A History of the Entomology of Michael P. Parrella T Ǥ ǡ ϐϐϐ Ǥ Dzdz ǡ Ǥ Ȁ Ǥ Ȅ ϐϐϐ Ǥ ϐϐ ǡ Ǥ͕͙ǡ͔͔͔ϐ ϐ ȋ ͖͔͔͔Ȍǡ ǡ Ǥǡǡ ǡ ϐϐ ǡ Ǥ ϐ Dz dzȄ ϐǦϐ Ǥ ǡ ϐϐ American Entomologist • Volume 59, Number 1 Ǥ Mayflies: At the Beginning of Entomology and Fly Fishing ϐ ǡ ͔͔͗ Ǥ ǡThe Epic of Gilgameshǡ ͖ǡ͔͔͛ȋ ͖͔͕͕Ȍǡ ϐǤ ϐ klilǤ ȋ͖͔͔͗Ȍ ϐϐǡDzϐdz Ǥ Epic ǯǣ“Ever the river ϔǡϔϔ the water. On the face of the sun its countenance gazes, then all of a sudden nothing is there.” ϐǯǤ Epic, ǡ Ǥ EpicDz dzȋ͕͛͝͝ȌǤ the EpicϐǡMortogenesia 15 mesopotamicaȋǣȌǡ Dz dzϐ ǡ Ȁ Ǥ Dzϐǡdz ϐǦϐǤϐDzϐdz Compleat Anglerȋ͕͚͚͛Ȍǡ ǡ Dzϐ ǦǤdzȋ ǤǤȌDzϐdz ȋEphemera danica Ȍȋ ͕͖͛͝ȌǤ ϐ Dzϐdz ȋǣȀȀǤ Ȁ͖͗͝Ȍǡ Ǥ ϐ ȋ͕͗͜͜Ȃ͕͜͜͜Ȍǡ ȋǣȀȀǤ Ȁ͘ȌHistory of Animalsǡ ͙͔͗ Ǥ Dzephémeronǡdz Dzdz ȋ͕͛͝͝ȌǣDz Ǧǡ ǡǢǡ ǡϐȋephémeronȌ ǣǡǡ ǡ ǡ ǥdz ϐ ȋ͖͔͔͜ȌǤǡϐ ȋǤ͕Ȍǡ Ǥǡ ϐ Ǥ ϐ ϐ ϐ ȋ Ǥ͕͙͛Ȃ Ǥ͖͙͗ȌǡDe Ǥ͙ǡ Ǥ ȋ͕͙͝͝Ȍȋ͖͔͔͝Ȍ ϐ ȋ Dzǡdz Dzǡdz DzdzȌ ǤǤǡϐǦ ϐǤǤǡ ϐϐ ϐȋ͕͚͗͝ȌǣDz ͖ǡ͔͔͔ ϐϐǦ Ǧ Ǧϐ Ǥ Ǥ ϐǡϐ ϐǤdz ȋ͖͔͔͙Ȍ ϐǡ ǣ ȋǡ ǡ ǡ ȌǤȋ͕͙͝͝Ȍ ϐȋȌǡ Ǥ ϐǯ ϐϐǡ ǡ ϐ OligoneuriellaȋǣȌǤǡ ϐ ϐϐ ϐǡ ϐ Ǥ ϐ ϐ ǡ ϐ Ǥ ϐ ϐ ϐ Ǥ ǡ ȋ Ȍ Ǥ ǡ ϐϐǤ ǡ ȋȌ ϐ͙͔͔Ǥ ǡϐ ϐǡ ͙͕ ȋǣȀȀ Ǥ Ȁ͖͗͝ȌǤ Figure 1. Hexagenia limbata (Serville), (Ephemeroptera: Ephemeridae) taken at the Fall River; Fall River Mills, CA. July 3, 2010. Photo by M. P. Parrella. 16 American Entomologist • Spring 2013 ϐϐ ǤǤǡϐ ϐ Ǥ Ǥ ǡ ǤǤ ͚͘͝ϐ ȋǤǤ ǤȀϐȀǦ ǦǤȌǡ ͖͝ ȋǡȌϐȋǣȀȀǤ Ȁ ͖͛ȌǤ ȋ͕͙͙͝Ȍǡ ϐϐǡ ǣDz ǡǡ ϐ Ǥdz ϐ ϐ ǤǤ ȋ͕͚͔͜Ȃ͕͖͔͝ǡ ǤǤȌǡ ϐ Ǥ Ǥ ȋ͕͖͜͝ȌȋǤ͖Ȍ ǯThe Fly-Fisher’s Entomologyȋ͕͚͗͜Ȍ ǡ ϐǤ ϐ ϐϐ Ǥ ͕͜ ǡ ϐ ϐ ǡǡ ϐ Ǥǡ ϐϐ Ǧ͕ǡ͔͔͔ Ǥ ǡ ǡǤ ǡǡA Treatyse of Fysshynge wyth an Angle (TFA)͕͚͘͝ǤThe Boke of St. Albansǡϐ ǡ͕͚͘͜ǡ ǯTFA edition of the Boke Ǥ The Contribution of Juliana Berners TFAȋ͕͖͕͝Ȍǡ Ǥǯ ϐϐǡ ϐ ǡ and The Boke of St. Albansϐ ȋ Ȍ ȋ ͖͔͔͗ȌǤǡ͔͔͜ ͕͙͔͗Ǥϐ TFA ϐϐ ͕͖ ϐ Ǥ ȋȌǡȋ Ȍǡ ȋȌǡȋȌǡ ȋȌǡ ȋȌǡȋȌǡ ȋȌǡȋȌǡȋȌǡȋȌ ȋȌǤϐȄ American Entomologist • Volume 59, Number 1 Figure 2. The Mayflies of Marbury (1832, reprinted 1976) and their imitations. 189 – Pheasant, 190 – Puffer, 191 – Parker, 192 – Professor, 193 – Blue Professor, 194 – Poor Man’s Fly, 195 – Queen of the water, 196 – Quaker, 197 – Quack Doctor, 198 – Red Fox, 199 – Romeyn, 200 – Red Ant, 201 – Red Head, 202 – Red Ash, and 203 – Reuben Wood. Each of these flies (in addition to hundreds of others) is given its own detailed origin and history. Ȅ ȋȌ ȋ͕͖͛͝ȌǤ ȋǡǡȌǤ TFAϐǢ ϐǦ Ǥ ǡ ϐ Ǥ ϐ ǯ ϐϐ ǢǡȋȌ Ǥ ǯȋ͕͕͜͝Ȍ ϐϐǡ TFAǡ Ǥ ϐ TFAϐǡ ǡǡ ǡ ϐȋ͖͔͔͝ȌǤ ϐ ϐϐ ǡ Ǥ Ǥ ϐDzdzȋ doone ȌǤ ǯǦ ȋ͖͔͔͗Ȍǡ Dzdz ϐǦϐ ϐǤ ǡ ȋ͕͚͗͜Ȍ Dzdz 17 ϐǡϐ ϐDzdzȋ ȌǤ ǡϐ ǡ ǯ ǡ ȋϐȌ Ǥ ǡǡϐ ȋǤǤǡȌ ǡ ȋ ǡ͖͔͔͘ȌǤ ͕͖ϐ ͙͔͔ ǤϐDzǡdz ȋ͖͔͔͗Ȍǣ Dzϐ Ǣ ǡ ȋ͕͕͛͝ȌǦ ͕͔͛͝Ǥdz Markham Leads to Isaac Walton and Charles Cotton: The Compleat Angler ͕͙͔ǡ ϐ ϐ ȋ ͕͖͛͝ ͖͔͔͗ Ȍǡ ͕͚ TFAǤǯȋ͕͚͕͘Ȍ A Discourse of the Generall Art of Fishing, with the Angle, Or Otherwiseǡ ϐ ȋ ͕͙͛͝ȌǤ ϐ ϐǡ ϐ ȋ͕͖͕͝ȌǤ ǯȋȌ ϐϐTFAǡ͕͚͙͗ ǯThe Compleat Angler Ǥ ǡ ȋ ͖͔͔͗ȌǤ ϐ ǡȋ͖͔͔͗ȌǡDz ϐǤdz ǡ ϐTFAϐ Dzϐ ϐǤdz ϐ ǡ͕͝ ǡ Dzϐǡdz ȋ ǡ ǡǡ ǤȌǤ ϐThe Compleat Angler͕͚͚͛ǡ ϐǦϐ ϐ͚͙Ǥ ǡ ϐϐǤϐ ϐϐȋ ͕͕͜͝Ȍǡ ȋȌ ϐǤCompleat Anglerǡ͔͔͘ȋǤ͗ȌǤ ȋ͕͖͕͝Ȍϐǡ ǡ ǤDz dzϐ ǡ ǡ ǡǤ 18 ϐǡ ǡϐ ȋϐǦ Ȍ Ǥ ȋ ͕͖͜͝Ǣ ͖͔͔͚ȌǤ ȋ͕͖͜͝Ȍ ϐǤϐ ϐǤ ǡ ǡ Ǥȋ ǤȌ ǡǯ ϐ Ǥȋ͕͖͕͝ȌǣDz ǡ Ǥdz ϐϐ͖͔͔ ǯ The Compleat Anglerǡ ϐǡ ϐǤ ǡ ǡϐ Figure 3. Fly fishing during the time of Isaac Walton and Charles Cotton (late 1600s; from the Compleat Angler 1915 reprint). American Entomologist • Spring 2013 ϐϐ ȋϐȌǡ ϐǦ ȋ ͕͕͜͝ȌǤ Dz dz ϐ ϐǡ͕͚͛͘ ϐǡ ϐ ϐǤ Ǥ Alfred Ronalds and The Fly-Fisher’s Entomology Figure 4. The 30 most important flies as deemed by Bowlkers (1746): 1. Red fly; 2. Blue Dun; 3. March Brown; 4. Cowdung Fly; 5. Stone Fly; 6. Granam or Green Tail; 7. Spider Fly; 8. Black Gnat; 9. Black Caterpillar; 10. Little Iron Blue; 11. Yellow Sally; 12. Canon or Down Hill Fly; 13. Shorn Fly or Marlow Buzz; 14. Yellow May Fly or Cadow; 15. Grey Drake; 16. Orl Fly; 17. Sky Blue; 18. Caddis Fly; 19. Fern Fly; 20. Red Spinner; 21. Blue Gnat; 22 & 23. Large Red and Black Ants; 24. Hazel Fly or Welshmans’ Button; 25. Little Red and Black Ants; 26. Whirling Blue; 27. Little Pale Blue; 28. Willow Fly; 29. White Moth; and 30. Red Palmer. Ǥ ǡ ϐ ȋ͖͔͕͕ȌǤ ϐ ǡ ϐǤ ȋ͕͖͕͝Ȍǡ Ǥ Ǧ ǡ ȋ͕͚͛͘Ȍϐǡ ǯǯϐǤǯϐ Dz ǡ dz ȋ͕͖͕͝ȌǤ ǯϐ ͚͙ ǡ ǯ ϐǤ ǡϐ ǡ ǡ ǡ ǡȋǤ ͘ȌǤȋ͕͖͕͝Ǣ ȋ͕͚͕͜Ȍǡ ǤȌ American Entomologist • Volume 59, Number 1 Dz dz The Fly-Fisher’s Entomology ȋ͕͚͗͜ȌǤ ͛͘ϐ ϐǡ ϐ ǡȋǤ͙ȌǤ ȋ͖͔͔͗Ȍǡ ǡ ϐ ϐ ϐ Ǥǡǯ Ǣǡǡ ȋǣȀȀǤϐϐǤ ȀǤ ȌǤǡϐ ϐ ϐ ϐ ȋ Ǣ͕͖͕͝ȌǤ ǡ ϐǦϐ ϐ Ǥǡ ȋ͖͔͔͗ȌǤ ϐ Ǥ Figure 5. Mayflies from Ronalds (1836, reprinted 1990) and their imitations. 28a – Green Drake (May Fly, Cadow), 28b – Grey Drake (Glossy-winged Drake), 29 – Orange Dun. 19 Figure 6. Stewart’s (1857) flies tied with a palmer hackle (no wings). This is his “black spider,” a precursor to “wet flies” that are still tied today. ͕͚͗͜ ͕͚͔ǡ Ǥ Ǥȋ͖͔͕͕ȌDzThe Fly-Fisher’s Entomology ϐ Ǥdzǯ ϐ ϐ ǡϐ Ǥ ȋ͕͖͜͝Ȍǡϐ ȋǣ Ȍȋǡ͙͗Ȍ Ǥȋ͕͙͔͝Ȍ ȋǣ ȌǤ ǯ ͕͚͗͜ Dzdz ϐϐǤ ǡǦ Ǥ ǡȋ͖͔͕͕Ȍ ϐ ǤǤǡ Ǥǡǯ ǡ ǡ ȋ͕͛͗͝Ȍ ȋ ͕͖͖͝ȌǤ Stewart, Halford, Mosley, Skues, and Gordon: the Bridge from the U.K. to the U.S. ȋ͕͚͛͘Ȍ ȋ͕͚͗͜Ȍǡ ȋ͕͙͛͜Ȍ ϐǦϐ Ǥǡϐϐǡǡ ϐϐǤǡǡ ȋȌϐ ϐϐǤȋ͕͚͛͘Ȍ ϐ ͕͔͔ǡϐ ϐ ǡϐ ǦǤ ǡ Ǥ ϐϐ ϐǣ ȋ͕͙͛͜ȌǤ ǡ DzdzȋǤ͚ȌǤ ϐϐǡ ϐǡ ǡ ϐϐȋǤ Ǥ ȀǦǤȌǤ ϐϐǡ ϐǤϐϐǡ ǡϐ ǡϐȋǤǤǡ ȌǤ Ȁ Ȁ ȋDzdzDzdzϐǦ ϐȌǤϐǦ 20 ǡ ϐǡϐǡ ϐǤ ȋ ϐϐǦϐȌǤϐ ǡ Ǥ͕͔͘͜ǡϐ ϐϐǤ ǡ ͕͔͘͜Ȃ͕͚͔͜ǡϐ ǡ͕͔͜͜ǡϐ ϐǤ Ǥ Dz ϐϐdzȋ͕͖͕͝ȌǤ ϐϐǡ ϐ ȋ ͕͙͙͝Ȍϐ ϐϐǤǯDry Fly Entomologyȋ͕͛͜͝Ȍ ϐϐȋ ȌǤ ǡ ϐ ȋǤϐϐǤ ȀϐǤȌǤ ǯ ǡ ȋ͕͚͙͜Ȍ Ǥ Ǥ ȋ͕͗͜͜Ȍǡ ǡ Ǥ ǯ ȋ͖͔͕͕Ȍǡ ϐϐ ϐ Ǥ ǡ ǡȋ͕͖͕͝Ȍ Dzǯ Ǥ ϐǡ .dz ǡǯ Dry Fly Entomology ǡ ȋǤ͛ȌǤǯ Dz͕͕͔͝dzϐ Ȅϐ ϐ ϐ Ǥ ϐ ϐǡ Ǥ Dzϐ ǡ ϐ.dzǡ ϐǡ ǡ ϐϐǡ ȋ͕͖͕͝ȌǤ ϐϐ Ǥ ȋ͕͗͝͝ȌǡȋThe Dry-Fly Fisherman’s Entomologyǡ͕͖͕͝Ȍ Ǧ ǯ ǡ ȋǯ Ȍ ϐ ȋ ͖͔͕͕ȌǤ American Entomologist • Spring 2013 ǡ͕͖͕͝ ǡǡDz ϐǤ Ǩdz ϐϐǤǡ ǤǡϐǦϐ ϐ ǣϐǡ ϐǡǤ ǡϐ ͕͔͗͜Ǥϐ ϐϐ ȋ ͖͔͔͚Ȍǡ Ǥ ϐϐǡϐ ϐ ǯǡ Ǥ ǡ ϐ ǡ ϐϐ ϐϐ ǤǤȋ͖͔͔͛ȌǤ Ǧϐ ϐ Ǥϐ ǡ ǡ ǡǤ ǡϐ Figure 7. A beautifully drawn life cycle of Ephemerella ignita, the Blue-Winged Olive. Newly-born larva and eggs are shown in Fig. 1; a larva at the age of 3 months is shown in Figure 2; Figures 3 and 4 are the male and female subimago of this species, respectively (from Halford, 1897). ȋ͕͖͕͝Ȍǡ ȋǤ͜ȌǤǡ ǤǡǡDz Ǣ ϐ ǡ Ǥdz ǡϐϐ ϐǤ ǯ ǡϐǦ Ǥ ϐϐϐ ϐ ϐ Ǥ ϐϐ ȋ Ȍǡ ȋ͕͖͕͝Ȍ ϐȋ͖͔͕͔ȌǤ ϐ ϐǡϐ Ǥ ǡǤ American Entomologist • Volume 59, Number 1 Figure 8. Hand-drawn flies and their imitiations by West (1921). West was often criticized for the position of the wings (see text). 21 ϐϐȋ ͕͖͛͝ǡ͖͔͔͗ǡ͖͔͔͛ȌǤǡ ϐϐ Ǥ͕͔͔͝ǡ New York TimesǡDz ǡϐ ǯǡ Ǥ dzȋ͖͔͔͛ȌǤ Descent Into Confusion Figure 9. The Quill Gordon tied by Theodore Gordon, the progenitor of dry fly fishing in America. (photo by Valla [2009], with permission). Figure 10. A modern version of the Quill Gordon tied and photographed by M. Valla (2009) (with permission). This represents the Catskill style of dry flies. Ǥ ǤǤǡ ϐ ǡ ϐǤ ϐȋ ͕͖͛͝Ǣ͖͔͔͛ȌǤ ϐ ǡ ǡ ϐ ϐȋȌ ϐǤǡ ǡ ȋǤ͕͔͝ȌǤǡ Ȅϐ Ǥ ǡ ϐǤ ȋ͕͖͛͝ȌǡDz ϐ ǡ ϐ ϐǤ ϐ Ǥϐ ǤǮ ǡǯǤdz ϐǡ ϐϐǤ ǡ ϐϐ Ǥ 22 ϐ ǡ ǡ Ȁǡ ǡ ǡǡǡǡ Ǥ ϐ ϐǡ Ǥ ͕͛͝͝ǡ ǯ A Modern Dry-Fly Codeȋ͕͙͔͝Ȍ ǣ Dz Ǥǡ ǡ Ǥ ͕͚͜͜ǡ Ǥ ǡ Ȅ Ǥ ǡ ϐǡ ǡ Ǥdz ǣ Dzǣϐǡ ȋ Ȍǡ Ǥdz Entomology as a Young Discipline ϐ ǡ ȋ͕͖͛͝Ȍ Ǥ ǡǤ ǡDz Ȅ ǡ Ǯ ǡǯ ϐ.dz ǯ Dzdz ϐϐǤ ȋ͕͚͗͛Ǧ͕͚͔͜Ȍ ϐ ǡ ȋ͕͛͗͝Ǣ ͖͔͔͙ȌǤϐ American Entomologist • Spring 2013 Ǥ ǡ ϐȋ͕͘͘͝ȌǤ ȋȌ ϐϐ Ǥ ȀǡThe ͕͕͚͝ǡAmerican Trout Stream Compleat Angler ǡ Ǥ ϐ Insects. ǡ ͕͔͔ Systema Naturae ȋ͕͙͛͜ȌǤ Ǥ ͕͜ ϐ ǡ Ǥ ǡ ϐ ǡ ǡ ϐ ȋǤ͕͕ȌǤ ǡ± ȋ͖͔͔͛ȌǡDz ȋ ϐ Ȍ ϐǡϐ ȋ͕͛͗͝ȌǤ ǮǯǤdz Ǥ ϐ ǡ ϐ͖͔ ȋ ͕͕͜͝ȌǤ ǡ ͕͕͜͝ ȋ͕͛͗͝ȌǤ Dry Fly Entomologyȋ͕͛͜͝Ȍǡ Ǥ ϐ ǯ ͕͙͗͝ ǡ A Book of Trout Fliesǡ ϐ ǡǡǡ ǡ ϐ Ǥϐϐǡ ϐ ϐǤ ϐǤ Ǥǡǯ ǡ ȋ͕͛͘͝Ȍ ȋ͕͙͙͝ȌǤ ϐϐϐ Ǧ ϐϐϐ ȋ͕͚͙͜Ȍǡ Ǥ Ǥǡ ϐ ǣ The Situation Today Dzǡ ϐ ȋ͕͚͝͝Ȍǡ ǡ ǡǤdz Ǥ ǡ ȋ͕͖͜͝Ȍǣ Dz ǡ ϐ ϐ Ǥdz ǡ ϐ Ǥǡ ǡ ǡ ±±ǡ Ǥ ǡ ȋ ϐ Ȍ Ǥ ȋ͖͔͔͕Ȍǡϐ ϐ ͕͔͖͜ȋEphoron leukon Figure 11. A panel from Louis Rhead’s Trout Stream Insects (1916), showing the flies he drew and the patterns he Ȍǡǡ͚͝ designed to imitate the naturals. He devised his own classification scheme and none of his fly patterns have endured ϐ (see text). American Entomologist • Volume 59, Number 1 23 ϐ ǡϐǡ ǡ Ǥϐϐ Ǣ ͔͗Ȃ͔͘ ϐ Ǥǡ ϐ Ǥ ȋ͕͔͜͝Ȍǡ ȋ͕͕͜͝Ȍǡȋ͕͕͜͝Ȍǡ ȋ͕͖͜͝ȌǤ ǡ ϐ ϐ ȋ ͖͔͔͚ȌǤ ͕͚͔ ϐǡ DzǤǤǤǤǤdzȋ Ȍǣ ϐǦ ϐ ϐǫ ȋ͖͔͔͚Ȍ ǡ ϐϐ ȋȌ ǡ ȋ ͖͔͔͙Ǣ ͖͔͔͛Ǣ͖͔͔͝Ǣ ͖͔͔͝Ǣ͖͔͕͔ȌǤ ϐ ϐ Ǥȋ͕͖͕͝Ȍϐ ǡϐ Ǥ ȋ͖͔͕͕ȌǡDz Ǧ ϐ Ǥdz ȋ͕͖͛͝Ȍ ǡ ȋ ǡ ǡǦȌǡ ǡ ϐǤ ǡ ϐ Rhithrogena germanica Eaton ȋǣȌǤ ϐǤ ǡ ϐ Ǥ ϐǦϐ ǯ TSA. ȋ͕͖͕͝Ȍ ȋ͖͔͕͔Ȍǡȋ͕͖͕͝Ȍ ϐǢǡ Ǥ ȋ͕͚͕͜ȌǤȋ͕͚͛͘Ȍȋ͕͚͗͜Ȍ ǡ ǡ Baetis longicaudaǤ Ǥȋ͕͖͕͝Ȍ R. haarupiǡ R. germanicaȋǣȀȀǤ Ȁ 24 ͖͚͖͝Ȍȋ ǡ ͕͖͕͝Ǣ͖͔͕͕ȌǤ ϐ ǤǤǡ Ǥ ǤǤǤǤǡ ϐ ǤǡȋǤ Ǥ Ȁ Ǧ ǤȌǡ Ǥ ǡ ϐ ϐ Ǥ ϐǡ Ǥ ǡ Ȅ Ǥ ǡ ǡǡǦǡ Ǥǡ ǡ Ǥ ϐ ȋǤǤǤȀϐȀǦ ǦǤȌǡMaccaffertium vicariumȋȌǡ ǡ Ǥ ȋǣȀȀǤ Ȁ͗ȌȄ ǡ ǡ ǡ ǡǡ ǡ Ǥ ǤǤǡRithrogena morrisoniȋȌ ǡ Ǥ ϐǤ Ephemerella subvariaȋ Ȍ ȋǣȌǡ ǡϐ ǡ ϐǦϐ ǤǤ ȋ ͖͔͔͛ȌǤǡ Ǥǡ Ǥ Dz dz Empherella ȋ͖͔͕͔ȌǤ ǡǡ ϐϐ ϐϐǤǡϐϐ Ǥϐϐ Ǧǡ ǡ ǡ ϐ Ǥ Ǧ ǡ Ǧϐǡ Ǥǡȋ͖͔͕͕Ȍ Ǥ American Entomologist • Spring 2013 Scientific vs. Common Names (from Bosik 1997) ǡ ϐ ȋǣȀȀ ǤȀȌǤǡ ǡ ǯ ϐ Ǥǡ ϐ Ǥ ȋ Ȍǡ ϐ Ǥ ǡ ϐ Ǥ ǡ ȋǤǤǡ ϐϐȌǡ Ǥ ǡ ǡ ǡ Ǥǡ Ǥ Ǧ ϐ ǡ ǡ Ǥ͕͔͗͝ǡ ȋȌ Ǥ ϐ͕͔͜͝ ͕͖͘ Ǥǡ͖ǡ͔͔͔ ȋǤ ǤȀȀ ̴ȌǤ Dzϐ dz Ǥ ϐǡ ϐǡϐǤ ϐ ǡ ȋ͕͕͜͝Ȍ ϐ ϐ ϐ Ǥ ǡ ǤϐȋǤ ǤǤȀϐȀȌǦǦ ǡ Ǥ ǡϐ ϐǤ ǡ Ǥ ϐ ϐ ǡ ȋǣȀȀǤ Ȁ͜͝Ȍ ȋǣȀȀ Ǥ Ȁ͚͜ ͜Ȍ ϐ ϐǤ ǡ ȋ ǣϐȌ Ǥ ǡ ϐϐ Ǥ American Entomologist • Volume 59, Number 1 ϐϐǡ Ǥȋ͖͔͕͕ȌDzǦ dzϐǡ ǡDzǦ dzǤ ǡǤ ϐϐ ϐ ǡ Ǥ ȋ͖͔͔͙Ȍ ϐ Ǧǡ Dzdz Ǥ ϐ ϐ ȋǣȀȀǤ Ȁ͘͝ȌǤ ȋ ϐǡ ǡ ǡ ǤȌǤ ϐǡ ȋȌ ǡ ϐȋǤǤǡȌ ȋǤǤǡBaetis, Hexageniaǡ ǤȌǤ ϐǦϐ ǡ Ǥȋ͖͔͕͕Ȍ ȋ ϐϐȌǡϐϐ ȋ Ȍ Ǥ ǡ ϐϐ Ǥ ȋ͕͚͙͜Ȍϐϐϐ ǣ Dzϐϐǡ ǡ Ǥ Ǣ ǡ ǥ ǡ ǡ ϐ ǡ ǤǤǤ ǡ Ǥdz ͕͜Ǥ͖ǤǤ ϐϐǡ͖Ǥ͝ȋǣȀȀ Ǥ Ȁ͗ȌǤȋ͖͔͔͛Ȍ ϐ ϐ Ǧ ǡ ȋǤǤǡϐǡǡǡǡǡ ȌǤǦ ϐϐǡ ǤǦ 25 Ǥ ǡ ϐϐ Ǥ Ǥȋ͕͙͔͝Ȍȋ ͕͕͛͝Ȍϐϐ ǣ Dzϐ ϐ ϐ ϐǦ ǡϐ Ǥ ǡ ϐ ϐǡ ǡ ϐǡ ϐ ϐǤϐǦ ϐǤ ǡ ǡ Ǥdz Final Thoughts ϐϐ ǯTFAǤ ǡ ϐǦϐ ȋ͕͖͕͝Ȍǡȋ͖͔͔͗Ȍǡ ȋ͖͔͕͕ȌǤ ϐϐ Ǥǡ ϐ ϐ ǡ ϐ Ǥ ϐ ϐ ǡ Ǥ ǡ Ǥ ǡ Ǥǡ ǡ Ǥ ϐϐǣǤ ͕͕ǡ͔͔͔ ϐϐǤ ϐ ϐϐǡ ϐ Ǥǡ ϐ ϐ Ǥ ϐ ϐǡ Angler’s Almanac ȋ͕͙͕͜ȌǣDz ϐ 26 ǢǤ ǡ ϐϐǤdz References Cited ǡǤǡǤ͖͔͔͝Ǥϐǣ Ǥ ǡ ǡǤ ǡǤ͖͔͔͔Ǥ ϐǫ ϐ Ǥ ǡ ǤǡǡǤ ǡǤ͖͔͕͕ǤϐǣϐϐǤ ǡ͕͛ǤǣȀȀǤ Ȁ͖͔͕͕Ȁ͔͙Ȁ͕͛ȀǦ ǦǦǦϐǦϐȀ ǡǤǤ͕͙͝͝Ǥǣ Ǥ ǡǡǤ ǡǤǤ͕͚͘͝Ǥ ǤǤ ǡǡǤ ǡ ǤǤ͕͔͜͝Ǥǣ Ǥ ǡǡǤ ǡǤǤ͕͛͝͝Ǥ Ǥ ǡǤ ǡǤ͕͚͛͘ȋ͕͙͘͜ȌǤǯǣ ϐǦϐǡǡϐ ϐǡƬ Ǥǣ ϐ ǤǡƬǤǡǡǤǤ ǡǤǤ͕͙͕͜Ǥϐǯvade mecumǡǯ ͕͙͕͜ǤǤƬǤǡǡǤ ǡǤ͕͚͕͜Ǥǯvade mecumǡǡ ǡ ǡ ǡ ǡǡ ϐǤ ǡ͖͔͕͕Ǥ ǡǤ͖͔͔͙Ǥ ǯDe natura animaliumǤǤ͛͘ǣ͛͘͝Ǧ͗͘͜Ǥ ǡǤǤ͕͘͘͝Ǥ Ǥ ǡ ǡǤ ǤǤǤ͕͚͗͝Ǥ ϐǤǡ ǡ ǡǤǡǡ ͕͕͛͝Ǥ ǡǤ͖͔͔͙ǤǣϐϐǯǤ ǡ ǡǤ ǡǤǤǤ͖͔͔͛Ǥǣϐ ϐǯ Ǥ ǡǡǡǤ ǡǤǤ͕͗͜͜Ǥ ϐǤ ǡ ǣǤ ǡǤǤ͖͔͔͙Ǥ ǡ͕͛ǣ ͕͚͔͔Ǥ ͚͜ǣ͕͗͗Ǧ͕͘͘ǤǣȀȀ ǤȀȀ Ȁ̴̴Ǥ ǡǤ͖͔͔͙ǤϐǤ ǡ ǡǤ ǡǤ͕͛͘͝Ǥǣ ȋ͖͔͔͛ȌǤǡ ǡ Ǥ ǡǤǤ͖͔͔͗Ǥ ǣ ǡ ǤǤ ǡǡǤǤ ǡǤǤǤ͕͕͜͝Ǥ ǣǯϐǤ ǡǡǤ ǡǤǤǤ͖͔͔͘Ǥϐ ǣ ϐ Ǥ ǡ Ǥ ǡǤ ǡǤ͖͔͔͚ǤǦϐǣǯ Ǥ ǡ ǡǤ American Entomologist • Spring 2013 ǡǤǤ͕͛͜͝Ǥϐǣ ͕͔͔ϐϐǤƬǤǡǤǡǤǤ ǡǤ͖͔͕͕ǤϐϐǤǡ ǡǤǤ ǡǤǤ͕͖͕͝ǤǤ ƬǤǡǡǤǤ ǡǤǤ͕͙͗͝Ǥϐǣ Ǥ ǡ ǤǡǤ ǡ Ǥ͖͔͔͗Ǥ ϐϐǤǡ ǡ Ǥ ǡǤǤ͕͙͔͝Ǥǡǡǡǡǡ ǡ ϐǡǡǡϐ ǡ ͖ǡ͖͔͔ǤǤǤ ǡ ǤǡǤ ǡǤǤ͕͛͗͝Ǥ ǣ͕͔͔͜Ǧ͕͙͜͝ǡǤ͕͕͝Ǧ͕͙͘ǤInǤǤǡǤǤǡ ǤǤȋǤȌǤǤ Ǥ ǤǡǡǤ ǡǤ͕͙͛͜Ǥ§§ǡ ǡǡǡ ǡ ǡ ǡǡ ǤǤǡ ǡǤ ǡǤ͖͔͕͕ǤϐǤ ǤǣȀȀǤ Ȁ͖͔͕͕Ȁ͔͙Ȁ͕͚ȀǦ ǦǦ ǦǦϐȀ ǡǤ͕͙͝͝ǤϐǤ͕͖͔ǣ͗Ǧ͕͗Ǥ ǡǤǤ͕͖͜͝ǤϐǤ ǡ ǡ Ǥ ǡǤǤ͕͙͔͝ȋ͕͛͝͝ȌǤǦϐ Ǥ ǡǡǤ ǡ Ǥ͕͚͕͘Ǥ ϐ Ǥ ǡǤ ǡǤǤ͕͕͜͝Ǥ ǣϐǯ ǯ Ǥ ǡǡ Ǥ ǡǤǤ͖͔͔͕Ǥǣ͖͔͔ ϐǡǤ͖͕Ǧ͙͗ǤIn ǤǤȋǤȌǤ ȀǡǡǤ ǡǤ͕͖͛͝Ǥ ǤǤǡ ǤǡǤ ǡǤǤǤǤ͕͙͛͝ǤϐǤ ǡ͖͛ǤǣȀȀǤ Ǥ ȀȀ ȀȀ ͕͕͖͖͗͗͘Ȁ͕ȀǤ ǡǤ͕͚͙͜Ǥ ǣ ǦϐǤ ǡǤǣǤ ǡǤǤǤǤȋǤȌǤ͕͚͝͝Ǥ ǢǤȀ ǡǡǤ ǡǤǤ͕͖͖͝Ǥ Ǥ ǡǤ ǡ Ǥ ͕͖͕͝Ǥ Ǧϐ ϐǯ Ǥ ƬǡǤǤ ǡǤ͕͗͝͝Ǥ ϐǤ Ƭ ǤǤǤ ǡǤ͖͔͔͝Ǥǣ Ǥ ǡǡǤ ǣȀȀǤǤ Ȁ ǡǤǡǤǤǡǤ ǡǤǤ͖͔͔͚Ǥ EnϐǦϐǤ ͙͖ȋ͖Ȍǣ͖͛Ǧ͛͘Ǥ ǡǤ͕͖͜͝Ǥǣϐ ǤƬǡǡǤ ǡǤ͕͕͚͝ȋ͖͔͔͝ȌǤ Ǧ ǣ ϐ American Entomologist • Volume 59, Number 1 ǡ Ǥ ǡǡǤ ǡǤ͕͚͗͜ȋ͕͔͝͝ȌǤϐǦϐǯǤ ϐǡ ǡǤ ǡǤǤ͕͚͖͜Ǥϐ ϐ ǡ ϐ ϐǡ Ǥ Ǥ ǡǤ ǡ Ǥ ͖͔͔͛Ǥ ϐǦϐ Ǥ ǡ ǡǤ ǡǤ͖͔͔͚Ǥǣǡ ϐϐϐǤ ǡ ǡǤ ǡǤ ǤǡǤ͕͙͙͝Ǥ ǣ Ǥ ǡǤ ǡ ǤǤǤ͕͖͕͝Ǥϐ Ǥ ǡǤǡ Ǥ͖͕͛Ǥ ǡǤ͕͛͝͝ǤϐȋȌǣ ǡǤ͙͕͕Ǧ͙͕͗ǤInǤƬǤ ȋȌǤƬ ǣǡ ǡ Ǥ ǡǡ͕͙͝͝Ǥ ǡǤǤ͕͖͜͝Ǧ͕͔͗͜ǤǤ ǡǡǤǤ ǡǤǤ͕͙͛͜ȋ͕͔͛͝ȌǤ Ǧϐ Ǥ ǡǡǤǤ ǡǤǤ Ǥ͕͕͛͝Ǥ Ǥ ͖͔͔͔ǡǤ ǡǤ͖͔͕͔Ǥǣϐϐǯ ϐ Ǥ ǡ ǡǤ ǡǤǤ͕͛͗͝Ǥ ǣ͕͔͔͛Ǧ ͕͕͙͜ǡǤ͙͝Ǧ͕͕͜ǤInǤǤǡǤǤǡǤǤ ȋǤȌǤǤǤ ǤǡǡǤ ǡǤǤȋǤȌ͕͕͛͝Ǥ Ǥ ǡǡǤ ǡǤ͖͔͔͝ǤǦϐǤǡ ǡǤ ǡǤ͖͔͔͛ǤϐǤǡǡǤ ǡǤ͕͚͙͗ȋ͖͔͔͗ȌǤ ǯ Ǥ Ǥǡ ǡǤ ǡǤ͕͚͚͛ȋ͕͙͛͜ǡ͕͕͙͝ȌǤ ǯ ǣ ǡ ϐǦǡϐǡϐ Ǥǡ ǡǡǤǤ ǡ Ǥ ͕͖͕͝ ȋ ͖͔͕͔ȌǤ ϐ ǤǡǡǤ ǡǤ͖͔͔͚ǤǤ ǡǤǤ ǡǤ͕͖͜͝Ǥ Ǥǡ ǡ Ǥ ǡǤǤ͖͔͔͜ǤϐǤ ǣǡ͕͚Ǥ Ǥ ȀȀ͖͔͔͜Ȁ͔͝Ȁ̴̴̴ϐǤ Michael P. Parrella ǡǡǤ DzdzȋǤ͖͔͔͚ȌǤ Ǥ 27
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