Annals of the entomological works of Darwin.” Yet, a record or obituary never appeared in the An-nals or the Bulletin of the Entomological Society of France: Darwin’s death was not recognized in print by the Society.

French entomologists ignored these new aspects of biology, which could have influenced and advanced their work in insect taxonomy. They did not try to develop experimental approaches to insect breeding that would allow them to test the importance of species variability; such studies could have opened the door to understanding evolutionary processes. Very little data exists within the French entomological scientific press that would shed light on exactly how entomologists viewed species evolution. In fact, from their published papers during this period, it is very difficult today to determine whether French entomologists even adopted a clear and definite position regarding Lamarckism and Darwinism. Appar-ently, their majority (especially the members of the entomological Society in Paris) appear to not have been interested in how species evolve, although this was not the case for some entomologists in the country (Carton 2011).

Author’s Note. This article is partially adapted from a translation of Chapter 8 of Entomologie, Darwin et Darwinisme (Carton 2011) (Fig. 4).

Acknowledgements We wish to acknowledge Dr. F. Nijhout,

Department of Biology, Duke University, Durham, NC, for providing the butterfly pictures.

References CitedCarton, Y. 2011. Entomologie, Darwin et

Darwinisme. Hermann, Paris, 241 pp.Chevrolat, A. 1862. Discours du président,

Bulletin de la Société entomologique de France, séance du 12 février 1862, pp. vi-vii.

David, J., P. Gibert, and B. Moreteau. 2004. Evolution of reactions norms. In T.J. De-Witt and S.M. Scheiner (eds.), Phenotypic plasticity: functional and conceptual ap-proaches. Oxford Press, UK.

Giard, A. 1896. Discours du président, Bulle-tin de la Société entomologique de France, séance du 8 janvier 1896, pp. 2-7.

Gosse, Isid. S. de. 1847. Histoire naturelle drolatique et philosophique des profes-seurs du Jardin des Plantes, des aides

naturalistes, préparateurs, etc., ed. Sandré, Paris. Huxley, T.H. 1888. On the reception of the Origin of Species, pp. 173-204.

In Darwin, F., The life and letters of Charles Darwin, vol. II. John Mur-ray, London.

Jourdheuille, C. 1895. Sur les recherches récentes du Dr Standfuss, séance du 27 février 1895, Bul. Soc. Entomol. Fr., LXIV, pp. LXVII-LXX.

Kritsky, G. 2008. Entomological reactions to Darwin’s theory in the nine-teenth century. Ann. Rev. Entomol. 53: 345-360.

Latreille, P.A. 1832. Discours prononcé le 29 février 1832, à l’ouverture de la première séance de la Société entomologique, Ann. Soc. Entomol. France, t. I, pp. 22-34.

Loison, L. 2010. Qu’est-ce que le Néolamarckisme ? Ed. Vuibert, Paris, Paris, 248p.

Loison, L. 2011. French roots of french Neo-lamarckisms, 1879-1985. Journal of the History of Biology, 44, 713-744.

Mac-Lachlan, R. 1867. Notes générales sur les variations de Lépidoptères, traduit de l’anglais avec annotation par MM Maurice Girard et J. Fallou, Annales de la Société entomologiques de France, pp. 323-350.

Peyerimhoff, P. de. 1932. La Société Entomologique de France (1832-1931), Livre du Centenaire, pp. 1-86, ed. The French Entomological Society.

Pictet, A. 1911. Un nouvel exemple de l’hérédité des caractères acquis, Arch. Sci. Phys. Nat., 31: 561-563.

Pictet, A. 1918-1921. I. Observations biologiques sur Porthesia similis Fuessl. pendant quatre générations, Bull. Soc. Lépidopt. Genève, IV, pp. 186-201; II. Expériences de génétique avec Porthesia similis et d’autres lépidoptères, Bull. Soc. Lépidopt. Genève, IV, pp. 202-220.

Piguet, H. 2007.1a vie et l’œuvre de Pierre-André Latreille, «prince de l’entomologie» (1762-1833), Société Entomologique. France, 21 p.

Standfuss, M. 1900. Etudes zoologiques expérimentales sur les lépidop-tères. Résultats principaux obtneus jusqu’à la fin de 1898. Rédaction française par H.-F. Deckert, Ann. Soc. Entomol. Fr. 69: 82-101.

Dr. Yves Carton is Vice President of the Entomological Society of France and Directeur de Recherches Emérite of the Laboratoire Evolution, Génomes et Spéciation, Equipe IRD-CNRS-INRA, CNRS 91198 Gif/Yvette and Université Paris 11 Orsay. E-mail: yves.carton@legs.cnrs-gif.fr

Fig. 4. Cover of the book Entomologie, Darwin and Darwinisme.