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.

Europe

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).

Details are in the caption following the image — **Figure 1**
Open in figure viewer PowerPoint

(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).

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.

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.

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.

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 hatches and eats the prey. Fabre removed the egg and prey from a Sphex cell and placed them outside; the female returned with another prey and repaired the cell but ignored the egg and prey placed outside (1879–1907:I, 170–171). In another experiment, he removed both egg and spider from the cell of a Pelopoeus wasp; the female continued bringing back more spiders (they were removed as she searched for more), for up to 20 spiders, without her noticing removal of either egg or other spiders (1879–1907, V:32–49). Unlike Lubbock and Forel, Fabre saw no lessons for humanity in insect societies. He thought “animals were less like people than they were machines” (1918), echoing Descartes. However, modern researchers have found that the self-organizing behavior of ants, bees, and termites can indeed provide insights useful in human self-organizing social behavior (Miller 2007, 2010).

John Ray, as noted in part 18 (Egerton 2005:308–309), had a female moth (probably Pachys betularia) in a cage on the night of 29 May 1693, when his wife noticed and caught two males that had flown into an open window and were attempting to reach the female. Ray correctly deduced that they had smelled the female, and he reported this discovery of pheromones (but did not coin the term) in his Historia Insectorum (1710:117, translated by Raven 1942:395). Fabre, who seemed unaware of Ray's report, had essentially the same experience (Fabre 1949:75–86). On the morning of 6 May 1874, a female great peacock moth emerged from her cocoon in his laboratory, and he habitually put her under a wire gauze bell jar and returned to his other work. At nine p.m. his son screamed, “Come quick” (in French), and Fabre entered his room and saw male great peacock moths flying about. Still others were in the laboratory where the female was confined. Unlike Ray, Fabre was an experimentalist, and he cut off antennae from males captured before releasing them to see if they would return the next night. Only one did. He marked others captured to see if they would, but his success was only slightly better. The female lived eight days, during which time he captured, marked, and released 150 males. He continued his experiments each May for two more years.

John Lubbock's father sent John to Eton at age 11, but removed him after three years and made him a partner in the family bank, since banking was interfering with his father's mathematical studies of tides (Pumphrey 1958:51). Fortunately, a friendly neighbor, Charles Darwin, was glad to assist in John's informal education at home (Patton 2007:18–22). After Darwin persuaded the father to buy John a microscope, John returned the favor by drawing barnacles for Darwin's barnacle monographs. They were very close friends for 40 years. Lubbock contributed new information to the first, second, and fourth editions of On the Origin of Species (Somkin 1962).

Lubbock published a study on Daphnia reproduction (1857) that won him admission into the Royal Society of London. He published nine studies on ants, bees, and wasps in the Linnean Society of London's Zoology Journal, 1874–1881 (listed in Donisthorpe 1924:167, Patton 2007:258–259), which he then collected in modified form into Ants, Bees and Wasps: a Record of Observations of the Habits of the Social Hymenoptera (1882, 17 other editions until 1929). The book has nine chapters on ants and one each on bees and wasps; and it has a three-page bibliography of previous studies on these social insects. Lubbock noted that his studies differed from earlier ones because he marked individual insects with painted dots or wing snips and observed them in social situations. Perhaps he published this technique before Fabre adopted it, though Fabre might have adopted it independently. Lubbock was also first to keep ant nests for more than a few months, and he devised containers, which he described but did not name (Donisthorpe 1924:159 called them “Lubbock nests”; Pumphrey 1958:55 called them “observation nests”), that enabled him to keep the nests for up to eight years (Lubbock 1899:2–4). He took two glass panes, 10 inches square, separated by a wooden frame around the perimeter that was only 1/10 to1/4 inch thick. The inside space was filled with soil, and obviously one wooden edge had to be removable, for the sake of the introduction of ants and then of food and water, though he gave no details. These nest containers were thin enough to enable him to observe the movements of individual ants, and Lubbock's house had 30 to 40 of them at various times. His marked individuals enabled him to dispense with estimates of longevity given by previous investigators. He had workers that lived for 7 years and two queens, of different genera, lived nearly 14 and nearly 15 years. Eggs reportedly hatched in about 15 days, but his took 6–7 weeks; larvae reportedly grew for 6 or 7 weeks, but he found that Myrmica ruginodis entered a pupa in less than a month, though other species took longer (1899:6–7, 11–12). Although all of Lubbock's chapters on ants relate to ecological topics, the most directly relevant are Chapter 3 on ant relations to plants, and Chapter 4 on other animals and ants.

He began Chapter 3 by noting that ants have not exerted as strong an influence over flowering plants as have bees, since bees fly from one plant to another, whereas ants move about on the same plant and then move on to another plant of whatever species, whereas bees stick with the same plant species during any period of collecting nectar or pollen. Belt found that flowers have contrivances to prevent random insects like ants from gaining entry into their nectaries. Lubbock discussed the various means different plant species use to effect this (Lubbock 1899:52–56), and he reviewed Belt's other discoveries. He cited Forel's discovery at an anthill, that returning ants brought back small caterpillars and other insects at the rate of 28 per minute, from which Forel calculated that the anthill consumed 100,000 insects a day, which Lubbock cited in his chapter on ants and plants under the presumption that plants benefited from that consumption, though he did not consider whether the 100,000 included some predatory and parasitic insects that also destroy other insects. Ants in southern regions store grain for the winter, but not those in northern Europe (1899:59–61). Lubbock did not explain why, but presumably northern ants hibernate in winter and therefore do not consume food then.

Lubbock fed his captive ants mainly honey, but he knew that in the wild, English ants were mostly predators and scavengers. He quoted Savage, “On the Habits of the Driver Ants” (1847) of Africa (Lubbock 1899:63–65), which eat all animals in their path, and he compared them to the army ants of Brazil, Eciton vastator and E. erratica, described by Bates (Egerton 2011c:42–43) and Belt (see above). Both groups of ants are blind and travel at night. Belt observed a spider and a harvestman Phalangium avoid army ants, but other insects became victims. Belt found a spider species that mimicked the form, color, and movements of ants and at first deceived him. Mites live in ant nests and parasitize them, as do some species of Diptera in the family Phoridae. Lubbock was especially interested in the positive relations between ants and other animals. He commented (1899:67) that it had long been known that certain species of aphids are “cows” of some ants. Aphids suck juices from plants, but then must expel large quantities of liquids. However, they wait for the ants that protect them to come and stroke them with their antennae before expelling it. He explained that aphid “honey” is sticky, and is relinquished to ants not just for protection services but also because it would be a messy waste that could encumber the aphids. He praised Rev. William Gould's An Account of English Ants (1747), but pointed out that the brown eggs that Gould identified as eggs for male Lasius flavus were actually aphid eggs, which this ant species tended as it tended its own eggs. Furthermore, the ants gathered the eggs from the plants where the aphids laid them and carried them into their nest, where they were protected during winter. Lubbock found in the nest of Lasius flavus “four or five species of aphis, more than one of which appears to be as yet undescribed” (1899:73). He also found several other species of insects that lived in ant nests unmolested by the ants, some of which had no detected benefit for the ants, one of which ate ant eggs, but other species secreted a substance that ants might eat. The claviger beetle is blind and seems to have lost the ability to feed itself and was fed by ants. He thought there was much about these relationships between ants and cohabiting other insects yet to be learned. Swiss naturalist Jean-Pierre Huber (1777–1840) discovered the existence of slavery in ants, explained in his Recherches sur les moeurs des Fourmis indigènes 1810. Lubbock quoted on this from the English edition of Huber's book. Lubbock did not attempt to explain how slavery evolved among ants, but he did comment on its consequences (1899:82).

Polyergus fufescens present a striking lesson of the degrading tendency of slavery, for these ants have become entirely dependent on their slaves. Even their bodily structure has undergone a change: the mandibles have lost their teeth, and have become mere nippers, deadly weapons indeed, but useless except in war. They have lost the greater part of their instincts: their art, that is, the power of building; their domestic habits, for they show no care for their own young, all this being done by the slaves; their industry—they take no part in providing the daily supplies; if the colony changes the situation of its nest, the masters are all carried by the slaves on their backs…

Lubbock was able to see all this for himself, because Forel had sent him a Polyergus colony which he observed for over four years.

Forel was the son of a prosperous Swiss surveyor and farmer; his great-uncle, Alexis Forel, was a botanist and entomologist. As a child, Auguste was allowed to collect insects “on the condition that I let him show me how to kill the creatures painlessly” (Forel 1937:33). At this news, his joy was “indescribable,” but:

I was to have even greater cause for rejoicing. Not only did my father give me a copy of old Réaumur's work, Mémoires pour servir à l'histoire des insectes; not only did my great-uncle most kindly come to my assistance, identifying my beetles and leaf-bugs whenever I applied to him, but in the same year my grandmother herself came to me and said: ‘Just think, I once had a dancing-partner who was a great lover of ants. He always scolded me when I destroyed the creatures, which used to eat my candied fruits; he was such a kind man, and such a lover of animals. He gave me his book; I never finished reading it, but it is said to be a work of great scientific value. Here, take it, but…never torment the little creatures again as you have done hitherto.” With these words she gave me the well-known book of Pierre Huber, Recherches sur les moeurs des fourmis indigenes (Geneva, 1810), with the author's dedication to my grandmother. It was the first edition. I didn't read this book, I devoured it!

In a flash all the mysteries of ant life, which had hitherto been enigmas to me were revealed. Huber's book literally became my Bible, and I swore that I would one day become his successor as the historian of ants.

Forel read about slave-owning ants, which he had already observed, and at age 11 he had also seen something that Huber had missed—lestobiosis, the habits of killer and robber ants. It became the subject of his first scientific paper. His studies on ants fit into a Swiss tradition begun by Charles Bonnet in the 1700s and including François Huber (studied bees) and his son Pierre (studied ants) earlier in the 1800s (Sartori and Cherix 1983, Drouin 1992, 2012, Sleigh 2007:25). Although Forel became a physician and psychiatrist, his interest in ants remained strong throughout his life. His Les fourmis de la Suisse appeared in 1874, but his Le monde social des fourmis (four volumes) and Recherches sur les moeurs des fourmis indigenes appeared during the 1900s. Remembered for his studies on ant habits, he also described over 3500 new species (Pilet 1972:74).

William Morton Wheeler, whose own studies on ants began in 1900 (Evans and Evans 1970:126), had an opportunity to visit Forel in Switzerland in June 1907. Forel took him to see some of his favorite collecting grounds and told him of other places to visit (Evans and Evans 1970:144–145). In 1930, Wheeler reviewed Forel's book, The Social World of the Ants Compared with That of Man, and he offered this retrospective commentary (Wheeler 1930).

The observations which he made during his childhood and youth were brought together and published in 1874 as a quarto volume, the “Fourmis de la Suisse,” which is one of the landmarks of myrmecology, and has lost none of its value during all the years that have elapsed. This was only the beginning of his penetrating studies of the ants, which were carried on without interruption till 1922, when illness and fading eyesight compelled to write his last myrmecological paper and to dispose of his very valuable collections to the Museum of Geneva. Since 1874, he has published hundreds of important monographs and notes, containing descriptions of several thousand species, subspecies and varieties from all parts of the world and treating of every aspect of ant structure, distribution, psychology, and behavior. But all of this was only a delightful avocation!

Nevertheless, this avocation had a serious aspect. Forel came out of World War I with greater optimism about ant societies, which ran smoothly from instinct, than he had for human societies, which depend heavily upon culture (Lustig 2004:288–291). As a psychiatrist, he had no illusions about human potentials for destructive behavior, and as a myrmecologist he understood that natural selection had given ants instincts which placed the welfare of the colony above that of individuals. The best he could do was to argue that the success of ants should inspire us to emulate them in seeing the advantages of cooperation, and also learn the lesson of World War I (as now named) to see the high price of not cooperating. He had suggested a world government before the idea of a League of Nations was developed, and the human dilemma after 1918 made the idea compelling (Forel 1928:II, 337–352).

North America

Although initially entomology progressed more slowly in America than in Europe, the United States and Canada were largely undeveloped in 1800, and they gained momentum throughout the century. (Mexico is omitted here.) America began inventing itself as the British industrial revolution reached its shores, and it adapted more easily to industrialization than did continental Europe, since there was less to tear down to make way for new technology and factories. Entomology grew with the new nation, with Philadelphia and Cambridge being early centers.

Frederick Valentine Melsheimer (1749–1814) reached America two years before the Hessian fly. He was born in the Dukedom of Brunswick, where his father was superintendent of forestry (Osborn 1937:13–23, Mallis 1971:9–12, Stentsinger 1973). He attended the University of Helmstaedt and became acquainted with entomologist A. W. Knoch. In 1776 he was appointed chaplain to a regiment of mercenaries for Britain who came to America. He and his regiment were soon prisoners of war, and he resigned, married, became a minister in Hanover, Pennsylvania, and had 11 children. He corresponded with Knoch in Brunswick and sent him over 700 American insects, and Knoch acknowledged his assistance when publishing on them and dedicated his book (1801) to him. Melsheimer compiled A Catalogue of Insects of Pennsylvania (60 pages, 1806), which was really a list of 1363 species of beetles, and was the first separate entomological publication in America.

Melsheimer made a careful collection of Coleoptera and Lepidoptera, which had labels on the pins. He interested two of his sons in his project, and after his death his collection and library went to the older son, Rev. Johann Friedrich Melsheimer, who planned to collaborate with Thomas Say on a catalog of Coleoptera of North America; however, J. F. Melsheimer died in 1829 and Say in 1834, without publication. In 1842, the younger son, Dr. Frederick Ernst Melsheimer, and a few others formed the Entomological Society of Pennsylvania—the first entomological society in America—which lasted about a decade (Sorensen 1995:15–31). A lack of involvement in economic entomology may have contributed to its demise. The society wanted to publish the Coleoptera catalogue, and the Smithsonian Institute published it in 1853; it was written by F. E. Melsheimer, and revised by S. S. Haldeman and John L. LeConte (Osborn 1937:15–16). In 1859, some members of the previous society formed the Entomological Society of Philadelphia. In 1864 Dr. Melsheimer sold the family's world-wide insect collection of 14,774 specimens to Louis Agassiz for Harvard's Museum of Comparative Zoology for $150 (Hagen 1884).

While entomology was becoming established in the East, exploration continued in the West. John Kirk Townsend (1809–1851) was the first known collector of northwestern insects, 1834–1836 (Hatch 1949:1–2, Mearns and Mearns 2007, Egerton 2011a:80). Townsend was an ornithologist and various specialists described his insect collection. In 1841, two naturalists with the U.S. Exploring Expedition, Charles Pickering (1805–1878) and Titian R. Peale (1800–1885) collected from the Columbia River south to California, but no report on their insects was published (Hatch 1949:2, Egerton 2011b:154–160). Other expeditions followed, with more collections and publications (Hatch 1949:2–5).

Academic entomology

The Melsheimers had no professional entomological affiliation, but subsequent notable entomologists were usually employed by government or educational institutions.

William Dandridge Peck (1763–1822) was born in Boston and graduated from Harvard in 1782 (Essig 1931:729–732, Mallis 1971:13–16, Sorensen 1995:8–10, 1999). Although his father was a naval architect, they went to live on a farm in Maine, where William became interested in insect pests. In 1796 he won $50 and a gold medal from the Massachusetts Society for Promoting Agriculture for an article on the life history of the cankerworm. Other papers established his reputation, and in 1805 he became the first professor of natural history at Harvard. He was sent to England, France, and Sweden to study for three years. He had several strokes before he died, which impaired his teaching, but he inspired one student to become an entomologist. Thaddeus William Harris (1795–1856) was the son of Rev. Thaddeus Mason Harris, author of Natural History of the Bible. The son was an undergraduate at Harvard in 1811–1815 and obtained an M.D. there in 1820 (Howard 1930:30–35, Essig 1931:651–653, Mallis 1971:25–33, Sorensen 1995:11–13, Elliott 1997, 1999, 2008). He later commented about Peck: “It was this early and much esteemed friend who first developed my taste for entomology, and stimulated me to cultivate it” (Mallis 1971:27). Harris' father had been librarian at Harvard in 1791–1793, and Harris was appointed to the same position in 1831 and held that position for the rest of his life. He began publishing articles on economic entomology in the New England Farmer in 1823, and he aspired to the chair of natural history, but it went to the botanist Asa Gray in 1842. Nevertheless, as we saw in part 39 (Egerton 2011:252), he was teaching a course in natural history in the mid 1830s, and Thoreau was one of his students (and friends).

Academic entomology got off to a good start at Harvard with Peck and Harris in the first half of the 1800s, but by the third quarter of the 1800s Harvard had only botanist Asa Gray and zoologist Louis Agassiz, both outstanding, but not entomologists. Agassiz founded the Museum of Comparative Zoology in 1859 (Winsor 1991), where he had several graduate assistants who studied entomology. Alpheus Spring Packard, Jr. (1839–1905) graduated from Bowdoin College in 1861, and in the fall he entered Harvard's Lawrence Scientific School and became a museum assistant in entomology (Cockerell 1920, Essig 1931:727–729, Dexter 1957, 1997, Mallis 1971:296–302, Norland 1974, Bocking 1999). Packard was the son and grandson of presidents of Bowdoin College, in Brunswick, Maine. Packard also studied medicine at Harvard, yet he received his M.D. degree from Maine Medical School, at Bowdoin. He then served as an assistant surgeon with the Maine Veteran Volunteers during the Civil War, 1864–1865. Afterwards he taught entomology at Maine State Agricultural College, Massachusetts Agricultural College, and Bowdoin College (Cockerell 1920:194). In 1867, Packard and two colleagues founded the American Naturalist.

Packard had worked on the Maine Geological Survey during the summers of 1861 and 1862, he worked on the Kentucky Geological Survey in the summer of 1874, and he worked with the Hayden Survey of the West for the U.S. Geological Survey in the summers of 1875 to 1877. He had been introduced to the cave fauna of Mammoth Cave, Kentucky in the summer of 1871, after the meeting of the American Association for the Advancement of Science in Indianapolis. The Louisville and Nashville Railroad invited the scientists at the meeting to take an excursion to the cave. Packard was fascinated by the cave fauna, and that interest motivated his involvement with the Kentucky and Hayden surveys, both of which afforded him the opportunity to study cave invertebrate faunas in more detail (Bocking 1988). Packard became a leader of American “neo-Lamarckians”—he coined the term and wrote a biography of Lamarck (1901)—and he saw the blind cave fauna as evidence for his theory of evolution (Bocking 1988). Neo-Lamarackians believed that the environment played a strong role in shaping evolution, and disagreed with the Darwinians who stressed random variation and competition. His extensive studies led to a monograph on the invertebrates of North American caves (1888), which is important for ecology as well as for evolutionary theory. His example of studying cave fauna to investigate evolution is still being followed (Culver et al. 1995).

Another example of Packard's ecological interests is his article, “Nature's Means of Limiting the Numbers of Insects” (1874). He discussed birds and parasitic insects as nature's means to check the increase of insect pests. This was far from being a new subject, but his compilation is nevertheless an interesting indication of the knowledge of that time. Packard had previously speculated in print that few birds seemed to eat canker worms, and that apparently motivated C. J. Maynard of Ipswich to open the stomachs of some 3000 songbirds, and he sent Packard a list of species that he found did eat canker worms. He also mentioned other pest species that he found his birds had eaten, and Packard quoted Maynard's information. This letter apparently prompted Packard to solicit similar information from other ornithologists, and he quoted from a reply from Thomas M. Brewer. Packard also found a relevant article by Perris (discussed previously), “Les Oiseaux et les Insectes,” (1873), from which Packard quoted extensively in English translation. Packard then discussed moles, field mice, and frogs that eat insects. Finally, he discussed insects that parasitize other insects, and he suggested that the European Hessian fly and wheat midge might be controlled by importing parasites that attack them in England and France.

In 1878, Packard became Professor of Zoology and Geology at Brown University. An active researcher, he wrote five textbooks on entomology, four on zoology, and six monographs on Lepidoptera. His Guide to the Study of Insects, and a Treatise on Those Injurious and Beneficial to Crops 1869 has 702 pages. Packard's bibliography lists 579 publications (Cockerell 1920:207–236). He was a leading American entomologist at a prestigious university and one of the editors of a leading natural history journal with numerous publications (Dexter 1981). Nevertheless, as a trainer of entomologists he was overshadowed by Comstock at Cornell (see the following paragraphs). The reason was probably because Cornell was both a university and an agricultural college, and many entomology students were oriented toward applied aspects of entomology, so they went to Cornell rather than to Brown.

In 1863, Agassiz hired Philip Reese Uhler (1835–1913) as curator of insects in the Museum of Comparative Zoology, and Uhler stayed four years (Mallis 1971:205–208, Ignaciuk 1997). In 1867, Agassiz hired German entomologist Hermann August Hagen (1817–1893) to replace Uhler (Essig 1931:643–646, Mallis 1971:119–126). Besides numerous articles and two monographs, Hagen had published Bibliotheca Entomologica (two volumes, 1862–1863). In 1870 Hagen became Harvard Professor of Entomology, the first so titled in America (Howard 1930:73). In 1874, Hagen invited 12 men to meet in his house to form the Cambridge Entomological Club (Matthews 1974). He had a series of graduate student assistants until he was incapacitated by a stroke in 1890.

By then, leadership in academic entomology was shifting to a new university, Cornell. John Henry (“Harry”) Comstock (1849–1931) was born on a Wisconsin farm, but after his father died en route to the California gold fields, his mother took Harry back to her home in New York State. There she worked as a live-in nurse and he grew up with relatives and friends (Essig 1931:575–578, Needham 1946, Comstock 1953:2–34, Mallis 1971:126–138, Smith 1976, Henson 1996, 1999b). His early formal education was limited, and his youth might have been grim if he had not had a pleasant personality and a willingness to work hard. One summer he worked as a cook on a Great Lakes ship, and while stopping over in Buffalo, he spent $10 for a copy of the illustrated edition of Harris' Insects Injurious to Vegetation Harris 1841), and it gave direction to his life.

Comstock entered Cornell in autumn 1869, a year after it opened. It was a land-grant college at Ithaca, New York, whose federal endowment was supplemented by the wealth and land of the remarkable Ezra Cornell, advised by Andrew D. White, who became its first president. Comstock had gotten malaria in the summer, and it forced him to drop out. He returned in autumn 1870, and in January 1871 he became assistant to Professor Bert Green Wilder (1841–1925), who taught vertebrate zoology. President White had asked Louis Agassiz to recommend someone to head natural history, and Agassiz had recommended Wilder, who had degrees from both Harvard's Lawrence Scientific School and Medical College. Comstock and Wilder were impressed with each other when they met. However, Comstock soon got typhoid fever, and Wilder and his wife nursed him back to health. There was no entomologist on campus, and Comstock studied it alone until spring 1872, when 13 students petitioned to have him teach them a course for credit on insects injurious to vegetation. Their petition was granted, and in the summer Wilder arranged for Comstock to go to Harvard and study under Hagen—Hagen's first student. In spring 1873, the Cornell board of trustees appointed Comstock Instructor in Entomology, and he began teaching while still pursuing his own education. He graduated in June 1874, and Wilder took him along to teach entomology when he went to Peoria, Illinois to teach summer school.

Anna Botsford (1854–1930) grew up on a New York farm with a mother who loved flowers and taught her to know them (Needham 1946, Comstock 1953:53–73, Mallis 1971:131–135, Smith 1976, 1999, Bonta 1991:154–166, Norwood 1993:77–81, Henson 1996, 1999a). She attended private schools (where she rebelled against religious instruction) and entered Cornell in 1874. In 1875 she took Comstock's entomology course. She and Comstock ate in the same place, and he sat beside her. They collected insects together, and married on 7 October 1878. He became Assistant Professor in December 1876. In July 1878 he accepted Riley's invitation to study the cotton worm Alabama argillacea in Georgia while Riley studied it in Alabama (Comstock 1953:89–96). Afterwards Comstock recommended one of his capable students, Leland Howard, to Riley, and Riley hired Howard as his assistant. The Comstocks were unable to have children. They compensated by having an active social life and devoting much time to research and publishing on entomology. In 1879, Riley disagreed with the Commissioner of Agriculture, William Le Duc, and resigned as head of USDA's Entomology Division. Howard urged Harry to apply for the position; he did and was appointed. Although Comstock had a copy of his own notes on the cotton worm, Riley took his notes and specimens when he left. Comstock needed them in order to write a report, but Riley ignored his request for them, and Le Duc had the U.S. Attorney General get them from Riley. Before publishing his cotton worm report, Comstock had requested additional funds from Congress to expand the work and personnel of the Entomology Division. Congress was unresponsive until the report was published and given to each Congressman on 18 May 1880. Congress was impressed with the 343-page report and voted the requested funds. Entomologists were also impressed (Howard 1929:135, 1930:58, Comstock 1953:117–124). While head of the Entomology Division, Comstock also conducted important studies on the insect pests of citrus trees in Florida and California. Anna made expert drawings of scale insects to accompany Harry's report, and he sent a copy to the French authority on Coccidae, Signoret, who replied: “The drawing of the male is magnificent. It was made by the hand of a master” (Comstock 1953:130). This encouragement led to Anna's career as an illustrator. She later went to New York City and studied engraving under a professional.

When a new presidential administration came to Washington in 1881, Riley returned as head of the Entomology Division and the Comstocks returned to Cornell, with a USDA grant to enable him to complete his report on scale insects, published in 1881 with Anna's illustrations (Comstock 1953:135–141). She earned her bachelor's degree in June 1885 with a thesis on the internal anatomy of a larval Corydalus cornutus. On 1 November 1888 the Comstocks published the first part of An Introduction to Entomology, with his text and her illustrations, of eight insect orders. Harry decided that his German language skills were inadequate for him to continue, and so he and Anna traveled to Germany, visiting zoologists in Heidelberg, Leipzig, Dresden, and Berlin. After they returned, Harry became dissatisfied with the first part of his Introduction, despite its popularity at Cornell and at other universities, and instead of writing a second part, they produced a new book, A Manual for the Study of Insects 1895. The Comstocks' popularity with students, faculty, fellow entomologists, and people of New York State enabled him to acquire the first university department of entomology, and Harry Comstock had his own building on campus, the Insectary, for office and raising insects, another first (Comstock 1953:161). Comstock founded and ran a successful research school at Cornell in evolutionary entomology, with an impressive list of distinguished graduates (Evans 1975:xxiii, Henson 1993).

Government entomology

An advantage of the United States, with each state having its own government, is that when one state innovated, others could see whether it was successful, and if so, then they could follow that example, making any improvements desired. North Carolina, Georgia, and Pennsylvania founded universities in the mid 1790s, with other states soon following. North Carolina established the first geological survey, in 1823, and other states quickly followed, sometimes adding a natural history survey of plants and animals (Hendrickson 1961). Another example of states' innovation is appointing state entomologists, in which New York led the way.

States established geological surveys, 1823–1840s, to identify mineral wealth, and some states expanded their survey to include native flora and fauna (Merrill 1920, Hendrickson 1961). Massachusetts was one of those that expanded the scope, in two stages. Congregational minister and Amherst Professor of Chemistry and Natural History Edward Hitchcock (1793–1864) was in charge of the survey (Aldrich 1972, Guralnick 1972). In Hitchcock's Report on the Geology, Mineralogy, Botany and Zoology of Massachusetts (Amherst, 1833), Thaddeus Harris published a “Catalogue of Insects,” which listed and classified 2350 species. Harris later wrote a separate volume on insects. Influenced by Peck's strong interest in agricultural pests, Harris had already published articles on economically important species. His mandate was to make his volume useful for agriculture. Not all of the survey volumes attracted a broad readership, but Harris' Treatise on Some of the Insects of New England, Which Are Injurious to Vegetation (1841, new editions 1852, new editions 1862, new editions 1970) was quite influential. A later edition was illustrated—possibly influenced by the fine illustrations in John Curtis' Farm Insects—with black-and-white illustrations and eight color plates. As an example of Harris' accounts, he quoted from several authors on its life history, reviewed the early evidence against the Hessian fly coming from Europe, and cited more recent evidence of it being found near Toulon in France, near Naples in Italy, and on the island of Minorca (1841:422).

During the 1800s in America, agriculture grew as fast, or faster, than industrialization, and economic entomology flourished. In the 1840s, there were about a dozen American entomologists, in the 1850s about 30, in the 1860s over 40, and in the 1870s about 800 (Sorensen1995:38). However, Sorensen's last figure is questionable, as the Naturalists Directory for 1880 reported 436 entomologists (Smith 1989:13), which was still more than for any other zoological discipline. Some entomologists were in universities, as discussed above; states began appointing state entomologists, and the federal government sometimes had one at the Smithsonian Institution, and in 1881 when Riley went to the USDA Division of Entomology, it had four entomologists under him (Weber 1930:8–9). Although both numbers, 800 and 436, apparently included Canada, most of these entomologists were probably amateurs. Natural history became a popular hobby, which benefited entomology (Barnes 1985). The establishment of land-grant colleges in 1862 and of state agricultural experiment stations tied to those colleges in 1887 were momentous events for entomology as they were for phytopathology, discussed in part 44 (Egerton 2012c).

In 1854, the federal government hired its first entomologist and New York hired the first state entomologist. The federal entomologist was Englishman Townsend Glover (1813–1883), who came to America in 1836 to visit relatives and stayed (Howard 1930:35–43, Mallis 1971:61–69, Sorensen 1995:72–74, Cammarata 2004). He married into a family that owned a large estate on the Hudson River, and Glover merged his interest in insects with running a farm and orchard. He attracted favorable attention at agricultural meetings and fairs by displaying a large collection of models of fruits. His display of them in Washington led to his being hired as entomologist by the agricultural division of the U.S. Patent Office, where he published annual reports, 1854–1858 (Weber 1930:3–7). He resigned in 1859 to teach at the Maryland Agricultural College (now University of Maryland), but in 1863, he became the first entomologist at the newly created USDA, and he subsequently published 17 annual reports. The one for 1866 was only 18 pages, un-illustrated. However, Glover was a fine artist, and his 19-page report for 1867 is well illustrated with his drawings (Fig. 7b), and in 1877 his 60-page report was illustrated with five well-executed plates. Glover was quite interested in developing an agricultural museum (displaying his models), but he failed to understand the need for an authoritative collection of insect specimens and did not build one. He did take several trips into the South to study insect pests, but when agricultural insect crises emerged in the 1870s, he did not adequately respond and resigned in 1878 for health reasons.

In Canada, the Dominion Department of Agriculture appointed its first entomologist, James Fletcher, in 1883, to advise farmers about insect pests (Palladino 1996:51–54). Fletcher established a network of over 400 farmers to report on insect pests. He worked closely with William Saunders of the Entomological Society of Ontario to prepare a report on agricultural colleges and experimental stations in Canada and the United States for the Dominion Parliament. It was impossible to tie experimental stations to agricultural stations, as in the United States, because of stipulations in the British North America Act, and William Weld, editor of Farmers' Advocate and Home Magazine, argued that Canada did not need experimental stations because Canadians could just use reports published by the American stations. Nevertheless, Parliament established five experimental farms across Canada. Canada's higher education system was slow to train applied entomologists, and those hired for the experimental farms were trained either in Britain or the United States.

New York appointed the first state entomologist, Asa Fitch (1809–1879), in 1854 (Howard 1930:43–50, Essig 1931:632–634, Mallis 1971:37–43, Rezneck 1972, Barnes 1988, 1999, Sorensen 1995:71–72, Palladino 1996:23–24, Sterling 1997b). He was born in Salem, New York, north of Albany, and lived all his life there on his family farm. His father, Dr. Asa Fitch, had a successful practice and a large library. The son entered Rensselaer School in Troy in 1826. It had been founded by Stephen Van Rensselaer in 1824 to train teachers of science and its applications. After a year there, he studied medicine in Albany and New York City because of the poor chance of finding work as an entomologist. However, he found time to practice medicine, run the family farm, and study insects. The New York State Agricultural Society was formed in 1832, and many of Fitch's more than 140 publications were in its Transactions (his bibliography is in Barnes 1988:76–82). His most important publications were 14 annual “Reports on the Noxious, Beneficial, and Other Insects of the State of New York” (1855–1872). Fitch successfully wrote them for two audiences: farmers and entomologists. These reports were so popular the state legislature often reprinted them, and they became a model for other states and were highly valued in Europe. In 1858 he was awarded a gold medal by the French Section of Natural History of the Imperial and Central Society of Agriculture, and he was elected a corresponding member of entomological societies in Philadelphia, Germany, and Russia (Barnes 1988:60). He occasionally lectured on entomology at his alma mater, Rensselaer School (now Rensselaer Polytechic Institute).

In the 1840s, wheat was an important crop in New York State, but it was besieged by two insect pests, the Hessian fly and the wheat fly or midge. Fitch's publications in 1845 and 1846 were devoted to these species (Fig. 7). He thought the wheat fly was a European species, which it may have been, though not the one he identified (Barnes 1988:63–64). Working on that assumption, he concluded that it was more destructive in America than in Europe. He reasoned that in Europe it had parasites that kept its numbers down, but that these parasites had not come to America with the fly. After he began corresponding with John Curtis, he wrote to him in May 1855 and requested he send some parasites to him in Salem. Curtis thought this was a good idea, but not having any live wheat fly parasites, he took the suggestion to a meeting of the Entomological Society of London and requested help from its members. Nothing came of it. Barnes (1988:64) and Doutt (1964) thought this was probably the first suggestion of biological control by introduced parasites, though we have seen (Riley 1931, Egerton 2009:185–186) that Erasmus Darwin had made such suggestions in 1800. Fitch also saw that no American insects fed on the naturalized toadflax and suggested that European insects that feed on it be imported. Benjamin Walsh generalized from these two suggestions from Fitch: “Whenever a Noxious European Insect becomes accidentally domiciled among us, we should at once import the parasites and Cannibals that prey on it at home” (Walsh 1866a; quoted from Barnes 1988:65). After Fitch died, his collection of over 120,000 specimens was sold, some to the State Museum in Albany and some to the U.S. National Museum.

Illinois and Missouri appointed state entomologists in 1867 and 1868, respectively. Coincidentally, both had immigrated from England. Benjamin Dann Walsh (1808–1869) had collected insects in England (Tucker 1920, Howard 1930:50–53, Mills 1958:90–91, Mallis 1971:43–48, Sorensen 1995:74–75, Garber 1999). He had been Charles Darwin's classmate at Cambridge University, and, as he wrote to Darwin later, he had admired Darwin's beetle collection at Cambridge (Darwin 2001–2008, XII:161). They corresponded after Walsh settled in America. He came to America at age 30 with his wife, bought a farm in northwest Illinois, and farmed for 12 years. Then he ran a lumber yard in Rock Island, 1851–1858. Before retiring, he built a row of brick tenements, and then became a landlord, with time to study insects. In 1860, he gave a fascinating lecture on insects to the Illinois State Horticultural Society's annual convention, followed by several articles on insects in agricultural newspapers. In 1862–1866 he published a dozen articles in proceedings of the Boston Society of Natural History and the Philadelphia Entomological Society (Henshaw 1889). His “On Certain Entomological Speculations of the New England School of Naturalists” (1864a) defended Darwin's theory of evolution from the attacks by Louis Agassiz, James Dwight Dana, and their disciples.

It was one of five of his publications that he sent to Darwin, who responded appreciatively (21 October 1864, Darwin 2001–2008, XII:374–375). Darwin welcomed letters from any competent naturalists, since they usually became eager members of his de facto research institute, which he ran through the mail. He and Walsh exchanged photographs, and Walsh commented that he could still see the Darwin he knew at Cambridge behind the beard and despite the balding head (1 March 1865, Darwin 2001–2008, XIII:69). Darwin also sent Walsh a copy of his impressive paper, “On the Sexual Relations of the Three Forms of Lythrum salicaria” (1865), and although Lythrum is a plant, it addressed the same topic as Walsh's “On Dimorphism in the Hymenopterous genus Cynips” (1864b), which he had sent Darwin. Walsh also thanked Darwin for having John Lubbock review Walsh's Cynipidos theory in the Natural History Review (January 1865:138–141). Walsh had “no possible chance out in this uncivilized region to get a sight of Bates's Paper on mimetic Lepidoptera,” and asked Darwin to request that Bates send him that or other papers. Darwin replied (27 March 1865 [2001–2008, XIII:95–96]) that he had passed on Walsh's request to Henry Walter Bates and also thanked Walsh for his paper on willow galls (Walsh 1864c), which interested Darwin. He also wanted a copy of any paper Walsh published on wingless insects. In response to Darwin's question about the evolution of gall-making, Walsh replied (29 May1865 [Darwin 2001–2008, XIII:155–156):

My idea as to the mode in which Cecidomyia acquired its gall-producing poison is that originally they were all without it, as are to this day the Hessian fly (C. destructor) & the Wheat-midge (Ced. Tritice). But the individuals by Variation acquired the power of secreting a minute portion of poison so as to irritate the plant slightly & cause a slight additional flow of sap, & a better nidus for the future larva, whereby they gained an advantage over their fellows & so on according to your theory.

On 9 July, Darwin wrote (2001–2008, XIII:193) that he was pleased that Walsh liked Bates' paper on mimicry (1861) and recommended that Walsh also read Bates' Travels. Taking Darwin at his word, that he was interested in insect galls, Walsh sent him, on 12 November, a summary of his rather complex experiment to demonstrate that Cynipide insects generally do not move far on a tree from one generation to another (Darwin 2001–2008, XIII:305–306). Darwin wished him success with his experiment and thanked him for sending his paper (1865) on a potato bug, which Darwin thought “uncommonly well done” (19 December, 2001–2008, XIII:327). Leland Howard later seconded this opinion (1930:53). On 17 July 1866, Walsh commented further on the spread of insect galls (Darwin 2001–2008, XIV:244)

You mentioned some time ago [19 December 1865] the case of a foreign gall-fly having suddenly spread over England. Was it not a species that made its gall on the leaf, so that leaf & gall together might be blown great distances by the wind? I have often remarked that our “Oak-apples” are carried by the wind hundreds of yards with the living insect in them; but the species that make their galls in the twig, so that they are part & parcel of the twig itself, infest the same tree year after year, without spreading, except very slowly indeed sometimes, to adjoining trees.

Walsh thought Darwin would be pleased with the second part of his willow gall paper (1866b) because it showed that “Gall-making insects must have originated according to your theory.” Walsh became editor of the second (and last) volume of the Practical Entomologist 1867 and sent Darwin a copy. Darwin gladly read it, but he expressed skepticism of Walsh's claim in it about male Lucanidae (9 August 1867 [Darwin 2001–2008, XV:345])

…are you sure that the Lucanidae use their great jaws to hold the females in copulation; I always thought that they used them in fighting with other males, & I am nearly sure that this is the case.

Walsh did not reply promptly, but the skepticism expressed above did not shake Darwin's confidence in Walsh's reliability, for on 14 February 1868 he sent Walsh a series of questions, which he hoped Walsh might answer, on sexual selection in insects (2001–2008, XVI:114). Three days later, he wrote an apology for not having looked though Walsh's papers he already had before writing that letter, but he also added more questions that had arisen since 14 February (17 February [Darwin 2001–2008:128]). Walsh replied on 25 March to all three of Darwin's letters (Darwin 2001–2008, XVI:313–317). As for the jaws of male Lucanidae, Walsh did not doubt that they “may be used for pugnacious as well as amatory purposes …” Darwin was quite pleased with Walsh's detailed letter: “it really is a mine of wealth to me” (13 April [Darwin 2001–2008, XVI:399]). With such encouragement, Walsh sent still more details on 1 May (Darwin 2001–2008:460–462).

In 1867 Walsh became State Entomologist for Illinois (Decker 1958:105–108, 113–115), and he published a First Annual Report on the Noxious Insects of Illinois 1867. Commenting on it later, Howard (1930:53) stated: “Few people can write the way Walsh wrote, and he had a way of stressing important points that was masterly.” Howard said that Walsh's bibliography ran to 385 titles. After his accidental death in 1869, his collection of 10,000 insects reached the Chicago Academy of Science in time to be destroyed by Chicago's 1871 fire.

Charles Valentine Riley (1843–1895) seemed more ambitious than his older countryman, Walsh (Howard 1930:53–57, Essig 1931:741–745, Mallis 1971:69–79, Ordish 1972:28–32, Sorensen 1995:75–77, Sawyer 1996:6–15, Smith and Smith 1996, Wassong 1997, Smith 1999). Mallis called him “one of the greatest and most controversial figures in the history of American entomology.” He had a passion for collecting and drawing insects from an early age. Although he was illegitimate and his father died in debtor's prison, he was nevertheless educated in London, Dieppe, France, and Bonn, Germany, all before age 17, when he came to America. He worked on a northern Illinois farm for three years, and studied insects there. By August 1863, the Colorado potato beetle Leptinotarsa decemlineata (Say) had reached Illinois (via railroads), and Riley sent a description, with illustration of its life stages, to the Prairie Farmer, the foremost American farm journal, published in Chicago. He soon became its entomological editor and corresponded with other entomologists. In 1868 he became the state entomologist for Missouri, having been recommended to the legislature by Walsh. Riley remained there nine years, 1868–1877, and issued nine important annual reports that, like Fitch's, appealed to both farmers and entomologists (Henshaw 1889). Also in 1868, Walsh and Riley founded the American Entomologist, and after Walsh's death, Riley became sole editor. Riley wrote a book about the Colorado potato beetle (1876), which contained his illustration of its life stages (Fig. 10). It was reprinted in England (1877) to reassure Britons it was unlikely to arrive with American potatoes. However, suppliers to insect collectors were breeding it in England until a law stopped them (Ordish 1972:3–4). This beetle is now thought to be a native of Mexico (Casagrande 1987:142). It had already become a serious pest in Kansas, Nebraska, and Iowa before it reached Illinois via the transcontinental railroad in 1865 (Allen 2008:60); it reached the Atlantic in 1874. Some farmers got their chickens to pick the beetles from the potato plants, but most farmers resorted to picking them off by hand. There were many varieties of potatoes, and some were found to be less vulnerable than others. In his article on this beetle, Walsh (1866a) described a newly invented horse-drawn potato beetle collector.

Walsh (1866a) identified stink bugs and ladybird beetles as enemies of the potato beetle. Riley (1869), who had become State Entomologist for Missouri in 1868, described a parasitic fly, Lydella doryphorae (now Doryphoraphaga doryphorae), which destroyed 10% of the second generation of potato beetles and 50% of the third generation in his garden. Yet he attributed a decline in Iowa potato beetles to depredations of ladybird beetles. Riley also described four species of ladybird beetles, four bugs, four ground beetles, and a blister beetle that occasionally preyed on the potato beetle. His list of enemies later climbed to 22, but D. doryphorae was most important. The Canadian entomologist Charles James Steward (1838–1932), who would later become professor of entomology and zoology at the Ontario Agricultural College, reviewed the 22 enemies and concluded they could not adequately control the potato beetle (1872). By 1872 Riley was recommending the use of Paris green, an arsenic paint, which became the first widely used insecticide, against the potato beetle (Casagrande 1987:144).

Reports of the Colorado potato beetle depredations were soon eclipsed by news of the even greater devastation of crops by the Rocky Mountain locust Melanoplus spretus. Walsh had officially named and described it in 1866, though the specific name had been used for it earlier (Lockwood 2004:28–29). There had been occasional local grasshopper plagues reported since the mid 1700s in the East and in California (Schlebecker 1953:86), but grasshoppers, unlike locusts, do not migrate long distances. Yet, this locust differed from potato beetles in not migrating east as farmers migrated west. It lived in an arid environment, and it only became a problem as farmers settled prairies and plains. The Selkirk Settlement in the Red River Valley, now in Manitoba, suffered locust attacks of crops in 1818, 1819, and 1822 (Riegert 1980:24–26). The Mormons in Utah experienced locust plagues in 1854, 1855, and in the 1860s (Lockwood 2004:44–45). In 1856, locusts were eating crops in Canada's Manitoba, Saskatchewan, and Alberta, and soon also in Minnesota (Riegert 1980:27, Sorensen 1995:129). Wild birds, chickens, and turkeys would eat locusts, but if they gorged themselves, it could be fatal. Farmers learned that peas, beans, and corn were less vulnerable than wheat to locust attack (Lockwood 2004:55–58).

Locust plagues of the 1870s were the most dramatic event in the history of American entomology (Sorensen 1995:127–130). By 1873 locust problems were so serious that there were calls for government intervention. Entomologist John Lawrence LeConte (1825–1883), speaking before the American Association for the Advancement of Science, urged reorganization of the USDA to use science more effectively to assist farmers (Essig 1931:680–685, Mallis 1971:242–248, Sorensen 1995:10–11). Governors of Iowa and Minnesota asked for federal assistance, and in 1874, so did governors of Kansas and Nebraska. On 3 March 1876, Riley published a letter in the St. Louis Globe-Democrat advocating a commission to study injurious insects (Dexter 1979:28). The Minnesota governor convened a meeting of Midwestern governors in Omaha on 26 October 1876, and they were impressed by a talk Riley delivered (Lockwood 2004:86). He and LeConte also appeared before agricultural committees of the U.S. House and Senate in 1877 and proposed formation of a entomological commission to deal with the locust threat. Congress did so, and Riley became its head, with Cyrus Thomas and Alpheus Packard as the other commissioners on the U.S. Entomological Commission.

Cyrus Thomas (1825–1910) never attended college, though he became both a lawyer and a Lutheran minister (Howard 1930:14–15, Essig 1931:770–772, Mallis 1971:50–52). He served as botanist and entomologist for Hayden's U.S. Geological Survey of the Territories, 1869–1873, taught at the Illinois State Normal School, 1874–1876, and served as State Entomologist for Illinois from 1875 to 1882, when he became entomologist at the Smithsonian Institution (Mills 1958:93–94). He was appointed to the Commission at the insistence of his brother-in-law, Illinois Senator John A. Logan (Sorensen 1995:134–135).

The Entomological Commission was rather successful. Ralph Dexter (1979:30–32) has summarized its five reports, 1878–1890, the briefest of which ran to 396 pages, and the longest to 957 pages, with illustrations and maps (see Fig. 17). Instead of the commissioners fighting over power and prestige or how to conduct their work, they divided the western territory into three sections and each used his own judgment in the work for his section. Although lacking the term, they pioneered the use of “integrated pest management,” which was only named a century later, when it became clear that insecticides alone would not provide long-term insect control (Lockwood 2004:110–111). The Rocky Mountain locust was their main focus, but other pests were also investigated, and commissioners delegated research to other entomologists as needed. They explained the differences between this locust and those of Eurasia and Africa and from the other grasshoppers of America. The reports also described its biology, distribution, migration, and enemies. The breeding grounds were 300,000 square miles of high plateau east of the Rocky Mountains in Colorado, Wyoming, Montana, Idaho, western North Dakota, and in Canada. Hot weather affected buildup of swarms, and wind controlled their direction. Entomologists and farmers could use this knowledge to vanquish their foe. There was even a discussion of the brain, histology, and what we call its ecology. The last major locust invasion in Canada was in 1875 and in the United States in 1877. Norman Criddle collected the last living specimens, male and female, in Manitoba, Canada on 19 July 1902 (Lockwood 2004:128). The reason for the extinction may have been a change in agricultural practices, perhaps influenced by Commission advice, or perhaps the disappearance of buffalo from the breeding range, or both (Sorensen 1995:147). The commission was a helpful stepping stone for Riley and Packard to better positions (Palladino 1996:25), and could have been so for Thomas as well, had he chosen to continue working in entomology.

In 1878, when Glover resigned due to ill health, Riley was appointed chief of USDA's Division of Entomology. He persuaded Congress to increase funding for the Commission, but Commissioner of Agriculture William Le Duc disapproved of Riley going directly to Congress rather than going through Le Duc's office, and fired Riley. Le Duc replaced Riley with Cornell professor John Henry Comstock, but after two years, Riley returned under a new presidential administration and remained there until 1894. It was natural for him to bring to his USDA division the problem-oriented approach that he used at the Entomology Commission. In doing so, his division became the prototype for other science bureaus (Dupree 1957:161–162, Sorensen 1995:140). The Entomology Division's budget steadily increased under Riley, though it rarely exceeded $30,000 (Weber 1930:11). In 1885 the Division of Entomology established a branch of economic ornithology and mammalogy, which in 1886 became a separate USDA division.

During the 1600s and 1700s, some European immigrants brought grapevines to America and attempted to establish vineyards, but all failed except in California. During the 1800s, American farmers began successfully domesticating native grapes (Ordish 1972:19–23). No one knew why European vines failed and American vines flourished, and no one reported aphids sucking on the roots. In 1867, the French agricultural press began reporting diseased grapevines. This happened after American grapevines had been imported for several years. The mayor of St-Rémy asked the Société d'Agriculture de l'Hérault to investigate. The society appointed a three-man commission in July 1868 that was more successful than Prime Minister Peel's commission established in 1845 to investigate the Irish potato blight (Egerton 2012:312). This vine commission included viticultural scientist Félix Sahut and Montpellier Professor of Medicine and Pharmacy Jules-Émile Planchon (1823–1888). Sahut discovered aphids on vine roots and showed them to Planchon, who named the aphid Rhizaphis vastatrix (Ordish 1972:35–38, Paul 1996:19). In 1869, Planchon and his brother-in-law, J. Lichtenstein, decided that this aphid seemed to be the same as Fitch's leaf aphid Pemphigus vitifolii, and Riley went to France in 1869 and confirmed their identity. Ordish (1972:41) stated that Fitch's specific name should have been retained, and some entomologists do use it, but that vastatrix was too widely used to be eliminated. But entomologist Victor Antoine Signoret (1816–1889) thought the aphid might attack the roots because the vine was weakened by disease, and he transferred vastatrix to the Phylloxera genus (Ordish 1972:43–44). However, Planchon stood his ground, and another commission in 1869 agreed with him. In 1873, Planchon went to America and Riley took him on a five-week tour of vineyards and entomologists, and he collected vines that resisted the pest, though they turned out not suited to the French soil; another Frenchman came over to get resistant stocks suited to French soils (Ordish 106–107). Planchon and Riley introduced into France an American mite, Tyroglyphaus phylloxerae, which preys on Phylloxera, and it became established, but without significant impact on Phylloxera populations (Howard and Fisk 1911:24). To save their own grape varieties, Europeans grafted them onto stems of aphid-resistant American varieties (Paul 1996, Campbell 2004:xix, Gale 2011).

When Walsh wrote to Darwin on 1 May 1868, he passed along this information (Kritsky 1995:92)

The State of Missouri has just appointed a young friend of mine, C. V. Riley of Chicago, State Entomologist with a salary of $2,500 per annum. He is comparatively only a beginner in the science, but he is a hard working intelligent young fellow, understands German which I am ashamed to say I do not…

Walsh enclosed both his own and Riley's reprints. Darwin responded on 21 September, sending his thanks and his photograph for Riley. After Walsh's death in 1869, Darwin wrote to Riley on 1 June 1871 (Kritsky 1995:92)

I received some little time ago your report on noxious insects, and have now read the whole with the greatest interest. There are a vast number of facts, and generalizations of value to me, and I am struck with admiration at your powers of observation.

The discussion on mimetic insects seems particularly good and original.

In response to such praise, Riley sent Darwin his state reports for the next six years, and the marginalia in Darwin's copies show that he read them closely and cited Riley's observations in his own books (Kritsky 1995:92–94). Riley visited Darwin at Down House in 1871 and 1875, and after Darwin's death in 1882, Riley wrote an appreciative essay on Darwin's contributions to entomology (Riley 1882).

In 1887 the Fruit Growers' Convention met at Riverside, California and invited Riley to discuss the severe infestation of the cottony-cushion scale Icerya purchasi (Caltagirone and Doutt 1989:3). There had been previous attempts to import predators or parasites of pest insects (Howard and Fiske 1911:18–22), and Riley told the growers that the cottony-cushion scale was an accidental import from Australia, and that someone should go there to find its predators or parasites, but that Congress had limited his department's investigations to the United States. He suggested they discuss this with their Congressmen, which they did. This led to the discovery that the State Department was going to send a representative to an international exposition in Melbourne, and one of the entomologists from USDA, Albert Koebele (1852–1924) was sent as the U.S. representative (Howard 1930:500–503, Essig 1931:274–279, 673–680, Sawyer 1996:11–16). In Australia, Koebele discovered that the vedalia beetle Rololia cardinalis preys on I. purchasi, and that a fly Cryptochetum iceryae parasitizes it. Koebele sent 129 of these beetles and 12,000 flies to an agent in California, where the beetles were placed on a screened-in orange tree infested with I. purchasi. The vedalia beetle was extremely successful in destroying the I. purchasi and also in reproducing, so that there were soon beetles that could be distributed throughout citrus-growing regions. In areas too cool for vedalia beetles, C. iceryae flourished and checked I. purchasi populations (Caltagirone and Dott 1989:6). This first significant venture into biological control was probably the most successful. Although I. purchasi is still in the orchards, so is Rololia cardinalis, which continues to keep its numbers low (Waldbauer 2009:8).

Introducing vedalia beetles was such a spectacular success story that biological control became very popular (Sawyer 1996, Allen 2008:70–72), and led Koebele into a career of biological control in Hawaii (Palladino 1996:77–78). However, Riley and Koebele had beginner's luck: many introductions have not worked and some have backfired (Krebs 1985:410–412, Simberloff and Stiling 1996, Davis 2009:153–154). In 1893, Riley was President of the Washington Entomological Society, and his presidential address was a survey of parasitism, including the use of parasites to control pests (Riley 1893). In 1894, Riley and an assistant, Henry Hubbard, traveled to Montserrat in the Caribbean in search of parasites of citrus pests, and when they returned, Riley was chastised for violating Department travel restrictions. He resigned in disgust and moved over to the Smithsonian Institution, only to die in a bicycle accident on 14 September 1895 (Smith and Smith 1996:237).

By the late 1800s, America led the world in applied entomology, and “pure” entomology also flourished. Virtually all entomologists were either in government service or in universities. In 1889, they founded the American Association of Economic Entomologists (Howard 1930:109–114). Riley had suggested doing so in the January issue of Insect Life, and when Canada's Entomologist, James Fletcher, visited Washington, D.C., in July, he and Howard drafted a constitution, and it was organized in Toronto in August. The Association founded the Journal of Economic Entomology in 1907. This association lasted until 1953, when it merged with the Entomological Society of America, founded in 1906. The 1890s were quite eventful for American entomology, because of newly discovered insect pests on the one hand—gypsy moths in Massachusetts in 1889, San Jose scales in the East in 1893, and cotton boll weevils in Texas in 1894 (Howard 1930:115–134)—and on the other hand, the establishment of agricultural experiment stations with entomologists who could study these pests (Palladino 1996:27–29).

French naturalist-astronomer Leopold Trouvelot, living in Medford, Massachusetts, imported eggs of gypsy moths Porthetria (now Lymantria) dispar in 1868 or 1869, while investigating raising American silkworms, despite the damage that gypsy moths did in Eurasia (Forbush and Fernald 1896:3–7, 273–284, 1977). Some eggs or larvae blew out the window (no screens), which Trouvelot promptly reported, but no damage was reported until 1889 (when Trouvelot was back in Paris). In June 1889, gypsy moths began defoliating trees in Medford and a specimen was taken to the secretary of the Massachusetts State Board of Agriculture for identification, who was unable to identify it and sent it to the Hatch Experiment Station at Amherst (Forbush and Fernald 1896:32–44). In December, the president of the Massachusetts Agricultural College wrote to the governor suggestions for extermination, which he communicated to the legislature in 1890, on which the legislature acted, authorizing a salaried commission. Edward H. Forbush was appointed to head the eradication effort, under the guidance of State Entomologist and Professor of Zoology at the Massachusetts Agricultural College Charles H. Fernald (1838–1921). In 1896 they co-authored a 600-page report on the life history and eradication of gypsy moths which Howard (1930:119) praised as a model for others. It included an extended discussion of birds and insects that preyed on the moths (Forbush and Frenald 1896:203–243, 375–406). However, gypsy moths are now established in northeastern USA and in southeastern Canada (Elkinton and Liebhold 1990).

Comstock discovered and bestowed the scientific name on the San Jose scale (Aspidiotus [now Quadraspidiotus] perniciosus) in the Santa Clara Valley in California in summer 1880, where it infested fruit trees (Howard 1930:119–124,). It spread rapidly along the Pacific coast. Its origin was long unknown, but Charles L. Marlatt eventually traced it to China, and it entered America on imported fruit trees. In August 1893, specimens were sent from Charlottesville, Virginia to the USDA, where Howard identified it and raised an alarm. Within six years, however, it had spread throughout the eastern and midwestern states (Croker 2001:112–114). Entomologists and orchardists developed a lime–sulfur and mineral oil supressant that controlled, but did not eliminate it. It remains a serious pest (Pedigo 1989:542).

In 1843, someone collected a weevil in Vera Cruse, Mexico, which a Swedish entomologist named Anthonomus grandis (Howard 1930:124–132). In 1871, a German entomologist discovered it in Cuba, but it did not attract attention until botanist Edward Palmer collected it at Monclova, Mexico, where the damage to cotton was so great that farmers switched to other crops. Palmer sent a specimen and letter to the USDA, where no one could identify it, and USDA sent it to Paris, where A. Sallé identified it. By 1894, the boll weevil was destroying the cotton crop in Corpus Christi, Texas, and USDA sent agent C. H. Tyler Townsend to investigate. He traveled from 15 November to 15 December through southern Texas and adjacent Mexico and submitted an alarming report. In subsequent years it spread steadily north and east. In 1901, USDA put Walter D. Hunter (1875–1925) in charge of boll weevil studies and eradication, to which he devoted the remainder of his life. One remedy was “use of an early-maturing variety of cotton and a forcing of the crop, bringing about an early harvest, and the destruction of all cotton standing in the field by the end of October” (Howard 1930:127), but alternation of crops and sprays proved more popular. Nevertheless, “Lint and cottonseed losses between 1909 and 1949 averaged $203 million annually” (Pedigo 1989:330).

Stephen A. Forbes (1844–1930), from Illinois, was only a year younger than Riley, yet his career was as much in the 1900s as 1800s, and he also worked on both sides of the academic–government divide (Howard 1931, Mallis 1971:55–60, Winsor 1975, Lovely 1995, Croker 2001). He studied at the Rush Medical College in Chicago and at the Illinois Normal University, Normal, (just north of Bloomington), and in 1871 became curator of the Natural History Museum of Normal University; he became Professor of Zoology in 1875. He was probably first to make systematic studies on wild animal diets, beginning with fishes in 1878 and birds in 1880 (Forbes 1878. 1880a, Lovely 1995:176–177, 388–389). There are both fish and bird species that eat insects, and it was a natural progression to go on to study the diets of insect species that also eat other insects (Forbes 1880b, Lovely 1995:188). He also determined which Illinois insects were harmful or beneficial to agriculture and horticulture (Forbes 1880, Lovely 1995:188–223). In 1877 the museum was moved to Springfield and he persuaded the board of education and the state legislature to turn the Normal Museum building into a state biology laboratory to investigate Illinois natural history (Kohler 2006:102–103). It became a model for other states to follow (Mills et al. 1958, 1977). In 1882 Forbes became state entomologist, without relinquishing his administration of the state laboratory, and he published Illinois entomology reports from 1883 to 1916 (Howard 1931:31–51). In 1884, the University of Indiana awarded him a Ph.D. based on his publications, and in 1885 he became Professor of Zoology and Entomology at the University of Illinois in Champaign, without relinquishing other positions. Forbes contributions were widely appreciated (Howard 1931:17), and he was the second president of Association of Economic Entomologists.

Forbes not only investigated the foods of a wide variety of species, he also discussed the ecological implications of his findings (while not yet using Haeckel's term, “oecologie”). “On some interactions of organisms” was his first general ecological essay, and he began by emphasizing the interrelatedness of species that live together (1880a):

The serious modification of any group, either in numbers, habits, or distribution, must modify, considerably, various other groups; and each of these must transmit the change in turn, or initiate some other form of change, the disturbance thus propagating itself in a far extending circle.

A specific example: He found that Carabidae beetles eat 57% animal matter and 43% of plant matter. Since these beetles prefer animal food, if a prey species has a population explosion, they feed almost entirely on the numerous species until its numbers are reduced to its usual numbers (Forbes 1880b:175–176, 1883, Lovely 1995:189–192).

The success of the vedalia beetle in controlling the cottony-cushion scale impressed American entomologists, and Forbes wondered if he could duplicate that success against the cinch bug Blissus leucopterus, a serious pest of wheat, corn, and other cereals (Pedigo 1989:107). He reported in 1882 on a bacterium that infects cinch bugs.

When Riley resigned as head of entomology at USDA, he was replaced by his assistant, Leland O. Howard (1857–1950), whose illustrious career as administrator lasted until 1927 (Mallis 1971:79–86, Hatch 1972). When Howard first became Riley's assistant in November 1878, Riley asked him to compile a manual on silk culture, which Howard easily did. In his autobiography Howard says that Riley published it under his own name, which was then the custom, but a custom Howard changed when he replaced Riley (Howard 1933:28–31). During the 1900s, Howard became “probably the world's foremost entomologist” (Hatch 1972). His “Study of Insect Parasitism…of the White-marked Tussock Moth, with an Account of Their Habits and Interrelations” is interesting for ecological interactions he described (1897:5)

The beginning of its rapid increase in our cities is nearly coincident with the beginning of the remarkable multiplication of the English sparrows after their introduction into this country, and there seems little doubt that this coincidence is really a matter of cause and effect. One of the early results of the introduction of the English sparrow was the practical extermination by this bird of the cankerworms, which at that time were the principal insect enemies of our city shade trees. The removal of the cankerworms afforded room for the multiplication of the tussock moth, which, from the fact that its larva is hairy, was not eaten by the sparrows, and consequently multiplied with rapidity. Furthermore, the tussock moth must be considered as one of those species which are becoming attached to cities—which are slowly altering their habits and accommodating themselves to city environment.

Washington, D.C., had a severe outbreak of this species, Orgyia leucostigma, during summer 1895, which prompted Howard's study. In summer 1896, however, the species was scarce, and the reason seemed to be that its previous abundance had created favorable conditions for its parasites. His study of the parasites revealed 17 species of primary Hymenoptera parasites, 6 species of primary Diptera, and also 13 species of secondary and 2 species of tertiary Hymenoptera parasites (Howard 1897:52). With native parasites controlling a native pest, no human action was needed.

Conclusions

Many insect species menace human welfare, and some species are beneficial to humanity. In the 1800s they were well studied from both perspectives, and also because insects are conspicuous and fascinating. Many studies explored ecological aspects of life histories. Entomological knowledge expanded immensely, which benefited ecology.

Literature guide

Literature on the history of entomology during the 1800s is extensive, and titles listed here are not exhaustive. First, general histories. Most recent, is entomologist Jacques d'Aguilar's well-illustrated Histoire de l'entomologie 2006. Although rather brief, the chapters are chronologically organized, and it has a brief but helpful bibliography. Most ambitious is History of Entomology 1973, edited by entomologists Ray Smith, Thomas Mittler, and Carroll Smith for the Entomological Society of America. It encompasses entomological history from antiquity to mid-1900s, with seven or so topical articles that include the 1800s. Edward Smith and Janet Smith's encyclopedic article (2003) covers all entomological history in 11 pages, with about half on the 1800s. Leland Howard's History of Applied Entomology 1930 has good country-by-country coverage for the 1800s and also contains 254 portraits on 51 plates. The year after Howard's anecdotal history appeared, Edward Essig published a 1000-page History of Entomology 1931, with no subtitle to indicate its scope: about half is on California, and although the rest is general history and biographical sketches, these parts also emphasize California. Essig's “Sketch History of Entomology” (1936) is a long bibliographical essay. Herbert Osborn's Fragments of Entomological History 1937 is limited to the 1800s and early 1900s and is mostly organized topically; his Brief History of Entomology 1952 was a third the length of Essig's but more balanced in coverage, since antiquity, but also topically organized. Edward Ross' essay on “Systematic Entomology” (1955) introduces 22 historical essays by different authors on particular groups, mostly orders of insects, studied 1853–1953. Historian of biology Charlotte Sleigh's Six Legs Better: A Cultural History of Myrmecology 2007 is international in scope, covers the period 1874–1975, and emphasizes ant studies by Auguste Forel, William Wheeler, and Edward Wilson. P. Gilbert's Compendium of the biographical literature on deceased entomologists 1977 is a worldwide bibliographical guide. The Dictionary of Scientific Biography (16 volumes, 1970–1980), edited by Charles Gillispie, has biographies of some entomologists, accessed through “entomology” in its index. Baron Charles Robert Osten Sacken was one of the very few entomologists of the 1800s to write a book-length autobiography (1903–1904). He lived in Russia, the United States, and Germany. The history of taxonomy and systematics might be the least ecologically exciting aspect of the history of entomology, but it also is important. Carl Lindroth (1973) has written an excellent survey for the period 1800–1859.

There are many works on the history of entomology in particular countries. Kai Henriksen's Oversigt over Dansk Entomologis Histoire (1921–1937) is detailed, well illustrated with portraits, and has a good bibliography. The Entomological Society of London was possibly the first lasting society (if both of its starts are counted), and its members in 1933 wrote its centennial history (Neave 1933), and Berit Pedersen (2002) has edited a guide to its archives. George Ordish spent his career studying garden and agricultural pests and wrote widely on aspects of the subject, including coverage for the 1800s in The Constant Pest: a Short History of Pests and Their Control (Ordish 1976:110–173), encompassing fungi and other pests as well as insects. He also wrote John Curtis and the Pioneering of Pest Control 1974; Ordish's The Great Wine Blight 1972 concerns one species. Historian Muriel Blaisdell's well-documented Darwinism and Its Data: The Adaptive Coloration of Animals 1992 actually begins early in the 1800s. Michael Salmon, a physician and amateur lepidopterist, has written a well-illustrated Aurelian Legacy: British Butterflies and Their Collectors 2000, about both amateur and professional collectors, authors, and artists. John Clark's Bugs and the Victorians 2009 is a social history of British entomology during most of the 1800s. Two biographical dictionaries appeared in 2004 that include British entomologists of the 1800s: Lawrence Goldman, editor, Oxford Dictionary of National Biography (60 volumes) and Bernard Lightman, editor, The Dictionary of Nineteenth-Century British Scientists (4 volumes). There are two reliable histories of French entomology, 1750–1950: Lhoste (1987) and Gouillard (2004). Jean Lhoste's emphasizes the 1700s and early 1800s, and Jean Gouillard's emphasizes the late 1800s and earlier 1900s and entomological societies. Auguste-Henri Forel lived 52 years in the 1800s, which are covered in his autobiography (1931, English translation 1937).

Harry Weiss' mimeographed The Pioneer Century of American Entomology 1936 provides a narrative history (despite the title) from 1588 until 1865. Melville Hatch's A Century of Entomology in the Pacific Northwest 1949 has only 50 pages, with 9 unnumbered plates and good bibliography. Arnold Mallis' American Entomologists 1971 is a helpful biographical dictionary, with sketches organized according to groups of insects studied. Historian Conner Sorensen's Brethren of the Net: American Entomology, 1840–1880 1995 is a well-documented account of a formative period of professional development in the United States. Jeffrey Lockwood has resurrected the memory of Locust: the Devastating Rise and Mysterious Disappearance of the Insect that Shaped the American Frontier 2004. Robert Spear has described The Great Gypsy Moth War 2005, about the effort to eradicate this invasive species in Massachusetts, 1890–1901. A good number of entomologists of the 1800s are included in American National Biography, John Garraty and Mark Carnes, editors (24 volumes, 1999). Leland Howard lived 43 years during the 1800s, covered in his autobiography (1933). Paolo Palladino's Entomology, Ecology and Agriculture 1996 ostensibly covers North America for the years 1885–1985, but he found it necessary to actually begin about 1850 in order to give the background for later developments. Samuel Henshaw has compiled valuable bibliographies on Walsh and Riley (1889) and on other contributions to American economic entomology during the 1800s (1895–1896). Nathan Banks then continued that bibliography down to 1 January 1905 (1898–1905).

Canadian history is covered in two works: Glen Wiggens, editor, Centennial of Entomology in Canada, 1863–1963 1966 and Paul Riegert, From Arsenic to DDT: A History of Entomology in Western Canada 1980. Riegert's book actually begins before the use of arsenic. The Wiggens volume celebrated both the centennial of the Entomological Society of Canada and the career of Professor Emeritus Edmund M. Walker.

Literature cited

P. Acot, editor. 1998. The European origins of scientific ecology (1800–1901). Two volumes. B. P. Hamm, translator. Éditions des Archives Contemporaines, Gordon and Breach, Amsterdam, The Netherlands.
Google Scholar
D'Aguilar, J. 2006. Histoire de l'entomologie. Delachaux et Niestlé, Paris, France.
Google Scholar
Alborn, T. L. 2004. John Lubbock, first Baron Avebury (1834—1913), banker, politician, and scientific writer. Oxford Dictionary of National Biography 34: 651–652.
Google Scholar
Aldrich, M. L. 1972. Edward Hitchcock (1793–1864), geology. Dictionary of Scientific Biography 6: 437–438.
Google Scholar
Allen, W. 2008. The war on bugs. Chelsea Green Publishing, White River Junction, Vermont, USA.
Google Scholar
Banks, N. 1898–1905. Bibliography of the more important contributions to American economic entomology. Parts 6–8: The more important writings published between June 30, 1888 and January 1, 1905. U.S. Department of Agriculture Division of Entomology, Washington, D.C., USA.
Google Scholar
Barnes, J. K. 1985. Insects in the new nation: a cultural context for the emergence of American entomology. Entomological Society of America Bulletin 31: 21–30.
10.1093/besa/31.1.21
Google Scholar
Barnes, J. K. 1988. Asa Fitch and the emergence of American entomology, with an entomological bibliography and a catalog of taxonomic names and type specimens. New York State Museum Bulletin 461. Albany, New York, USA.
Google Scholar
Barnes, J. K. 1999. Asa Fitch (1809–1879), entomologist, agriculturist, and historian. American National Biography 8: 36–37.
Google Scholar
Barton, R. 2004. John Lubbock (Lord Avebury) (1834–1913). Dictionary of Nineteenth-Century British Scientists 3: 1272–78.
Google Scholar
Bellevoye, A., and J. Laurent. 1897. Les plantations de pins dans la Marne et les parasites qui les attaquent. Bulletin de la Société d'Étude des Sciences naturelles de Reims 5: 70–126. Reprinted in Egerton 1977.
Google Scholar
Belt, T. 1888. The naturalist in Nicaragua. Edition 2. (Edition 1, 1874.) Edward Bumpus, London, UK.
Google Scholar
Belt, T. 1985. The naturalist in Nicaragua. University of Chicago Press, Chicago, Illinois, USA.
Google Scholar
Bethune, C. J. S. 1872. Report of the Entomological Society of Ontario for the year 1871. Hunter and Ross, Toronto, Ontario, Canada.
Google Scholar
Blaisdell, M. L. 1982. Natural theology and nature's disguises. Journal of the History of Biology 15: 163–189.
10.1007/BF00233013
Google Scholar
Blaisdell, M. L. 1992. Darwinism and its data: the adaptive coloration of animals. Garland, New York, New York, USA.
Google Scholar
Bocking, S. 1988. Alpheus Spring Packard and cave fauna in the evolution debate. Journal of the History of Biology 21: 425–456.
10.1007/BF00144090
Web of Science®Google Scholar
Bocking, S. 1999. Alpheus Spring Packard, Jr. (1839–1905), zoologist. American National Biography 16: 883–884.
Google Scholar
Bonta, M. M. 1991. Women in the field: America's pioneering women naturalists. Texas A and M University Press, College Station, Texas, USA.
Google Scholar
Brown, W. L., Jr. 1960. Ants, acacias and browsing mammals. Ecology 41: 587–592.
10.2307/1933346
Web of Science®Google Scholar
Burkhardt, R. W., Jr. 1973. Pierre-André Latreille (1762–1833), entomology, zoology. Dictionary of Scientific Biography 8: 48–49.
Google Scholar
Caltagirone, L. E., and R. L. Doutt. 1989. The history of the vedalia beetle importation to California and its impact on the development of biological control. Annual Review of Entomology 34: 1–16.
10.1146/annurev.en.34.010189.000245
Web of Science®Google Scholar
Cameron, J. W. M. 1973. Insect pathology. Pages 285–306 in R. F. Smith, T. E. Mittler and C. N. Smith, editors. History of entomology. Annual Reviews, Palo Alto, California, USA.
Google Scholar
Cammarata, P. 1997. Townsend Glover, 1813–1883, entomologist, illustrator, federal official. Biographical Dictionary of American and Canadian Naturalists and Environmentalists 308–310.
Google Scholar
Campbell, C. 2004. Phylloxera: how wine was saved for the world. Harper Collins, New York, New York, USA.
Google Scholar
Carpenter, M. M. 1945–1953. Bibliography of biographies of entomologists. American Midland Naturalist 33: 1–116, 50: 257–348.
Web of Science®Google Scholar
Carton, Y. 2011. Entomologie, Darwin et Darwinisme. Hermann, Paris, France.
Google Scholar
Casagrande, R. A. 1987. The Colorado beetle: 125 years of mismanagement. Entomological Society of America Bulletin 33: 142–150.
10.1093/besa/33.3.142
Google Scholar
Chalmers-Hunt, J. M. 1994. Entomological bygones or historical entomological collecting equipment and associated memorabilia. Archives of Natural History: 357–378.
10.3366/anh.1994.21.3.357
Google Scholar
Clark, J. F. M. 1992. Eleanor Ormerod (1828–1901) as an economic entomologist. British Journal for the History of Science 25: 431–452.
10.1017/S0007087400029599
Web of Science®Google Scholar
Clark, J. F. M. 1997. ‘The ants were duly visited’: making sense of John Lubbock, scientific naturalism and the senses of social insects. British Journal for the History of Science 30: 151–176.
10.1017/S0007087497002999
Web of Science®Google Scholar
Clark, J. F. M. 2004a. John Curtis (1791–1862). Dictionary of Nineteenth-Century British Scientists 1: 517.
Google Scholar
Clark, J. F. M. 2004b. William Kirby (1759–1850). Dictionary of Nineteenth-Century British Scientists 3: 1147–1148.
Google Scholar
Clark, J. F. M. 2004c. Eleanor Anne Ormerod (1828–1901). Dictionary of Nineteenth-Century British Scientists 3: 1507–1511.
Google Scholar
Clark, J. F. M. 2004d. William Spence (1782–1860). Dictionary of Nineteenth-Century British Scientists 4: 1883–1884.
Google Scholar
Clark, J. F. M. 2004e. William Spence (bap.1782–1860), political economist and entomologist. Oxford Dictionary of National Biography 51: 813–814.
Google Scholar
Clark, J. F. M. 2004f. John Obadiah Westwood (1805–1893). Dictionary of Nineteenth-Century British Scientists 4: 2136–2137.
Google Scholar
Clark, J. F. M. 2006. History from the ground up: bugs, political economy, and God in Kirby and Spence's Introduction to Entomology (1815–1856). Isis 97: 28–55.
10.1086/501099
CASPubMedWeb of Science®Google Scholar
Clark, J. F. M. 2009. Bugs and the Victorians. Yale University Press, New Haven, Connecticut, USA.
Google Scholar
Cockerell, T. D. A. 1920. Biographical memoir of Alpheus Spring Packard, 1839–1905. National Academy of Sciences Biographical Memoirs 9: 181–236 + portrait.
Google Scholar
Comstock, A. B. 1953. The Comstocks of Cornell: John Henry Comstock and Anna Botsford Comstock. G. W. Herrick and R. G. Smith, editors. Comstock Publishing Associates, Ithaca, New York, USA.
Google Scholar
Comstock, J. H. 1940. An introduction to entomology. Edition 3. Comstock Publishing Associates, Ithaca, New York, USA.
Google Scholar
Croker, R. A. 2001. Stephen Forbes and the rise of American ecology. Smithsonian Institution Press, Washington, D.C., USA.
Google Scholar
Culver, D. C., T. C. Kane and D. W. Fong. 1995. Adaptation and natural selection in caves: the evolution of Gammarus minus. Harvard University Press, Cambridge, Massachusetts, USA.
10.4159/harvard.9780674419070
Google Scholar
Curtis, J. 1824–1839. British entomology: being illustrations and descriptions of the genera of insects found in Great Britain and Ireland. 193 issues or 8 volumes. London, UK.
Google Scholar
Curtis, J. 1860. Farm insects: being the natural history and economy of the insects injurious to the field crops of Great Britain and Ireland. Blackie and Son, London, UK.
Google Scholar
Darwin, C. R. 1865. On the sexual relations of the three forms of Lythrum solicaria. Linnean Society of London (Botany) Journal of the Proceedings 8: 169–196. Reprinted in Darwin 1977. 2:106–131.
10.1111/j.1095-8312.1864.tb01085.x
Google Scholar
Darwin, C. R. 1887. The life and letters. F. Darwin, editor. Two volumes. D. Appleton, New York, New York, USA. An earlier English edition is in three volumes.
Google Scholar
Darwin, C. R. 1977. The collected papers. P. H. Barrett, editor. Two volumes. University of Chicago Press, Chicago, Illinois, USA.
Google Scholar
Darwin, C. R. 2001–2008. The correspondence. Volumes 12–16, for years 1864–1868. F. Burkhardt and D. M. Porter, editors. Cambridge University Press, Cambridge, UK.
Google Scholar
Davis, M. A. 2009. Invasion biology. Oxford University Press, Oxford, UK.
PubMedWeb of Science®Google Scholar
Decker, G. C. 1958. Economic entomology. Illinois Natural History Survey Bulletin 27: 104–126.
Google Scholar
Desmond, A., and J. Moore. 1981. Darwin. Warner Books, New York, New York, USA.
Google Scholar
Dexter, R. W. 1957. The development of A. S. Packard Jr. as a naturalist and an entomologist. Brooklyn Entomological Society Bulletin 52: 57–66, 101–112.
Google Scholar
Dexter, R. W. 1979. The organization and work of the U. S. Entomological Commission (1877–1882). Melsheimer Entomological Series 26: 28–32.
Google Scholar
Dexter, R. W. 1981. Contributions of Dr. A. S. Packard, Jr. to entomology. Bios 52: 4–7.
Google Scholar
Dexter, R. W. 1997. Alpheus Spring Packard, Jr. 1839–1905, naturalist, entomologist, evolutionist, university professor. Biographical Dictionary of American and Canadian Naturalists and Environmentalists 599–600.
Google Scholar
Donisthorpe, H. St. J. K. 1924. Entomology (ants). Pages 157–167 in U. Grant Duff. 1924. The life-work of Lord Avebury (Sir John Lubbock), 1834–1933. Watts, London, UK.
Google Scholar
Doutt, R. L. 1958. Vice, virtue, and the vedalia. Bulletin of the Entomological Society of America 4: 119–123. Reprinted in Egerton 1977.
10.1093/besa/4.4.119
Google Scholar
Doutt, R. L. 1964. The historical development of biological control. Pages 21–42 in Biological control of insect pests and weeds. P. De Bach, editor. Reinhold, New York, New York, USA.
Google Scholar
Drouin, J.-M. 1992. L'image des sociétés d'insectes en France ~a l'epoque de la Révolution. Revue de Synthèse 4: 333–345.
10.1007/BF03181191
Google Scholar
Drouin, J.-M. 2005. Ants and bees between the French Revolution and the Darwinian Revolution. Ludus Vitalia 13: 3–13.
Google Scholar
Drouin, J.-M. 2012. Three philosophical approaches of entomology. In T. Uebel and A. Brenner, editors. Research networking programme: the philosophy of science in a European perspective. European Science Foundation, Strasbourg, France.
Google Scholar
Dunlap, T. R. 1978. The triumph of chemical pesticides. Environmental Review 5: 38–47.
Google Scholar
Dunlap, T. R. 1980a. Farmers, scientists, and insects. Agricultural History 54: 93–107.
Web of Science®Google Scholar
Dunlap, T. R. 1980b. The gypsy moth: a study in science and public policy. Journal of Forest History 24: 116–126.
Google Scholar
Dupuis, C. 1974. Pierre André Latreille (1762–1833): the foremost entomologist of his time. Annual Review of Entomology 19: 1–14.
10.1146/annurev.en.19.010174.000245
Web of Science®Google Scholar
Egerton, F. N., compiler. 1977. Ecological studies on insect parasitism. Arno Press, New York, New York, USA.
Google Scholar
Egerton, F. N. 2005. History of ecological sciences, part 18: John Ray and his associates: Francis Willughby and William Derham. ESA Bulletin 86: 301–313.
10.1890/0012-9623(2005)86[301:AHOTES]2.0.CO;2
Google Scholar
Egerton, F. N. 2006. History of ecological sciences, part 21: Réaumur and his history of insects. ESA Bulletin 87: 212–222.
10.1890/0012-9623(2006)87[212:AHOTES]2.0.CO;2
Google Scholar
Egerton, F. N. 2008. History of ecological sciences, part 30: invertebrate zoology and parasitology during the 1700s. ESA Bulletin 89: 407–433.
10.1890/0012-9623(2008)89[407:AHOTES]2.0.CO;2
Google Scholar
Egerton, F. N. 2009a. History of ecological sciences, part 31: studies of animal populations during the 1700s. ESA Bulletin 90: 169–193.
10.1890/0012-9623-90.2.169
Google Scholar
Egerton, F. N. 2009b. History of ecological sciences, part 33: naturalists explore North America, mid-1780s–mid-1820s. ESA Bulletin 90: 434–487.
10.1890/0012-9623-90.4.434
Google Scholar
Egerton, F. N. 2010. History of ecological sciences, part 37: Charles Darwin's voyage on the Beagle. ESA Bulletin 91: 398–431.
10.1890/0012-9623-91.4.398
Google Scholar
Egerton, F. N. 2011a. History of ecological sciences, part 38A: naturalists explore North America, mid-1820s–1850s. ESA Bulletin 92: 64–91.
10.1890/0012-9623-92.1.64
Google Scholar
Egerton, F. N. 2011b. History of ecological sciences, part 38B. naturalists explore North America, 1838–1850s. ESA Bulletin 92: 153–184.
10.1890/0012-9623-92.2.153
Google Scholar
Egerton, F. N. 2011c. History of ecological sciences, part 39. Henry David Thoreau, ecologist. ESA Bulletin 92: 251–275.
10.1890/0012-9623-92.3.251
Google Scholar
Egerton, F. N. 2011d. History of ecological sciences, part 40: Darwin's evolutionary ecology. ESA Bulletin 92: 351–374.
10.1890/0012-9623-92.4.351
Google Scholar
Egerton, F. N. 2012a. History of ecological sciences, part 41: Victorian naturalists in Amazonia—Wallace, Bates, and Spruce. ESA Bulletin 93: 35–59.
10.1890/0012-9623-93.1.35
Google Scholar
Egerton, F. N. 2012b. History of ecological sciences, part 42: Victorian Naturalists abroad: Hooker, Huxley, Wallace. ESA Bulletin 93: 125–159.
10.1890/0012-9623-93.2.125
Google Scholar
Egerton, F. N. 2012c. History of ecological sciences, part 44: phytopathology during the 1800s, ESA Bulletin 93: 303–339.
10.1890/0012-9623-93.4.303
Google Scholar
Elliott, C. A. 1997. Thaddeus William Harris, 1795–1856, librarian, entomologist. Biographical Dictionary of American and Canadian Naturalists and Environmentalists 347–349.
Google Scholar
Elliott, C. A. 1999. Thaddeus William Harris (1795–1856), librarian and entomologist. American National Biography 10: 189–191.
Google Scholar
Elliott, C. A. 2008. Thaddeus William Harris (1795–1856): nature, science, and society in the life of an American naturalist. Lehigh University Press, Bethlehem, Pennsylvania, USA.
Google Scholar
Endersby, J. 2004. Edward Newman (1801–1876). Dictionary of Nineteenth-Century British Scientists 3: 1471–1472.
Google Scholar
Erickson, R. F. 2004. Adrian Hardy Haworth (1768–1833). Dictionary of Nineteenth-Century British Scientists 2: 918–919.
Google Scholar
Essig, E. O. 1931. A history of entomology. Macmillan, New York, New York, USA. 1965. Hafner, New York, New York, USA.
Google Scholar
Essig, E. O. 1936. A sketch history of entomology. Osiris 2: 80–123.
10.1086/368457
Google Scholar
Evans, H. E. 1975. The Comstock heritage. Pages xvii–xxv in D. Pimentel, editor. Insects, science, and society. Academic Press, New York, New York, USA.
10.1016/B978-0-12-556550-9.50009-1
Google Scholar
Evans, M. A., and H. E. Evans. 1970. William Morton Wheeler, biologist. Harvard University Press, Cambridge, Massachusetts, USA.
10.4159/harvard.9780674333383
Google Scholar
Fabre, A. 1921. The life of Jean Henri Fabre: the entomologist, 1823–1910. B. Miall translator. Dodd, Mead, New York, New York, USA.
Google Scholar
Fabre, J.-H. 1879–1907. Souvenirs entomologiques: études sur l'instinct et les moeurs des insectes. 10 volumes. Delagrave, Paris, France.
Google Scholar
Fabre, J.-H. 1918. The wonders of instinct: chapters in the psychology of insects. A. Teixeira de Mattos and B. Miall, translators. T. Fisher Unwin, London, UK.
Google Scholar
Fabre, J.-H. 1949. The insect world. A. Teixeira de Mattos, translator. E. W. Teale, editor. Dodd, Mead, New York, New York, USA.
Google Scholar
Fitch, A. 1845. Insects injurious to vegetation. No. 2. Insects of the genus Cecidomyia, including the Hessian fly and wheat fly. American Quarterly Journal of Agriculture and Science 1: 255–269.
Google Scholar
Fitch, A. 1846. The Hessian fly: its history, character, transformation, and habits. New York State Agricultural Society Transactions 6: 316–372.
Google Scholar
Foote, Y. 2004a. Edward Newman [pseud. Rusticus] (1801–1876), naturalist. Oxford Dictionary of National Biography 40: 629–630.
Google Scholar
Foote, Y. 2004b. John Obadiah Westwood (1805–1893), entomologist and paleographer. Oxford Dictionary of National Biography 58: 318–319.
Google Scholar
Forbes, S. A. 1877, 1879. The food of birds. Illinois Horticultural Society Transactions 10: 37–44, 12: 140–145.
Google Scholar
Forbes, S. A. 1878. The food of Illinois fishes. Illinois Laboratory of Natural History Bulletin 1: 71–89. Reprinted in Forbes 1977.
Google Scholar
Forbes, S. A. 1880a. On some interactions of organisms. Illinois Laboratory of Natural History Bulletin 1: 3–17. Reprinted in Forbes 1977.
Google Scholar
Forbes, S. A. 1880b. Notes on insectivorous Coleoptera. Illinois Laboratory of Natural History Bulletin 1: 153–160. Reprinted in Forbes 1977.
Google Scholar
Forbes, S. A. 1882. Bacterium a parasite of the cinch bug. American Naturalist 16: 824–825.
PubMedWeb of Science®Google Scholar
Forbes, S. A. 1883. The food relations of the Carabidae and Coccinellidae. Illinois Laboratory of Natural History Bulletin 1: 33–64. Reprinted in Forbes 1977.
PubMedGoogle Scholar
Forbes, S. A. 1895. On contagious disease in the cinch-bug (Blissus leucopoterus, Say). Department of Agriculture of the State of Illinois Transactions 24: 16–150.
PubMedGoogle Scholar
Forbes, S. A. 1977. Ecological investigations of Stephen Alfred Forbes. F.N. Egerton, compiler. Arno Press, New York, New York, USA.
Google Scholar
Forbush, E. H. and C. H. Fernald. 1896. The gypsy moth Porthetria dispar (Linn.): a report of the work of destroying the insect in the Commonwealth of Massachusetts, together with an account of its history and habits both in Massachusetts and Europe. Wright and Potter, Boston, Massachusetts, USA. 1977. Arno Press, New York, New York, USA.
Google Scholar
Forel, A.-H. 1928. The social world of ants, compared with that of man. C. K. Ogden, translator. Two volumes. Putnam, London, UK.
Google Scholar
Forel, A.-H. 1937. Out of my life and work. B. Miall, translator. George Allen and Unwin, London, UK. French edition, Lausanne, 1931.
Google Scholar
Freeman, R. B. 1980. British natural history books, 1495–1900: a handlist. Wm. Dawson and Sons, Folkestone, UK.
Google Scholar
Gale, G. 2011. Dying on the vine: how Phylloxera transformed wine. University of California Press, Berkeley, California, USA.
Google Scholar
Garber, J. 1999. Benjamin Dann Walsh (1808–1869), entomologist. American National Biography 22: 561–563.
Google Scholar
J. A. Garraty, and M. C. Carnes , editors. American national biography. 24 volumes. Oxford University Press, New York, New York, USA.
Google Scholar
Gerardi, M. H., and J. K. Grimm. 1979. The history, biology, damage, and control of the gypsy moth Porthetria dispar (L.). Fairleigh Dickinson University Press, Rutherford, New Jersey, USA.
Web of Science®Google Scholar
Gilbert, P. 1977. A compendium of the biographical literature on deceased entomologists. British Museum (Natural History), London, UK.
Google Scholar
Gilbert, P. 2004. Edward Donovan (1768–1837), natural historian and artist. Oxford Dictionary of National Biography 16: 552.
Google Scholar
C. C. Gillispie, editor. 1970–1980. Dictionary of scientific biography. 16 volumes. Charles Scribner's Sons, New York, New York, USA.
Google Scholar
Glover, T. 1867. Report of the entomologist. Pages 27–44 in Report of the commissioner of agriculture for the year 1866. Government Printing Office, Washington, D.C., USA.
Google Scholar
Glover, T. 1868. Report of the entomologist. Pages 58–76 in Report of the Commissioner of Agriculture for the year 1867. Government Printing Office, Washington, D.C., USA.
Google Scholar
Glover, T. 1878. Report of the entomologist and curator of the museum. Pages 89–148 + 5 plates in Report of the Commissioner of Agriculture for the year 1877. Government Printing Office, Washington, D.C., USA.
Google Scholar
L. Goldman, editor. 2004. Oxford Dictionary of National Biography. 60 volumes. Oxford University Press, Oxford, UK.
Google Scholar
Gouillard, J. 2004. Histoire des entomologists français (1750–1950). Société nouvelle des Éditions Boubée, Paris, France.
Google Scholar
Grant Duff, U. 1924. The life-work of Lord Avebury (Sir John Lubbock) 1834–1913. Watts, London, UK.
Google Scholar
Guralnick, S. M. 1972. Geology and religion before Darwin: the case of Edward Hitchcock (1793–1864). Isis 63: 529–543.
10.1086/351001
Web of Science®Google Scholar
Hagen, H. A. 1884. The Melsheimer family and the Melsheimer collection. Canadian Entomologist 16: 191–197.
10.4039/Ent16191-10
Google Scholar
Harris, T. W. 1841. A report on the insects of Massachusetts injurious to vegetation. Folsom, Wells, and Thurston, Cambridge, Massachusetts, USA. 1970. Arno Press, New York, New York, USA.
Google Scholar
Hatch, M. H. 1949. A century of entomology in the Pacific Northwest. University of Washington Press, Seattle, Washington, USA.
Google Scholar
Hatch, M. H. 1972. Leland Ossian Howard (1857–1950), applied entomology. Dictionary of Scientific Biography 6: 524–525.
Web of Science®Google Scholar
Haworth, A. H. 1807. Review of the rise and progress of the science of entomology in Great Britain. Entomological Society of London Transactions 1: 1–69.
Google Scholar
Hendrickson, W. B. 1961. Nineteenth-century state geological surveys: early government support of science. Isis 52: 357–371.
10.1086/349475
Web of Science®Google Scholar
Henshaw, S., compiler. 1889. Bibliography of the more important contributions to American economic entomology. Parts 1–3: the more important writings of Benjamin Dana Walsh and Charles Valentine Riley. U.S. Department of Agriculture Division of Entomology, Washington, D.C., USA.
Google Scholar
Henshaw, S., compiler. 1895–1896. Bibliography of the more important contributions to American economic entomology. Parts 4–5. Of government and state entomologists, and of other contributions. Part 4: A–K. Part 5: L–Z. U.S. Department of Agriculture Division of Entomology, Washington, D.C., USA.
Google Scholar
Henson, P. M. 1993. The Comstock research school in evolutionary entomology. Osiris 8: 158–177.
10.1086/368722
Google Scholar
Henson, P. M. 1996. The Comstocks of Cornell: a marriage of interests. Pages 112–125, 311–313 in Creative couples in the sciences. H. M. Pycior, N. G. Stack, and P. G. Abiram , editors. Rutgers University Press, New Brunswick, New Jersey, USA.
Google Scholar
Henson, P. M. 1999a. Anna Botsford Comstock (1854–1930), educator and scientific illustrator. American National Biography 5: 305–306.
Google Scholar
Henson, P. M. 1999b. John Henry Comstock (1849–1931), entomologist. American National Biography 5: 312–313.
Google Scholar
Hollerbach, A. L. 1996. Of sangfroid and sphinx moths: cruelty, public relations, and the growth of entomology in England, 1800–1840. Osiris 11: 201–220.
10.1086/368760
Web of Science®Google Scholar
Hooper, J., and Y. Foote. 2004. John Curtis (1791–1862), entomologist. Oxford Dictionary of National Biography 14: 767–769.
Google Scholar
Howard, L. O. 1897. A study of insect parasitism: a consideration of the parasites of the white-marked tussock moth, with an account of their habits and interrelations, and with descriptions of new species. U. S. Department of Agriculture Division of Entomology Technical Series No. 3. Washington, D.C., USA. Reprinted in Egerton 1977.
Google Scholar
Howard, L. O. 1929. The rise of applied entomology in the United States. Agricultural History 3: 131–139.
Google Scholar
Howard, L. O. 1930. A history of applied entomology (somewhat anecdotal). Smithsonian Institution, Washington, D.C., USA.
Google Scholar
Howard, L. O. 1931. Biographical memoir of Stephen Alfred Forbes, 1844–1930. With bibliography compiled by H. C. Oesterling. National Academy of Sciences of the United States of America Biographical Memoirs 15: 1–54 + portrait. Reprinted in Forbes 1977.
Google Scholar
Howard, L. O. 1933. Fighting the insects: the story of an entomologist; telling of the life and experiences of the writer. Macmillan, New York, New York, USA.
Google Scholar
Howard, L. O, and W. F. Fiske. 1911. The importation into the United States of the parasites of the gypsy moth and the brown-tailed moth. U. S. Department of Agriculture Bureau of Entomology Bulletin 91, Washington D.C., USA. 1977. Arno Press, New York, New York, USA.
Google Scholar
Huber, P. 1810. Recherches sur les moeurs des fourmis indigènes. J. J. Paschoud, Paris, France.
Google Scholar
Ignaciuk, S. 1997. Philip Reese (Rhees) Uhler, 1835–1913, geologist, entomologist, librarian. Biographical Dictionary of American and Canadian Naturalists and Environmentalists 791–792.
Google Scholar
James, M. J. 1973. The new Aurelians: a centenary history of the British Entomological and Natural History Society. British Entomological and Natural History Society, London, UK.
Google Scholar
Janzen, D. H. 1966. Coevolution of mutualism between ants and acacias in Central America. Evolution 20: 249–275.
10.1111/j.1558-5646.1966.tb03364.x
CASPubMedWeb of Science®Google Scholar
Kirby, W., and W. Spence. 1815–1826. An introduction to entomology; or, elements of the natural history of insects. Four volumes. (Edition 7, 1856.) Longman, London, UK.
Google Scholar
Kirby, W., and W. Spence. 1843. An introduction to entomology; or, elements of the natural history of insects. Edition 6. Two volumes. Longman, Brown, Green, and Longmans, London, UK. Reprinted 1846 in one volume. Lea and Blanchard, Philadelphia, USA.
Google Scholar
Kohler, R. E. 2006. All creatures: naturalists, collectors, and biodiversity, 1850–1950. Princeton University Press, Princeton, New Jersey, USA.
Google Scholar
Krebs, C. J. 1985. Ecology: the experimental analysis of distribution and abundance. Edition 3. Harper and Row, New York, New York, USA.
Google Scholar
Kritsky, G. 1995. Darwin, Walsh, and Riley: the entomological link. American Entomologist 41: 89–95.
10.1093/ae/41.2.89
Google Scholar
Legros, G. V. 1913. Fabre: poet of science. B. Miall , translator. Century, New York, New York, USA. 1971. Horizon Press, New York, New York, USA.
Google Scholar
Lhoste, J. 1987. Les entomologists français (1750–1950). L'Office pour l'Information Ecoentomologique (OPIE), [Paris?], France.
Google Scholar
B. Lightman, editor. 2004. The dictionary of nineteenth-century British scientists. Four volumes. University of Chicago Press, Chicago, Illinois, USA.
Google Scholar
Lindroth, C. H. 1973. Systematics specializes between Fabricius and Darwin: 1800–1859. Pages 119–154 in R. F. Smith, T. E. Mittler and C. N. Smith , editors. History of entomology. Annual Reviews, Palo Alto, California, USA.
Google Scholar
Lockwood, J. A. 2004. Locust: the devastating rise and mysterious disappearance of the insect that shaped the American frontier. Basic Books, New York, New York, USA.
Google Scholar
Lovely, R. A. 1995. Mastering nature's harmony: Stephen Forbes and the roots of American ecology. Dissertation. University of Wisconsin-Madison, Wisconsin, USA.
Google Scholar
Lubbock, J. 1882. Ants, bees, and wasps. Kegan Paul, Trench, London, UK.
Google Scholar
Lubbock, J. 1899. Ants, bees, and wasps: a record of observations on the habits of the social Hymenoptera. Revised edition. D. Appleton, New York, New York, USA.
Google Scholar
Lubbock, J. 1929. Ants, bees, and wasps: a record of observations on the habits of the social Hymenoptera. J. G. Myers , editor. E. P. Dutton, New York, New York, USA.
Google Scholar
Lustig, A. 2004. Ants and the nature of nature in Auguste Forel, Erich Wasmann, and William Morton Wheeler. Pages 282–307 in L. Daston and F. Vidal , editors. The moral authority of nature. University of Chicago Press, Chicago, Illinois, USA.
Google Scholar
Mallis, A. 1971. American entomologists. Rutgers University Press, New Brunswick, New Jersey, USA.
Google Scholar
Marchal, P.-A. 1897. L'équilibre numérique des espèces et ses relations avec les parasites chez les insectes. Société biologique de Paris Compte Rendus 49: 129. Cited from Thompson, W. R. 1939. Biological control and the theories of the interactions of populations. Parasitology 31:299–388.
Google Scholar
Marchal, P.-A. 1907. Utilization des insects auxiliaries entomophages dans la lutte contre les insects nuisibles à l'agriculture. Institut National d'Agronomie series 2, 6: 281–354.
Google Scholar
Marchal, P.-A. 1908. The utilization of auxiliary entomophagous insects in the struggle against insects injurious to agriculture. Popular Science Monthly 72: 352–370, 406–419. Reprinted in Egerton 1977.
Google Scholar
Matthews, J. R. 1974. History of the Cambridge Entomological Club. Psyche 81: 2–37.
10.1155/1974/82714
Google Scholar
Mays, R. 2004. Moses Harris (1730–c.1788), entomologist and artist. Oxford Dictionary of National Biography 25: 456–457.
Google Scholar
Mearns, B., and R. Mearns. 2007. John Kirk Townsend: collector of Audubon's western birds and mammals. Barbara and Richard Mearns, Dumfries, UK.
Google Scholar
Merrill, G. P. 1920. Contributions to a history of American state geological and natural history surveys. Smithsonian Institution, United States National Museum Bulletin 109, Washington D.C., USA. 1978. Arno Press, New York, New York, USA.
Google Scholar
Miall, B. 1895. The natural history of aquatic insects. (Edition 2, 1903.) Macmillan, London, UK.
Google Scholar
Miller, P. July 2007. Swarm theory. National Geographic 212, number 1: 126–147.
Google Scholar
Miller, P. 2010. Smart swarm. Collins, London, UK.
Google Scholar
Mills, H. B. 1958. From 1858 to 1958. In A century of biological research. [special issue of:] Illinois Natural History Survey Bulletin 27:85–103. 1977. Arno Press, New York, New York, USA.
Google Scholar
Mills, H. B., et al. 1958. A century of biological research. Illinois Natural History Survey Bulletin 27:81–234. 1977. Arno Press, New York, New York, USA.
Google Scholar
Moore, D. T. 2004. William Kirby (1759–1850), entomologist and naturalist. Oxford Dictionary of National Biography 31: 768–769.
Google Scholar
S. A. Neave, editor. 1933. The history of the Entomological Society of London, 1833–1933. Entomological Society of London, London, UK.
Google Scholar
Needham, J. G. 1946. The lengthened shadow of a man and his wife. Scientific Monthly 62: 140–150, 219–229.
Web of Science®Google Scholar
Norland, C. E. 1974. Alpheus Spring Packard, Jr. (1839–1905), entomology. Dictionary of Scientific Biography 10: 272–274.
Google Scholar
Norwood, V. 1993. Made from this earth: American women and nature. University of North Carolina Press, Chapel Hill, North Carolina, USA.
Google Scholar
Nussac, L. de. 1907. Pierre André Latreille à Brive. Steinheil, Paris, France.
Google Scholar
Ordish, G. 1972. The great wine blight. J. M. Dent and Sons, London, UK.
Google Scholar
Ordish, G. 1974. John Curtis and the pioneering of pest control. Osprey, Reading, UK.
Web of Science®Google Scholar
Ordish, G. 1976. The constant pest: a short history of pests and their control. Charles Scribner's Sons, New York, New York, USA.
Google Scholar
Osborn, H. 1937. Fragments of entomological history, including some personal recollections of men and events. Author, Columbus, Ohio, USA.
Google Scholar
Osborn, H. 1952. A brief history of entomology: including time of Demosthenes and Aristotle to modern times with over five hundred portraits. Spahr and Glen, Columbus, Ohio, USA.
Google Scholar
Osten Sacken, C. R. 1903–1904. Record of my life work in entomology. University Press/John Wilson and Son, Cambridge, Massachusetts, USA. Part 3, 1904. (No publisher), Heidelberg, Germany.
Google Scholar
Packard, A. S., Jr. 1869. Guide to the study of insects, and a treatise on those injurious and beneficial to crops. Naturalists Book Agency, Salem, Massachusetts, USA. Trauber, London, UK.
Google Scholar
Packard, A. S., Jr. 1874. Nature's means of limiting the numbers of insects. American Naturalist 8: 270–282.
10.1086/271315
Google Scholar
Packard, A. S., Jr. 1883. The Hessian fly and its transformations. Pages 198–248 in Third Report of the United States Entomological Commission. Government Printing Office, Washington, D.C., USA.
Google Scholar
Packard, A. S., Jr. 1888. The cave fauna of North America, with remarks on the anatomy of the brain and origin of the blind species. National Academy of Sciences Memoirs 4:1–156 + 27 plates. Reprint 1977. Arno Press, New York, New York, USA.
Google Scholar
Packard, A. S., Jr. 1901. Lamarck, the founder of evolution: his life and work, with translations of his writings on organic evolution. Longmans, Green, New York, New York, USA.
10.5962/bhl.title.56432
Google Scholar
Palladino, P. 1996. Entomology, ecology and agriculture: the making of scientific careers in North America, 1885–1985. Harwood Academic Publishers, Amsterdam, The Netherlands.
Google Scholar
Pasteur, G. July 1994. Jean Henri Fabre. Scientific American 271, number 1: 74–80.
Google Scholar
Patton, M. 2007. Science, politics, and business in the work of Sir John Lubbock: a man of universal mind. Ashgate, Aldershot, UK.
Google Scholar
Paul, H. W. 1996. Science, vine, and wine. Cambridge University Press, Cambridge, UK.
Google Scholar
Pedersen, B., editor. 2002. A guide to the archives of the Royal Entomological Society. Ashgate, Aldershot, UK.
Google Scholar
Pedigo, L. P. 1989. Entomology and pest management. Macmillan, New York, New York, USA.
Google Scholar
Perris, E. 1851–1870? Histoire des insects du pin maritime. M. Buquet, Paris, France.
Google Scholar
Perris, E. 1873. Les oiseaux et les insects. Société Royale des Sciences de Liége Memoires 3. Cited from Packard, A. S., Jr. 1874. Nature's means of limiting the numbers of insects. American Naturalist 8: 270–282.
Google Scholar
Peyerimhoff, P. de. 1932. La Société entomologique de France. Pages 1–86 in Livre du Centenaire. Société entomologique de France, Paris, France.
Google Scholar
Pilet, P. E. 1972. Auguste-Henri Forel (1848–1931), medicine, neurology, entomology. Dictionary of Scientific Biography 5: 73–74.
Google Scholar
Pumphrey, R. J. 1958. The forgotten man—Sir John Lubbock, F. R. S. Royal Society of London Notes and Records 13: 49–58 + portrait.
10.1098/rsnr.1958.0005
Google Scholar
Rauschenberg, R. A. 1972. Adrian Hardy Haworth (1768–1833), entomology. Dictionary of Scientific Biography 6: 184–185.
Google Scholar
Raven, C. E. 1942. John Ray, naturalist: his life and works. (Edition 2. 1950.) Cambridge University Press, Cambridge, UK.
Google Scholar
Ray, J. 1710. Historia insectorum. A. and J. Churchill, London, UK.
Google Scholar
Rezneck, S. 1972. Asa Fitch (1809–1879), economic entomology, medicine. Dictionary of Scientific Biography 5: 11–12.
Google Scholar
Richard, G. 1973. The historical development of nineteenth and twentieth century studies on the behavior of insects. Pages 477–502 in R. F. Smith, T. E. Mittler and C. N. Smith , editors. History of entomology. Annual Reviews, Palo Alto, California, USA.
Google Scholar
Riegert, P. W. 1980. From arsenic to DDT: a history of entomology in western Canada. University of Toronto Press, Toronto, Ontario, Canada.
Google Scholar
Riley, C. V. 1869. First annual report on the noxious, beneficial, and other insects of the state of Missouri. Ellwood Kirby, Jefferson City, Missouri, USA.
Google Scholar
Riley, C. V. 1876. Potato pests: being an illustrated account of the Colorado potato-beetle and the other insect foes of the potato in North America, with suggestions for their repression and methods for their destruction. Orange Judd, New York, New York, USA.
Google Scholar
Riley, C. V. 1877. The Colorado beetle, with suggestions for its repression and methods of destruction. George Routledge and Sons, London, UK. Reprint of Riley 1876.
Google Scholar
Riley, C. V. 1882. Darwin's work in entomology. Biological Society of Washington D.C. Proceedings 1: 70–80.
Google Scholar
Riley, C. V. 1893. Parasitism in insects. Entomological Society of Washington Proceedings 2: 397–431. Partly reprinted in Acot 1998, I:285–296.
Google Scholar
Riley, C. V. 1894. Report of the entomologist. Pages 199–226 + 3 plates in Report of the Secretary of Agriculture for 1893. Government Printing Office, Washington, D.C., USA.
PubMedGoogle Scholar
Riley, C. V., A. S. Packard Jr., and C. Thomas. 1878. First annual report of the United States Entomological Commission for the year 1877 relating to the Rocky Mountain locust. Government Printing Office, Washington, D.C., USA.
Google Scholar
Riley, W. A. 1931. Erasmus Darwin and the biological control of insects. Science 73: 475–476.
10.1126/science.73.1896.475
CASPubMedGoogle Scholar
Ross, E. S. 1955. Systematic entomology: introduction. Pages 485–495 in A century of progress in the natural sciences, 1853–1953: published in celebration of the centennial of the California Academy of Sciences. California Academy of Sciences, San Francisco, California, USA. 1974. Arno Press, New York, New York, USA.
Google Scholar
Salmon, M. A. 2000. The Aurelian legacy: British butterflies and their collectors. University of California Press, Berkeley, California, USA.
Google Scholar
Sartori, M., and D. Cherix. 1983. Histoire de l'étude des insects sociaux en Suisse à travers l'oeuvre d'Auguste Forel. Sociéte Entomologique de France Bulletin 88: 66–74.
Google Scholar
Sawyer, R. C. 1996. To make a spotless orange: biological control in California. Iowa State University Press, Ames, Iowa, USA.
Google Scholar
Schlebecker, J. T. 1953. Grasshoppers in American agricultural history. Agricultural History 27: 85–93.
Google Scholar
Schmitt, S. 2010. Lacepède's syncretic contribution to the debates on natural history in France around 1800. Journal of the History of Biology 43: 429–457.
10.1007/s10739-010-9227-7
PubMedWeb of Science®Google Scholar
Schwerdtfeger, F. 1973. Forest entomology. Pages 361–386 in R. F. Smith, T. E. Mittler, and C. N. Smith , editors. History of entomology. Annual Reviews, Palo Alto, California, USA.
Google Scholar
Shteir, A. B. 2004. Pricilla Wakefield (neé Bell: 1751–1832). Dictionary of Nineteenth-Century British Scientists 4: 2079–2080.
Google Scholar
Simberloff, D., and P. Stiling. 1996. How risky is biological control? Ecology 77: 1965–1974.
10.2307/2265693
Web of Science®Google Scholar
Sleigh, C. 2007. Six legs better: a cultural history of myrmecology. Johns Hopkins University Press, Baltimore, Maryland, USA.
Google Scholar
Smith, E. H. 1976. The Comstocks and Cornell: in the people's service. Annual Review of Entomology 21: 1–25.
10.1146/annurev.en.21.010176.000245
Web of Science®Google Scholar
Smith, E. H. 1989. The Entomological Society of America: the first hundred years, 1889–1989. Entomological Society of America Bulletin 35: 10–32.
10.1093/besa/35.3.10
Google Scholar
Smith, E. H. 1990. Anna Botsford Comstock: artist, author, and teacher. American Entomologist 36: 105–113.
Google Scholar
Smith, E. H. 1999. Charles Valentine Riley (1843–1895), entomologist. American National Biography 18: 513–515.
Google Scholar
Smith, E. H., and J. R. Smith. 1996. Charles Valentine Riley: the making of the man and his achievements. American Entomologist 42: 228–238.
10.1093/ae/42.4.228
Google Scholar
Smith, E. H., and J. R. Smith. 2003. History of entomology. Pages 509–519 in V. H. Resh and R. T. Cardé , editors. Encyclopedia of insects. Academic Press, New York, New York, USA.
Google Scholar
R. F. Smith, T. E. Mittler and C. N. Smith , editors. 1973. History of entomology. Annual Reviews, Palo Alto, California, USA.
Google Scholar
Somkin, F. 1962. The contributions of Sir John Lubbock, F. R. S., to the Origin of Species: some annotations to Darwin. Royal Society of London Notes and Records 17: 183–191.
10.1098/rsnr.1962.0017
Google Scholar
Somkin, F. 1973. Sir John (Lord Avebury) Lubbock (1834–1913), entomology, anthropology, botany. Dictionary of Scientific Biography 8: 527–529.
Google Scholar
Sorensen, W. C. 1995. Brethren of the net: American entomology, 1840–1880. University of Alabama Press, Tuscaloosa, Alabama, USA.
Google Scholar
Sorensen, W. C. 1999. William Dandridge Peck (1763–1822), entomologist. American National Biography 17: 229–230.
Google Scholar
Spear, R. J. 2005. The great gypsy moth war: the history of the first campaign in Massachusetts to eradicate the gypsy moth, 1890–1901. University of Massachusetts Press, Amherst, Massachusetts, USA.
Google Scholar
Steinhaus, E. A. 1956. Microbial control—the emergence of an idea: a brief history of insect pathology through the nineteenth century. Hilgardia 26: 107–160. Reprinted in Egerton 1977.
10.3733/hilg.v26n02p107
Google Scholar
Stentsinger, R. 1973. Frederick Valentine Melsheimer: parent of American entomology. Entomological Society of Pennsylvania, University Park, Pennsylvania, USA.
Google Scholar
Sterling, K. B. 1997a. Charles James Stewart Bethune (1838–1932), entomologist, clergyman. Biographical Dictionary of American and Canadian Naturalists and Environmentalists 78–79.
Google Scholar
Sterling, K. B. 1997b. Asa Fitch, 1809–1879, entomologist. Biographical Dictionary of American and Canadian Naturalists and Environmentalists 270–271.
Google Scholar
Théodoridès, J. 1971. Jean Henri Fabre (1823–1915), entomology, natural history. Dictionary of Scientific Biography 4: 503–505.
Google Scholar
Thompson, W. R. 1939. Biological control and the theories of the interactions of populations. Parasitology 31: 299–388.
10.1017/S0031182000012865
Google Scholar
Tucker, E. A. 1920. Benjamin D. Walsh—first state entomologist in Illinois. Illinois State Historical Society Transactions 54–61.
Google Scholar
Van Riper, A. B. 2004. Thomas Belt (1832–1878). Dictionary of Nineteenth-Century British Scientists 1: 177–178.
Google Scholar
Varma, C. S. 2009. Threads that guide or ties that bind: William Kirby and the essentialism story. Journal of the History of Biology 42: 119–149.
10.1007/s10739-008-9156-x
PubMedWeb of Science®Google Scholar
Wakefield, P. 1816. An introduction to the natural history and classification of insects; in a series of letters. Darton, London, UK.
Google Scholar
Waldbauer, G. 2009. Fireflies, honey, and silk. University of California Press, Berkeley, California, USA.
Google Scholar
Walsh, B. D. 1864a. On certain entomological speculations of the New England school of naturalists. Entomological Society of Philadelphia Proceedings 3: 207–249.
Google Scholar
Walsh, B. D. 1864b. On dimorphism in the Hymenopterous genus Cynips, with an appendix containing hints for a new classification of Cynipidae and a list of Cynipidae, including descriptions of several new species inhabiting the oak-galls of Illinois. Entomological Society of Philadelphia Proceedings 3: 443–500.
Google Scholar
Walsh, B. D. 1864c. On the insects, Coleopterous, Hymenopterous, and Dipterous, inhabiting the galls of certain species of willow. Entomological Society of Philadelphia Proceedings 3: 543–664.
Google Scholar
Walsh, B. D. 1865. The new potato-bug, and its natural history. Practical Entomologist 1: 1–4.
Google Scholar
Walsh, B. D. 1866a. Imported insects—the gooseberry sawfly. Practical Entomologist 1: 117–125.
Google Scholar
Walsh, B. D. 1866b. On the insects, Coleopterous, Hymenopterous and Dipterous, inhabiting the galls of certain species of willow—part 2d and last. Entomological Society of Philadelphia Proceedings 6: 223–288.
Google Scholar
Wassong, J. 1997. Charles Valentine Riley (1843–1895), entomologist, journalist. Biographical Dictionary of American and Canadian Naturalists and Environmentalists 674–675.
Google Scholar
Weber, G. A. 1930. The Bureau of Entomology: its history, activities, and organization. Brookings Institution, Washington, D.C., USA.
Google Scholar
Weiss, H. B. 1936. The pioneer century of American entomology. Author, New Brunswick, New Jersey, USA.
Google Scholar
Wheeler, W. M. 1930. Review: A. Forel, The social world of the ants compared with that of man. Journal of Social Psychology 1: 170–177.
PubMedGoogle Scholar
G. B. Wiggens, editor. 1966. Centennial of Entomology in Canada, 1863–1963: a tribute to Edmund M. Walker. University of Toronto Press, Toronto, Ontario, Canada.
Google Scholar
Wilson, D. L. 2004. Edward Donovan (or O'Donovan: 1768–1837). Dictionary of Nineteenth-Century British Scientists 2: 604–605.
PubMedGoogle Scholar
Winsor, M. P. 1972. Stephen Alfred Forbes (1844–1930), biology. Dictionary of Scientific Biography 5: 69–71.
Google Scholar
Yavetz, I. 1987. Theory and reality in the work of Jean Henri Fabre. Rivista di Storia della Scienza 4: 111–155.
Google Scholar
Yavetz, I. 1988. Jean Henri Fabre and evolution: indifference or blind hatred? History and Philosophy of the Life Sciences 10: 3–36.
PubMedWeb of Science®Google Scholar
Yavetz, I. 1991. Theory and reality in the work of Jean-Henri Fabre. History and Philosophy of the Life Sciences 131: 33–72.
Google Scholar

Acknowledgments

I thank Drs. Jean-Marc Drouin, Muséum National d'Histoire Naturelle, Paris, Anne-Marie Drouin-Hans, Université de Bourgogne, and Professor Emeritus Sidney A. Ewing, Veterinary Parasitology, Oklahoma State University, Stillwater (all retired).

Citing Literature

Volume94, Issue1

January 2013

Pages 36-88

This article also appears in:

Frank Egerton’s History of Ecological Sciences

Journal list menu