4-16. Phylogenetic relationships among insect lineages as proposed by Martynov (1938).

4.17. The distinguished, albeit eccentric, insect morphologist Guy Chester Crampton (1881-1951) of the University of Massachusetts. Crampton's devotion to his subject was total: He surrounded himself in his small apartment with thousands of vials of specimens. Photo: Special Collections and Archives, W. E. B. DuBois Library, University of Massachusetts, Amherst.

4.18. Three giants of entomological science; from left to right, Howard E. Hinton (1913-77), noted insect biologist; Robert E. Snod-grass (1875-1962), the most famous comparative arthropod anatomist; and Sir Vincent B. Wigglesworth (1899-1994), the prominent insect physiologist. Photo taken approximately 1956. Photo: G. W. Byers, University of Kansas Natural History Museum (UKNHM).

4.19. Frank M. Carpenter (1902-94), curator of fossil insects at Harvard's Museum of Comparative Zoology for 60 years and the most influential paleoentomologist of his generation. Photo taken 1956. Photo: G. W. Byers, UKNHM.

(1897a,b), Franz Klapalek (1904, 1905); and Charles Woodworth (1906, 1907, 1930).

Picking up the reins of Tillyard and Martynov were the paleoentomologists Frank M. Carpenter (1902-94: Furth, 1994) of the United States (Figure 4.19) and Boris B. Rohdendorf (1904-77) of the Soviet Union (Figure 4.20). While Carpenter focused on the Paleozoic fauna and conservatively approached the assignment of taxa to orders, Rhodendorf prolifically described taxa, naming numerous new groups. In contrast to his predecessor Martynov, Rohdendorf produced volumes of cursory descriptions, which the current generation of Russian paleoentomologists is extensively revising. Carpenter's methodical and careful studies gained him tremendous insight into the early evolution and diversification of insects. Perhaps his greatest contributions were the synthesis of his findings on the North American Paleozoic insect fauna with those taxa described from Carboniferous and Permian deposits from elsewhere in the world. Carpenter's conservatism led him to reduce the number of extinct orders, from approximately 50 to nine. He used as minimal criteria evidence from venation and mouthparts, but also recognized large paraphyletic groups, like Protorthoptera, and was too much of a taxonomic lumper.

Meanwhile, Ryuichi Matsuda (1920-86: Ando, 1988) revived the tradition of Crampton and Snodgrass and prepared several volumes on the comparative morphology of insects, either refining previous observations or providing alternative views for those expressed by earlier authors (Matsuda, 1965, 1970, 1976). Like his predecessors, Matsuda did not provide an explicit phylogenetic outline of the insect orders, but he did contribute considerably to the understanding of homologies across the orders.

Certainly the most influential insect systematist was Willi Hennig (1913-76: Schlee, 1978) (Figures 1.23, 4.21). Aside from developing cladistic methodology, Hennig was also the first to apply this critical method of analysis to the phylogeny of insects. Hennig's major accounts in 1953 and 1969 constitute even to this day some of the greatest advances in insect phylogeny (Figure 4.22). These works were updated posthumously in an extensively annotated and edited volume (Hennig, 1981). Hennig was simultaneously a paleontologist and liberally used information from fossil insects to shape his hypotheses of primary homology, character polarity, and evolution.

4.20. Boris B. Rohdendorf (1904-77), a prolific student of Martynov whose work is now being extensively revised by a new generation of Russian paleoentomologists. Photo taken 1964. Photo: G. W. Byers, UKNHM.
4.21. Willi Hennig (1913-76), founder of phylogenetic systematics and renowned entomologist. Hennig also produced very important work in paleoentomology and biogeography. Photo: G. W. Byers, UKNHM.

Perhaps the single most important paper in systematic entomology is the 1975 review by Niels P Kristensen (Figure 4.23), who revised and built upon Hennig's work. Kris-tensen's (1975) review and updated accounts (Kristensen, 1978a, 1981, 1989a, 1991, 1995, 1999a) have provided our modern concepts for ordinal relationships of Recent insects. Shortly after the appearance of Kristensen's classic paper, H. Bruce Boudreaux, a mite specialist from Louisiana, provided his own interpretation of arthropod phylogeny, with a particular emphasis on insect relationships (Boudreaux, 1979). His compendium is another important source of characters regarding insect ordinal relationships, although his classification is not entirely supported by more recent analyses.

The number of workers interested in insect phylogeny has proliferated greatly in the last couple of decades, most of them focusing on molecular data or individual Recent orders of hexapods. Despite the fervor generated by new molecular advances, morphology and paleontology are as relevant as ever. Indeed, their importance is perhaps even more acutely felt today than they have been in preceding years, and synergistic studies are more and more the norm. One might even say that paleoentomology is experiencing a renaissance with numerous new students entering the field. Today the most

4.22. (facing page) Relationships among major insect lineages as proposed by Hennig (1953). This phylogenetic hypothesis was the first one produced for insect orders using Hennig's phylogenetic methodology. Photo: Deutsche Entomologische Institut.

prominent paleoentomologists actively researching the evolution of insect orders include Jarmila Kukalova-Peck, Alexandr P Rasnitsyn, André Nel, and Rainer Willmann. The study of fossil insects and insect phylogeny is perhaps more active than ever before, but the two are rarely married. That is why our approach here was to fold extinct insects into a framework of relationships with Recent species.

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