History of Ecological Sciences, Part 45: Ecological Aspects of Entomology During the 1800s

2013; Ecological Society of America; Volume: 94; Issue: 1 Linguagem: Inglês

10.1890/0012-9623-94.1.36

ISSN

2327-6096

Autores

Frank N. Egerton,

Tópico(s)

Biological Control of Invasive Species

Resumo

Click here for all previous articles in the History of the Ecological Sciences series by F. N. Egerton An extensive literature on the history of entomology (see Literature Guide below) provides the basis for a discussion of aspects of ecologically relevant entomology for the 1800s, with two topics deferred to later parts of this history: diseases of insects and insects as vectors of human disease, to part 46 on parasitism and the germ theory of disease; and pollination ecology and domestic bees, to part 52 on symbiosis. We saw in parts 21 and 30 (Egerton 2006, 2008) that there were many important studies on insects during the 1700s, and in parts 33 and 41 (Egerton 2009b, 2012a) we met notable entomologists of the 1800s: Thomas Say (1787–1834) and Henry Walter Bates (1825–1892). Alfred Russel Wallace (1823–1913), discussed in parts 41–42 (Egerton 2012a, b), also notably contributed to entomology. Charles Darwin (1809–1882) had collected beetles at Cambridge and insects on the voyage of the Beagle; he discussed insects in the Origin of Species, and he studied insects in relation to pollination and insectivorous plants (Riley 1882, Egerton 2010, 2011b, Carton 2011). Some of these men might themselves be classified as "stamp collectors," but taxonomy and systematics were especially important for entomology if it were ever to become a sophisticated science, because it was essential for zoologists to distinguish whatever species they studied so others could verify or refute claims made about particular species. One historian (Clark 2006:28) calls the Introduction to Entomology (four volumes, 1815–1826, edition 7, 1856), by William Kirby (1759–1850) and William Spence (1783–1859), a founding text for entomology in English. Unlike the works in French and German, there had not been any work so comprehensive published in English before. In his "Preface" to the first edition, Spence explained: "One principal cause of the little attention paid to Entomology in this country," was that there had been "ridicule so often thrown upon the science." Botanists were "sheltered now by the sanction of fashion," but "in the minds of most men, the learned as well as the vulgar, the idea of the trifling nature of this pursuit is so strongly associated with that of the diminutive size of its objects, that an Entomologist is synonymous with every thing futile and childish." If that had been a problem, Kirby and Spence's treatise improved the entomologists' status. However, both Clark and Spence gave a misleading impression of English interest in entomology before 1817, assuming books are a measure of interest. The Aurelian (1766) and other works written by Moses Harris (1730–about 1788; Salmon 2000:115–117, Mays 2004) on butterflies were popular enough to go through several editions (Freeman1980:157); Edward Donovan (1768–1837 [born O'Donovan]; Salmon 2000:129–131, Gilbert 2004, Wilson 2004) published The Natural History of British Insects in 16 volumes (1792–1813) with 569 color and 7 black-and-white plates (Freeman 1980:113); and Adrian Hardy Haworth (1767–1833, Rauschenberg 1972, Salmon 2000:127–129, Erickson 2004) found sufficient market for his Lepidoptera Britannica (1803–1828) that all four volumes were published. Haworth also wrote a 69-page "Review of the Rise and Progress of Entomology in Great Britain" (1807). There were about 250 British entomologists in 1800 and about 500 in 1850 (Clark 2009:11). The year after Spence's complaint about ridicule of collectors, Pricilla Bell Wakefield (1751–1832), who wrote popular books on science (Freeman 1980:353–354, Shteir 2004), published An Introduction to the Natural History and Classification of Insects for children (Ordish 1976:121–122), which she is unlikely to have done if she and her London publisher had not been confident of its sale, though one might argue that she was capitalizing on the popularity of Kirby and Spence's first volume. It would be fairer to say that Kirby and Spence broadened the popularity of butterfly collecting to include other British insects. Commercial establishments were quick to provide equipment and supplies to collectors (Chalmers-Hunt 1994). Yet, as recently as 1902, Canadian entomologist Alston Moffat also complained that "the great majority of every community … seem to regard [entomology] as the frivolous pastime of a few harmless lunatics" (quoted in Riegert 1980:3). Kirby graduated from Cambridge University and followed the example of John Ray, William Derham, and Gilbert White in being a country parson, at Barham, who found time for serious study of insects within a natural theology context (Essig 1931:670–672, Salmon 2000:124–127, Clark 2004b, 2006, 2009:23–24, Moore 2004, Varma 2009). In contrast, Spence was a merchant and political economist at Hull, who was more practical minded (Salmon 2000:132–133, Clark 2004e, 2006, 2009:34–35). Kirby and Spence's Introduction to Entomology was popular because it was oriented to the public (Ordish 1976:123–126). Instead of the usual chapters of one insect order after another, as entomologists usually presented them, there were "letters" to the reader on topics of common interest, such as: 4. "Direct Injuries Caused by Insects (affecting Man personally)," including fleas, lice, and mosquitoes; 5. "Indirect Injuries Caused by Insects," to livestock; 6. "Indirect Injuries—continued," to crops; 9. "Indirect Benefits derived from Insects," as food for other animals, fertilization of plants. Other letters described the homes that social species construct, care of young, locomotion, self-defense, and hibernation. Kirby disposed of the concern that collecting insect specimens was cruel by arguing in letter 2 that insects do not feel pain (Kirby and Spence 1843, I:42–45, Hollerbach 1996:209–219). Spence wrote the first comprehensive account of insect diseases (1815–1826, IV:197–202, letter 44), but the only "diseases" to which they could assign a cause were those in which insects parasitized other insects (Steinhaus 1956:114–115, Cameron 1973:286). Agostino Bassi's demonstration (1835) that a silkworm disease, muscardine, was caused by a fungus is discussed in part 44 (Egerton 2012b:309–310). Ordish (1976:124–126) summarizes Kirby and Spence's discussion of insects that attack crops, but he comments that their approach was that of natural history, not eradication, and Kirby even inserted his view (not shared by Spence) that pest attacks were Divine judgment on human victims. However, preacher Kirby's emphasis was different in his 1833 two-volume Bridgewater treatise, On the Power, Wisdom and Goodness of God as Manifested in the Creation of Animals and in their History, Habits and Instincts (Blaisdell 1982:166–172). (a) William Kirby. Neave 1933: facing 125. (b) William Spence. Engraved by W. T. Fry from painting by J. J. Masquerier. Since Kirby and Spence were weak on combating insect pests, other books published in Britain filled the gap. In 1829, Joshua Major, a gardener, published A Treatise on the Insects Most Prevalent on Fruit Trees and Garden Produce, Giving an Account of the Different States They Pass Through, the Depredations They Commit, and Recipes for Their Destruction (Clark 2009:162–163). W. R. Shuckard translated from German Herman Burmeister's Manual of Entomology (London, 1836), but Ordish (1976:126) complains that its strength lay in physiology, not pest eradication. Jane and Mary Loudon translated from German V. Köllar's Treatise on Insects Injurious to Gardeners, Foresters and Farmers 1840, and J. A. Westwood supplied English annotations (Clark 2009:163). John Curtis (1791–1862) became known for two works: British Entomology (16 volumes, 1824–1839, 770 color plates), and Farm Insects 1860. He was the son of an engraver and became a skilled artist himself, though he could have learned limited skills from his father, who died in 1796 (Ordish 1974:3–4, Salmon 2000:138–139, Clark 2004a, Hooper and Foote 2004). He had a childhood interest in insects, and although he was poor, he attracted a patron who introduced him to Kirby. Kirby commissioned him to draw and engrave the plates for Introduction to Entomology (see Fig. 2). Curtis' British Entomology was outstanding because he excelled at both entomology and illustration. Since it was an expensive work, he published it in 193 issues. He drew and described the insects as he obtained them, and the work was not in systematic order, though most subscribers rearranged them into eight volumes in systematic order. He was inspired by Latreille's Genera crustaceorum et insectorum (four volumes, 1806–1809), and Curtis also intended to keep his illustrations and descriptions at the generic level. However, the announcement in 1826 of a rival work prompted him to change his plan with plate 150 and he began describing species within the genera (Ordish 1974:41–42). What he was accomplishing was widely appreciated, and he received honorary memberships in learned societies, and in 1842 Sir Robert Peel granted him a life pension of £100 a year. Even so, he was unable to afford membership dues for the Entomological Society of London until 1851. He was elected president in 1855, but resigned in 1857 because he could not afford to continue (Ordish 1974:92). Hymenoptera: 1. Sirex Gigas (center), 2. Evania appendigaster (right), 3. Nomada Marshamella (left); Diptera: 4. Pedicia rivosa (top), 5. Sericomyia Lapponum (bottom). Kirby and Spence 1843, II: Plate 4. Drawn and engraved by John Curtis. Curtis' biographer (Ordish 1974:78–79) believed Farm Insects founded economic entomology, which is true for Britain. However, Curtis himself, in complaining about the lack of economic entomology in Britain, cited works of Thaddeus Harris and Asa Fitch in America, Guérin Méneville and Bazin in France, Passerini in Italy, and others in Germany (Curtis 1860:xi–xii). Curtis' book was a slightly edited compilation of articles he had published in the Royal Agricultural Society's Journal (1841–1857). Ordish was aware of Major's book on fruit tree pests (1829) and Harris' Insects of Massachusetts Injurious to Vegetation (Harris 1841), but Major's book was limited in scope and Harris' book was organized according to insect systematics, and neither had illustrations. Curtis organized his book on a crop-by-crop basis, which was user friendly in the way that Kirby and Spence's Introduction had been, that is, according to the reader's desires rather than the author's education. But in 1855, Fitch (on whom, see below) began publishing his annual reports on New York insect pests, also organized by crops. Furthermore, in 1858, botany professor John Lindley praised Fitch's style and organization and urged Curtis to follow his example (Barnes 1988:60). Curtis' Farm Insects, with 16 color plates, 69 woodcuts, and a 524-page text, emphasized the importance of correct identification of insects in order to know the pests, their parasites, and predators. After explaining what was known about the life cycle of each species, he recommended what today is called integrated control—mechanical, chemical, and biological—though these weapons were rudimentary at the time (Ordish 1974:81–83). For example, his "biological control" of a turnip pest was the recommendation that the farmer hire about a dozen boys about age 10 to pick caterpillars off plants. Mr. Sells found that a 10-year-old could gather 180 caterpillars per hour, which, for an 8-hour day, means 9000 a week, or 90,000 a week for 10 boys, paid sixpence a pint, under a supervisor paid 2 shillings a day (1974:87). Young ducks ate the caterpillars. Curtis' Farm Insects was a good beginning of economic entomology in Britain. His successor was also a private individual, Eleanor Anne Ormerod (1828–1901), not the government (Clark 1992, 2004c, 2009:154–186). British entomology continued flourishing as science (as in Miall 1895), but not as applied science. Britain lacked the insect crises that challenged America, and when the British feared that the Colorado potato beetle might reach Britain, a London publisher (without permission) republished (1877) The Colorado Beetle, by expatriate Charles Riley, to explain how to deal with it if it arrived (Clark 2009:141). Ormerod grew up in a wealthy rural home in which her parents made no effort to find husbands for their three daughters (Clark 1992:433). She became interested in entomology in March 1852, when she obtained a copy of James F. Stephens' Manual of British Coleoptera or Beetles 1830. She was no doubt also encouraged by her brother Edward, who was writing a monograph on The British Social Wasps (Howard 1930:221). Entomology remained a hobby until Eleanor Ormerod responded to a plea by the Royal Horticultural Society (1868), published in the Gardeners' Chronicle and Agricultural Gazette, for someone to make a collection of insects harmful and helpful to British agriculture and horticulture. Next, she began publishing her 17 annual reports on methods of controlling insect pests, which her sister Georgiana illustrated (Howard 1930:221). These reports led to her becoming Honorary Consulting Entomologist to the Royal Agricultural Society of England. She also lectured at agricultural colleges, and in 1884 she published Guide to Methods of Insect Life, which later evolved into her Text-Book of Agricultural Entomology 1892. In 1900, the University of Edinburgh recognized her accomplishments with an honorary LLD degree, the first it gave to a woman. Economic entomology only became well established in Britain during World War I (Ordish 1976:149). Englishman Thomas Belt (1832–1878) had an early interest in nature, and at age 18 joined the Tyneside Naturalists' Club (Van Riper 2004). In 1852 he sailed to Australia and became an engineer at a gold mine. In 1862 he returned to England and joined the Geological Society of London. In 1868–1872 he managed a gold mine in Nicaragua, and afterwards wrote The Naturalist in Nicaragua 1874, which Darwin decided in that year was "the best of all natural history journals which have ever been published" (letter to Joseph Hooker, in Darwin 1887, II:366). Like Henry Bates, whose The Naturalist on the Rivers Amazon 1863 he often cited, Belt wrote about all aspects of nature, but especially about insects. Belt observed protective mimicry in the longicorn beetle Desmiphora fasciculate, covered with long brown and black hairs, resembling hairy caterpillars that birds avoid (Belt 1888:7, 1985). Foraging ants Eciton humata at times hunted in dense armies, or in columns when seeking nests of Hypoclines ants (Belt 1888:20–28). They seize Hypoclines eggs and larvae. Eciton species have small, imperfect eyes, or no eyes. E. humata seemed not to stay in one place more than four or five days, and temporary homes were in hollow trees or under logs. Belt put little pebbles on top of some E. humata and watched other ants rescue them, even going for help if needed. He thought Eciton ants were the most intelligent insects in Central America, with wasps and bees coming next. (a) John Curtis. Howard 1930: Plate 17. (b) Eleanor Anne Ormerod. Howard 1930: Plate 17. Bates had speculated that leaf-cutting ants Oecodoma used leaves to thatch tunnel entrances to their nests to avoid flooding (1864:11–18). Belt planted a garden and found leaf-cutting ants carrying off leaves of his vegetables (Belt 1888 [1985]:71–84). To drive them away, he dug up their nest, but then curiosity prevailed and he investigated their chambers where leaf cuttings were piled up, with fungi growing on them. Belt concluded that these ants ate the fungus, and he cited Fritz Müller's independent confirming conclusion. The remains of leaves after fungal growth served as food for Staphylinidae and other resident beetles. These ants carefully ventilate chambers and maintain constant temperature and humidity. Belt watched these ants cross a tramway for several days, with many crushed by wagon wheels; then they tunneled under the rails. (a) Nest of leaf-cutting ants. (b) Bull's-horn acacia thorns with ant hole near right tip. Belt 1888 [1985]:80, 218. Branches and trunks of bull's-horn acacias are covered with strong curved pairs of hollow thorns (Belt 1888:218–222). Small stinging ants Pseudomyrma bicolor make a hole near the tip of one thorn and live inside. During the rainy season, all thorns are occupied. Ants protect acacia leaves from browsing mammals and from leaf-cutting ants. In return, acacias provide shelter and also food (Belt 1888:219) The leaves are bi-pinnate. At the base of each pair of leaflets, on the mid-rib, is a crater-formed gland, which, when the leaves are young, secretes a honey-like liquid. Of this the ants are very fond … there is a still more wonderful provision of more solid food. At the end of each of the small divisions of the compound leaflet there is, when the leaf first unfolds, a little yellow fruit-like body united by a point at its base to the end of the pinnule. Examined through a microscope, this little appendage looks like a golden pear. Such observations made Belt "a pioneer in the study of tropical ecosystems" (Van Riper 2004:178). Modern ecologists have found the ant–acacia relationship as fascinating as Belt did (Brown 1960, Janzen 1966). Institutional support of entomology grew throughout the 1800s. Osborn (1952:24–27) has an international listing of early societies and periodicals. Salmon (2000:399–405) provides a detailed chronology of such societies in Britain. The Aurelian Society, in London, existed from 1801 until 1806. The Entomological Society of London, formed in 1807, published a volume of transactions (1807–1812), and faded away by 1822. In 1826, Edward Newman (1801–1876) was one of four founders of the Entomological Club (Endersby 2004, Foote 2004a), which published The Entomological Magazine, 1833–1838, under his editorship. It is the oldest existing entomological organization, with membership limited to eight. In 1833 the second Entomological Society of London was formed (Neave 1933, Pedersen 2002). Kirby was made honorary president for life, and he eventually presented his insect collection to it. Spence was president in 1847–1848 and vice-president in 1844–1846 and 1853. His son gave his library to the Society after his father's death. The Society's members would include Darwin, Wallace, and Bates. Darwin is listed as an original member (Neave 1933:121), though he was on the voyage of the Beagle in 1833: "Friends already had his name down for the Entomological Society, where he could be lionized for his tropical insects" (Desmond and Moore 1991:190). Not all members became evolutionists; a notable opponent was Oxford Professor John Obadiah Westwood (1805–1893), who was president three times and eventually made Honorary Life President (Neave 1933:131–132, Clark 2004f, Foote 2004b). The Society maintained peace by alternating evolutionists and nonevolutionists as presidents (Blaisdell 1992:242–277). Not all whose hobby concerned insects participated in the Entomological Society of London. The natural history community reflected the social classes of English society (James 1973:4). Wallace, Bates, and Spence, by virtue of scientific achievements, rose above their nonuniversity backgrounds and associated with that society's university men. Newman, by virtue of his editorship of natural history journals and authorship of numerous books (Freeman 1980:262–263), earned acceptance in the London Society and was elected to its Council (Neave 1933:9), but he was a tradesman who perhaps felt more comfortable with other tradesmen. In the 1850s and 1860s, regional entomological societies sprang up in London, and in 1872 Newman founded the South London Entomological Society that met at his house rather than at a pub, as the other regional societies did. It met there weekly until he died. In 1884 the name changed to the South London Entomological and Natural History Society, and in 1968 it became the British Entomological and Natural History Society. Explorer Alexandre Lefebvre (1798–1867) organized the Société Entomologique de France in Paris in 1832 and was its secretary until 1834 (Peyerimhoff 1932, Lhoste 1987:101–102, Gouillard 2004:19, Agular 2006:134–136). Pierre-André Latreille (1762–1833) was its first president, though he only lived another year (Nussac 1907, Burkhardt 1973, Lhoste 1987:48–51, Gouillard 2004:14–16, Agular 2006:76–79). Latreille was the "foremost entomologist of his time" (J. C. Fabricius, cited from Dupuis 1974:1), and a prominent influence in science generally (Schmitt 2010). He first attracted attention with his Précis des caractères génériques des insects (1796), and in 1798 he was appointed Assistant Naturalist at the Muséum National d'Histoire Naturelle. In 1830 he achieved a professorship. Although mainly a taxonomist (Wilson and Donner 1937:15–17), Latreille also studied insect anatomy and behavior. His Cours d'entomologie 1831 was probably the first general entomology textbook. The Société Entomologique began publishing its Annales in 1832, and later also a Bulletin; Osborn (1952:24) thought "these constitute one of the most important sources of entomological literature in existence." Five additional entomological societies arose in Europe by 1863 (Essig 1936:103, Richard 1973:478). Germany had more forests than any European countries west of it, and Germany pioneered forest entomology. Regensburg preacher Jacob Christian Schäffer (1718–1890) became alarmed at the damage done by a plague of the gypsy moth Porthetria (Lymantria) dispar in Saxony forests. He wrote a book (1752) describing its life cycle and illustrated its egg, caterpillar, feeding, pupation, and adult moth on a color plate. He attributed the moth plague to favorable weather, failure of natural enemies, and abundance of food. He was an amateur who laid a good foundation for this science (Schwerdtfeger 1973:363). A forestry magazine was founded in 1763. Several other authors wrote forestry books dealing with insects in the later 1700s. Johann Matthaus Bechstein (1757–1822), a theologian in Thuringia, was interested in hunting, forestry, birds, and insects. He founded a forestry school at Waltershausen in 1794 and became director of the forestry academy at Dreissigacker, 1801–1822. He began his prolific publishing with Gemeinnützige Naturgeschichte Deutschlands (1789–1895), followed by Naturgeschichte der Stubenvögel (1795) and Naturgeschichte der schädlichen Waldinsekten (Natural History of Injurious Forest Insects, 1798). He next collaborated with another parson, G. L. Scharfenberg, to edit Naturgeschichte der schädlichen Forstinsekten (three volumes, 1804–1805, 1042 pages, 13 plates), "the first reference book of forest entomology that deserves the name" (Schwerdtfeger 1973:367). Finally, Bechstein published a textbook, Forstinsectologie (1818, 551 pages) that showed a steady increase in knowledge over more than two decades of his studies. The first scientifically educated forest entomologist, Julius Theodor Christian Ratzeburg (1801–1871), was the son of a professor at the Berlin veterinary school and manager of the Royal Pharmacy (Schwerdtfeger 1973:367). The son studied medicine and pharmacy at the University of Berlin, taught there, and, with J. F. Brandt, compiled Medicinische Zoologie (two volumes, 1829–1833). In 1830, Ratzeburg, forester and mathematician, joined the faculty at a new forestry school at Eberswalde, 30 miles from Berlin. He collected literature on forest insects, supplemented it with information from foresters and naturalists, and had the ministry of forests ask all forest officials to send him information. They did, and his correspondence soon extended from St. Petersburg to London and from Sweden to Switzerland. He studied insects in forests, and he bred them in cages at different temperatures. Ratzeburg synthesized his findings in his great Die Forst-Insecten oder Abbildung und Beschreibung der in den Wäldern Preussens und der Nachbaarstaaten als schädlich oder nützlich bekannt gewordenen Insecten (The Forest Insects or Illustrations and Descriptions of those Insects which have Become Known as Either Harmful or Beneficial in the Forests of Prussia and the Neighboring States, three volumes, 1837–1844), a well-illustrated work that was still being praised, even in America and Russia, a century later (Schwerdtfeger 1973:369). He also summarized its information in a textbook, Die Waldverderber und ihre Feinde (The Forest Destroyers and their Enemies, 1841; edition 6, 1869, posthumous edition 7 edited by J. F. Judeich). He also described about 1000 species of Ichneumonidae in Die Ichneumonen der Forst-Insecten (three volumes, 1844–1852), these being "good insects" that parasitize species harmful to trees. H. Nördlinger began publishing Nachträge zu Ratzeburgs Forstinsekten (Supplements to Ratzeburg's Forest Insects) in 1848 and collected them into a second edition in 1880, and others also wrote volumes extending Ratzeburg's work (Schwerdtfeger 1973:371). Erasmus Darwin had discussed biological control of pests in 1800 (Egerton 2009a:184–186), and Kirby and Spence had cited his suggestions (Acot 1998, I:152–153). Later, French and Italian zoologists became interested in combating pest insects with predators or parasites (Howard and Fiske 1911:17–18). Professor Boisgiraud, of Poitiers, France, about 1840, placed carabid beetles Calosoma sycophanta on poplars attacked by gipsy moths Porthetria dispa, and the beetle controlled the moth population; he also destroyed earwigs in his garden by placing among them a rove beetle Staphylinus olens. These results were published by Professor of Zoology Nicolas Joly (1812–1885), University of Toulouse, in 1842. Joly's paper may have influenced Milan's Society for the Encouragement of Arts and Crafts to offer a gold medal in 1843 for the use of carnivorous insects to control a pest species of insect. Antonio Villa responded with a memoir (1844) that was praised by some, criticized by others, and soon forgotten. (a) Julius Theodor Christian Ratzeburg. Schwerdtfeger 1973:368. (b) Jean Henri Faber. In France, Edouard Perris (1808–1878) studied the buprestids, cerambycids, curculionids, and scolytids (Coleoptera) that attack Pinus maritime (Schwerdtfeger 1973:371–372, Lhoste 1987:266–268, Gouillard 2004:28–29). He had trees cut every month to follow these beetles' life cycles, described in his Histoire des insectes du Pin maritime (10 parts, dated 1851–1870, according to Schwerdtfeger, but 1852–1863 by Gouillard). In 1897 three Frenchmen published two studies on the populations of insect parasites. The Parisian zoologist Paul-Albert Marchal (1862–1942) wrote a brief paper, "L'équilibre numérique des espèces et ses relations avec la parasites chez les insects," arguing that oscillations in populations of crop pests, such as Hessian fly, oat midge, and army worm, was due to increase in their parasite's population following the increase in host population. When the host species' population crashed, the parasite population also crashed, which then allowed the host species to again increase (Thompson 1939:302). Ad. Bellevoye, member of the Société Entomologique de France, and J. Laurent, professor at the Lycée and at l'Ecole de medicine de Reims, published "Les plantations de pins dans la Marne et les parasites qui les attaquent." Their lengthy study provided "a mathematical theory of biological control" (Thomas 1939:304), though it attracted little attention in a provincial journal until discussed by Marchal (1907, 1908:353–355). Also in 1897, Leland Howard published on insect parasitism with observations and conclusions comparable to Marchal's, as discussed below. In the later 1800s and early 1900s, three notable European scientists investigated insect behavior (Richard 1973:482–485). They had very different backgrounds. Jean Henri Fabre (1823–1915) came from a poor rural family in southern France and had a long hard struggle to get an education and to support himself and his family (Legros 1913, Fabre 1921, Théodoridès 1971, Lhoste 1987:147–150, Pasteur 1994, Gouillard 2004:31–32, Agular 2006:114–116, 207). Sir John Lubbock (1834–1913) was the son of a British baronet-banker-mathematician, whose challenge was merely to make the most of his opportunities (Grant Duff 1924, Somkin 1973, Clark 1997, 2009:80–104, Alborn 2004, Barton 2004, Patton 2007). Auguste-Henri Forel (1848–1931) was a Swiss physician and psychiatrist who had developed an interest in ants as a child, which persisted into adulthood (Forel 1937, Pilet 1972, Lhoste 1987:150–152, Sleigh 2007:21–37). Although Fabre received a doctoral degree in science in 1854, he never held a university appointment and was self-consciously outside the mainstream of science. Constantine Rafinesque had been quite iconoclastic (Egerton 2009b:470–472, 2011a:69–70), and suffered neglect by other naturalists because of it, yet he nevertheless thought of himself as a member of the naturalist community. Fabre's publications were not hard to track down, as Rafinesque's were, but his published disdain for conventional naturalists undoubtedly led them to pay less attention to his writings than they might otherwise have. Nevertheless, Fabre had fervent defenders in the wider intellectual community, and Darwin admired his work. Fabre's strong interest in natural history predated publication of On the Origin of Species 1859, but his opposition to Darwin's theory of evolution by natural selection helped focus much of his research (Yavetz 1987, 1988). That did not preclude, however, his carrying out experiments that Darwin suggested to him. Each, of course, wanted to interpret Fabre's findings differently. Fabre believed that insects were endowed with instincts that could not be significantly modified. Two experiments that illustrate this involved wasps that sting and paralyze prey, which they store in a cell with an egg that hatch

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