146 5 spp. spp. 5309 Apioceridae Finding the Fly Tree of Life spp. Pachyneuridae 483 spp. 760 6187 7479 spp. spp. Bombyliidae Mydidae spp. 33 (bee flies) Bibionidae Cecidomyiidae spp. (mydas flies) Asilidae (march flies) (gall midges) 32 277 Lygistorrhinidae 136 spp. 392 spp. (robber flies) 20 spp. spp. 134 spp. Hilarimorphidae spp. Nemestrinidae Pantophthalmidae Xylophagidae Acroceridae (tangle-veined flies) 94 2302 (small-headed flies) spp. spp. Xylomyidae 2666 Ditomyidae Sciaridae 4164 spp. 390 16 spp. spp. spp. 32 (dark-winged fungus gnats) Stratiomyidae spp. Mycetophilidae ASILOIDEA 1 (soldier flies) spp. Scatopsidae Canthyloscelidae Rangomaramidae (fungus gnats) STRATIOMYOMORPHA (minute black Evocoidae 417 scavenger flies) 711 spp. spp. TABANOMORPHA Scenopinidae SCIAROIDEA Rhagionidae 60 (window flies) (snipe flies) spp. 6 59 BRACHYCERA spp. spp. Vermileonidae 945 (worm lions) Axymyiidae spp. Bolitophilidae 1 Keroplatidae spp. 159 (predaceous 4 spp. spp. 1131 Valeseguyidae fungus gnats) FLIES & FLY DIVERSITY spp. Anisopodidae Apsilocephalidae Therevidae 1 (wood gnats) hen you think of a "fly," chances are you picture one of only a few common pest species: perhaps the house fly (Musca domestica), the laboratory spp. (stiletto flies) W Oreoleptidae 34 spp. fruit fly (Drosophila melanogaster), or the malaria mosquito (Anopheles gambiae). In fact, there are over 154,000 known species of flies, most of which are rarely noticed by a Diadocidiidae casual observer, and never bother humans. They are found nearly everywhere on earth, from Antarctica to the rain forest. Please take a minute to explore the fascinating diversity of flies. BIBIONO- MORPHA What is a fly? The importance of flies Fly evolution 2958 Flies make up the order Diptera, meaning "two wings"; nearly The great biological diversity of flies makes them important to man and Dipterists (scientists who study flies) classify flies into about spp. 38 4450 spp. spp. Psychodidae 5 all other insects have four wings. "Higher" flies (Brachycera) have nature in many different ways. Although we may be disgusted by flies 150 different families. Each of these has its own unique structure Tanyderidae spp. Tabanidae (moth flies, sand flies) a highly developed flight mechanism, and some of these (such as breeding in our waste and filth, they have a very important role as and way of life. The first flies appeared over 250 million years ago, 50 (primitive crane flies) Perissomatidae spp. (horse flies) horse flies) are among the fastest and most agile insects in flight. decomposers in nature. Others are serious agricultural pests, feeding on before the time of the dinosaurs, and looked much like modern crane Pelecorhynchidae Unlike other insects, adult flies cannot chew their food. Instead, crops and livestock, or are beneficial parasites and predators of unwanted flies, but biggest "explosion" of fly diversity (especially in the group 124 their mouthparts are modified for either piercing and sucking (to pests. In addition, flies serve as pollinators. Many plants, including cacao (the Schizophora) has happened in the last 70 million years. Possible spp. Athericidae 324 spp. feed on blood or nectar), or sponging (to lap up fluids). Fly larvae source of chocolate), are dependent on fly pollinators. Some flies do deserve relationships between fly families are shown on this phylogeny ("family (watersnipe flies) Blephariceridae (maggots) are also distinctive in having no legs, and most also their bad name - bloodsucking flies such as mosquitoes serve as vectors for tree"), based on new research by an international team of dipterists. By (net-winged midges) have no visible head. Apart from these basic features, flies come dangerous diseases, including malaria, yellow fever, and leishmaniasis, and comparing anatomical structure and DNA sequences from each of the fly 2079 in a many different colors, shapes, and sizes. are responsible for millions of deaths annually. families, we are learning exciting new details about the evolution of flies. spp. Simuliidae (black flies) 182 spp. PSYCHODO- 11 Thaumaleidae MORPHA spp. 7236 (seepage midges) Atelestidae spp. 33 Dolichopodidae 174 7 spp. 35 spp. spp. (long-legged flies) spp. 321 3 Cryptochetidae spp. spp. Coelopidae 1 Rhinophoridae Braulidae 606 spp. 4003 Canacidae spp. Phaeomyiidae (seaweed flies) (woodlouse flies) spp. (bee lice) 1992 (beach flies) 104 Mystacinobiidae spp. spp. 74 Sciomyzidae Drosophilidae spp. (New Zealand 9622 278 spp. Ephydridae spp. Corethrellidae (marsh flies) 12 bat flies) (vinegar flies) Ptychopteridae spp. 1 (shore flies) (frog midges) Tachinidae spp. Milichiidae (phantom crane flies) Helcomyzidae (freeloader flies) 190 Mormotomyiidae spp. 9 1971 spp. spp. Oestridae 64 25 spp. Australimyzidae spp. (bot flies) Hybotidae Curtonotidae 57 spp. Dryomyzidae (quasimodo flies) 23 Chaoboridae 783 spp. CULICO- spp. (phantom midges) EMPI- MORPHA Conopidae Helosciomyzidae 1898 OESTROIDEA (thick-headed flies) spp. DOIDEA 2880 147 spp. 13 Calliphoridae spp. spp. (blow flies) Xenasteiidae Brachystomatidae 3094 40 Chloropidae spp. 48 spp. spp. (grass flies) Sarcophagidae Camillidae Diastatidae EPHYDROIDEA (flesh flies) 8 1 spp. spp. 3685 Huttoninidae 414 spp. Natalimyzidae spp. 1927 1896 spp. Culicidae (mosquitoes) spp. Scathophagidae Anthomyiidae Lauxaniidae (dung flies) 781 29 (root maggots) spp. spp. Hippoboscidae 3052 5 spp. 14 spp. Fergusoninidae (louse & bat flies) spp. (eucalypt gall flies) 350 2 SCIOMYZOIDEA Neminidae Acartophthalmidae spp. spp. 384 2977 Empididae spp. spp. (nobody flies) (dance flies) Chamaemyiidae Heterocheilidae Sepsidae 115 Agromyzidae spp. (silver flies) (ant-like scavenger flies) Celyphidae (leaf-mining flies) 23 (beetle flies) LAUXANIOIDEA spp. 5644 spp. Glossinidae 70 63 (tsetse flies) spp. Ceratopogonidae 34 spp. spp. (biting midges) 5210 Aulacigastridae 136 Lonchopteridae spp. spp. 1 Ropalomeridae 64 Asteiidae spp. (spear-winged flies) Muscidae spp. 93 Odiniidae Apystomyiidae 3 675 (house flies) spp. spp. spp. 66 350 spp. Anthomyzidae CYCLO- spp. Tachiniscidae Ulidiidae (picture-winged 59 RRHAPHA Pyrgotidae Pallopteridae 359 spp. flies) 504 spp. 252 (flutter flies) spp. 32 90 spp. 186 Opomyzidae spp. spp. spp. Fanniidae Lonchaeidae Periscelididae Carnidae Platypezidae Dixidae (lance flies) (small house flies) (flat-footed flies) 7054 (meniscus midges) 82 spp. spp. CALYPTRATAE Chironomidae Piophilidae (common midges) (skipper flies) 360 spp. 20 spp. 4711 Clusiidae 11 spp. (druid flies) Neurochaetidae spp. 3 TEPHRITOIDEA (upside-down Inbiomyiidae spp. Tephritidae 4 flies) spp. Ironomyiidae (true fruit flies) Opetiidae (ironic flies)

1162 175 spp. spp. 8 112 spp. 15214 10 Richardiidae 33 spp. spp. spp. Platystomatidae Teratomyzidae (signal flies) spp. Ctenostylidae Cypselosomatidae Neriidae (fern flies) Tipulidae (crane flies) (cactus flies)

4105 spp. 3 spp. Phoridae What do flies do? Marginidae 160 (scuttle flies) spp. 579 Trichoceridae LARVAE: ADULTS: spp. (winter crane flies) Micropezidae decomposers (stilt-legged flies) SCHIZOPHORA 7 pollinators spp. 1420 Nymphomyiidae blood- spp. fungi 320 feeding spp. Pipunculidae (big-headed flies) Psilidae 6016 15 spp. living plants spp. (rust flies) 7 predators spp. 5 Megamerinidae spp. Syrphidae Somatiidae Nannodastiidae 14 vertebrate (flower flies) spp. predators parasites 138 Deuterophlebiidae invertebrate spp. (mountain midges) Chyromyidae parasitoids 10 spp. vertebrate Syringogastridae Photo Credits: www.bugguide.net: Tom Murray (live shots, 55 families); Stephen Luk (Dixidae, Chamaemyiidae, Pipunculidae); Andrew Gale (Neriidae); Gayle & Jeanell Strickland (Calliphoridae, Sciomyzidae, Tanyderidae, Muscidae); Gary McDonald (Carnidae, parasites Scatopsidae); Phillip Kline (Oestridae); Kevin Hall (Pachyneuridae); Lynn & Gene Monroe (Granite Ridge Nature Institute) (Apioceridae); Steve Kortum (Nemestrinidae); Edward Trammel (Acroceridae, Culicidae, Syrphidae, Xylomyidae); Ilona Loser (Anthomyzidae); Lynette Schimming (Platypezidae); Keith Bayless (NCSU) (Ropalomeridae, Richardiidae); Shaun Winterton 191 (Apystomyiidae, Apsilocephalidae, Scenopinidae); Mike Boone (Pyrgotidae); www.diptera.info: Jorge Almeida (Phaeomyiidae, spp. Helcomyzidae, Hippoboscidae); Dmitry Gavrushin (Canthyloscelidae); Vida van der Walt (Curtonotidae); Albert de Wilde aquatic (Heterocheilidae); Takeyuki Nakamura (Nymphomyiidae); Ramon Batlle (Vermileonidae); Ben Hamers (Opetiidae, 1568 Tanypezidae); Arthur Anker (Pantophthalmidae); Walter Pfliegler (Thaumaleidae); other sources: Karsten Sund Diopsidae 728 spp. (www.sciaroidea.info) (Ditomyiidae, Diadocidiidae); Alex Wild (www.myrmecos.net) (Cryptochaetidae, Odiniidae, spp. Piophilidae); Matt Bertone (NCSU) (Chaoboridae, Lauxaniidae, Micropezidae, Rhinophoridae, Tabanidae); Alessandra (stalk-eyed flies) 8 21 47 Rung (Cal. Dept. Food & Agric.) (Aulacigastridae); Robert Copeland (ICIPE) (Mormotomyiidae); David McClenaghan spp. spp. spp. Sphaeroceridae (CSIRO Entomology) (Pelecorrhynchidae); Rod Morris, New Zealand Geographic (www.nzgeographic.co.nz) Poster: Isaac Winkler, North Carolina State University, Sept. 2011 (ver. 4) Heleomyzidae (Mystacinobiidae); www.munisanignacio.gob.pe (Tachiniscidae); Isaac Winkler (NCSU) (remaining specimen Nothybidae 5 (small dung flies) shots); from published literature: D.K. McAlpine (1997) Rec. Australian Mus. 49:167-194 (Gobryidae); Jaschhof This research was performed by the FLYTREE consortium, an international spp. Tanypezidae Strongylophthalmyiidae & Didham (2002) Studia Dipt. Suppl. 11(Rangomaramidae); Colless (1990) Ann. Soc. Entomol. Fr. 26:351-353 collaboration led by Brian Wiegmann (NC State University). (sun flies) (Valeseguyidae); Geoffrey Attardo (2008) PLoS Neglected Tropical Diseases, 2(3) (Glossinidae); Nadine Maynard, Funding provided by the US National Science Foundation (EF-0334948) Zloty et al. (2005) Syst. Entomol. 30:248-266 (Oreoleptidae); Jill Marie Mullett, Yeates et al. (2003) Syst. Entomol. Gobryidae 28:417-431 (Evocoidae); F. Zeledón, Buck (2006) Syst. Entomol. 31:377-404 (Inbiomyiidae). For more information, see: www.inhs.illinois.edu/research/FLYTREE Wiegmann et al. (2011) Proc. Natl. Acad. Sci. USA, doi: 10.1073/pnas.1012675108