Brief History Of Work

The history of entomology, although an engaging topic, has been more thoroughly covered elsewhere (Essig, 1931; Smith, 1973). The following is an account of those post-Darwinian authors who have contributed most significantly to our knowledge of insect phylogeny and the fossil record.

Even though early authors did consider the various affinities among major groups, like the quinarians of the early nineteenth century, it was not until Darwin provided his theory of evolution that systematists really tackled the phylogeny of insects in a major way. The first individual to depict a phylogeny for the insects was Ernst Haeckel (1834-1919), although he did not take into account fossil forms in any critical manner. Indeed, many of the early advances in paleo-entomology had not yet taken place while he was developing his system (Haeckel, 1866, 1890, 1909) (Figure 4.11). Haeckel's treatment was, moreover, superficial as he was more interested in the higher phylogeny of all life (and later in various racist ideas on the evolution of humans). Alpheus Hyatt (1838-1902) and Jennie M. Arms (1852-1937: later Jennie M. Sheldon) provided another early evolutionary tree of insects in a small guide for teaching entomology (Hyatt and Arms, 1890). Even though they did not explicitly include fossil taxa, their phylogeny did abstractly depict the surface of the globe with the living lineages arising from the disc of the Earth and progenitors of the orders coalescing beneath the disc. Examples of other authors giving early genealogical arrangements of the orders were Schoch (1884), Brauer (1885), Emery (1886), Packard (1869, 1886), Sharp (1895, 1898), Comstock (1888), and Comstock and Comstock (1895).

In the late 1800s, two authors, Samuel H. Scudder (1837-1911) and Charles Brongniart (1859-99), dramatically expanded the study of fossil insects and, as such, discussions of insect relationships and evolution. Although previous students of fossil insects such as Oswald Heer, Ernst F. Germar, P B. Brodie, and Christoph G. A. Giebel had cursorily described taxa from several deposits, none provided a significant context in which to understand their finds. Scudder worked on deposits of various ages ranging from the Late Carboniferous to the Pleistocene of North America (Figure 4.12), while Brongniart studied the Late Carboniferous fauna of Commentry, France. These authors brought for the first time the numerous, enigmatic Paleozoic lineages to the forefront of entomology. They were the first to highlight the importance of fossil forms for understanding insect evolution (Brongniart, 1885a, 1893; Scudder, 1885, 1886). Their monographs served as the foundation of paleoentomology that would be expanded upon by Anton Handlirsch (1865-1935) (Figure 4.13). Handlirsch can truly be considered the architect of paleoentomology and provided the first critical phylogenetic study of both living and fossil insects (Handlirsch, 1903, 1904). In a monumental work, Die Fossilen Insekten, of nearly 1,500 pages, Handlirsch synthesized all that was known of fossil insects and placed them into a phylogenetic classification with the Recent fauna (Handlirsch, 1906b, 1907, 1908). Handlirsch (1925, 1937, 1939) later slightly refined his system to reflect changes in his thinking during the intervening decades.

Although not as widely recognized as Handlirsch, Karl Börner (1880-1953) published papers on the phylogeny and

4.11. The first phylogenetic diagram depicting relationships among the insects and other arthropods. Insects were divided into Masticantia and Sugentia, for the masticating (chewing) and sucking insects, respectively, which is an artificial system. From Haeckel (1866).

4.12. Fossil butterfly, Mylothrites pluto, from Florissant, Colorado, from the first major work on fossil butterflies (Scudder, 1875). Samuel H. Scudder (1837-1911) was the first entomologist to specialize on fossil insects. He published sumptuous monographs on orthopterans and North American butterflies, as well as fossils. Photo: AMNH Library.

4.12. Fossil butterfly, Mylothrites pluto, from Florissant, Colorado, from the first major work on fossil butterflies (Scudder, 1875). Samuel H. Scudder (1837-1911) was the first entomologist to specialize on fossil insects. He published sumptuous monographs on orthopterans and North American butterflies, as well as fossils. Photo: AMNH Library.

classification of Recent insects at about the same time that the former was putting forth his major contributions to the subject. It can be easily said that some of Börner's concepts on insect ordinal relationships were ahead of his time. He was a careful comparative anatomist and made such distinctions as separating the silverfish from the bristletails and positing a closer relationship between Odonata and Neoptera based on the loss of a subimaginal molt (Börner, 1904, 1909).

Two other prominent paleoentomologists who contributed significantly to the ordinal classification of insects began producing papers around the time of Handlirsch's magnum opus: Robin J. Tillyard (1881-1937: Dunbar, 1937) (Figure 4.14) of Australia and Andreas V Martynov (1879-1938: Carpenter, 1938b) of Russia (Figure 4.15). Both of them, like Handlirsch, avidly studied the living insect fauna. Martynov, who was early interested in caddisflies and crustaceans, eventually took up the study of the great fossil insect deposits of the newly formed Soviet Union. Because Martynov was a keen comparative morphologist of living insects, he was adept at interpreting prolific fossils from diverse sites such as Karatau and Soyana. Martynov devel-

4.13. Anton Handlirsch (1865-1935), Director of the Naturhistorisches Museum in Vienna and the first major paleoentomologist. Photo: Deutsche Entomologische Institut.

oped a major classification of all insects based on his studies on fossils, in which he recognized the classical division of the winged insects into paleopterous versus neopterous forms among other supraordinal groupings (Figure 4.16), and which was simultaneously recognized by G. C. Crampton (discussed later). At the same time, Tillyard was developing his own ideas on insect evolution based on studies of the Paleozoic deposits of Australia and Kansas. In particular, Tillyard was the first individual to truly recognize the significance of the Lower Permian fossils recovered from Elmo, Kansas, which he monographed. It is little known that Tillyard was also a devotee of the occult, who believed that mysticism could be used to understand the lives of long vanished organisms. It has even been recounted that he once visited the Museum of Comparative Zoology with the idea of having a local Bostonian mystic bring back to life the huge Permian griffenfly specimens preserved there so that he could observe

4.14. Entomologist and paleontologist Robin John Tillyard (18811937), who was the first major worker on Permian insects. He sought a mystic to reveal the deep past so that he could see the giant Permian griffenflies in flight, but he was also a careful and thoughtful scientist. Photo: Science News.

their behavior (e.g., Evans and Evans, 1970). Certainly other paleoentomologists were also at work at this time, such as the prolific Theodore D. A. Cockerell (1866-1948: Weber, 1965), but none were as influential on paleoentomology or neoen-tomology.

While Tillyard and Martynov toiled away on Paleozoic deposits, several neontologists were at work attempting to construct a framework of insect evolution based on the comparative morphology of living forms. Guy C. Crampton (1881-1951: Mallis, 1971) (Figure 4.17) was a gifted, albeit obsessive, morphologist who filled his tiny apartment with vials of specimens and mountains of papers for his studies on the anatomy and phylogeny of insects. He published extensively during the early 20th century on the phylogeny and classification of insect orders (e.g., Crampton, 1924, 1928, 1931, 1938). His work was focused on the modern fauna, but he did consider the fossil record as it had been documented by Handlirsch, Tillyard, and others. Crampton's detailed studies of numerous invertebrates mirrored those of the more widely recognized Robert E. Snodgrass (1875-1962) (Figure 4.18) who wrote the definitive work, even to this date, on the morphology of insects (Snodgrass, 1935). While Snodgrass contributed perhaps more than any other person to our understanding of the anatomy of insects, his studies principally focused on the larger picture of arthropods among other invertebrates, the position of hexapods, and the deciphering of homologous structures across insect orders. Thus, Snodgrass provided a wealth of data for interpreting insect evolution but he did not provide a phylogenetic synthesis of the orders. Other researchers on the modern fauna worth mentioning include Anton Krausse (Krausse, 1906; Krausse and Wolff, 1919), Charles T. Brues and Axel L. Melander (Brues and Melander, 1915, 1932), Karl L. Escherich (Escherich, 1914: building upon Prell, 1912); John B. Smith

4.15. The Russian entomologist and paleontologist Andreas V. Martynov (1879-1938). His phylogeny of insect orders (Figure 4.16) was ahead of its time, and he proposed several major lineages now recognized (cf. Figure 4.24). From Carpenter (1938b).
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