, 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2

Where are we Now: Where are we Now? An Overview of the •Numbers Systematics of Flower • How we got in this Mess – A short history of Syrphidae Classification F. Christian Thompson • What we can learn from the Past Systematic Entomology Lab., USDA – Words of Wisdom for Today Washington, D. C. 2 September 2005 2 September 2005 Overview of flower Systematics 1 2 September 2005 Overview of flower fly Systematics 2

Names = Palaearctic 2,041 species Syrphidae 200 • 3 Subfamilies 180 160 • 14 Tribes 140 • 11 Subtribes 120

• 292 Genera and subgenera 100 Series1 Years •189 Genera 80 • 5,845 Species 60 • 8,807 Name records 40 20 – Per BioSystematic Database of Diptera 0 8 5 8 0 7 5 7 2 9 • Version 7.5 release date 15 September 2005 32 39 53 23 30 94 822 900 908 966 987 001 175 177 178 1797 1807 1 18 18 1846 18 186 186 1876 188 1893 1 1 1915 19 19 193 1944 195 195 1 1973 1980 1 19 2 Names 2 September 2005 Overview of flower fly Systematics 3 2 September 2005 Overview of flower fly Systematics 4

Names = Palearctic Names = New World 2,242 species

4000 160

3500 140 3000 120 2500 100

2000 Series1 80 Series1

Names Names 1500 60 1000 40 500 20 0 0

8 7 3 5 8 7 6 3 7 8 0 3 5 3 2 4 8 9 22 46 0 30 52 80 77 92 8 04 30 807 867 900 959 994 001 775 804 834 848 948 962 975 005 1758 1775 178 179 1 18 1832 1839 18 185 186 1 1876 188 1893 1 190 1915 1923 19 193 1944 19 1 196 197 19 198 1 2 175 1 179 1 182 1 1842 1 185 186 1869 18 1884 18 189 19 1912 1918 1924 19 1936 194 1 195 1 196 1 1982 1990 199 2 Years Years 2 September 2005 Overview of flower fly Systematics 5 2 September 2005 Overview of flower fly Systematics 6 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2 1 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2

Names = New World 4000 Gunilla’s 3500 Questions: 3000 2500 How did we

2000 Series1 get into this Names mess? 1500 1000 Explain your 500 classification?

0

4 5 4 8 4 6 5 04 42 12 42 54 90 99 823 863 869 877 930 948 962 1758 1775 1790 18 1 183 18 1848 185 1 1 1 1884 1892 1898 190 19 191 192 1 193 19 1 19 1 1968 1975 1982 19 19 200 Years 2 September 2005 Overview of flower fly Systematics 7 2 September 2005 Overview of flower fly Systematics 8

“Traditional*” Classification “Traditional” Classification

• Microdontinae • – Merodontini • – Spheginobacchiini – Brachyopini • Brachyopina – Bacchini – Pipizini – Rhingiini • Spheginina – Paragini – Callicerini – Cerioidini – Syrphini – Volucellini – Milesiini • Criorhinina – Eristalini – Toxomerini •Blerina • Sericomyiina • Temnostomina • Eristalinae (continued) • Helophilina • Tropidiina •Eristalina *Yes, Gunilla, a mess, but this is what we use for the • Milesiina BioSystematic Database of World Diptera. Maybe a • Xylotina Finn can clean it up ☺ 2 September 2005 Overview of flower fly Systematics 9 2 September 2005 Overview of flower fly Systematics 10

Linnaeus (1758) Fabricius 1775 - 1805 37 Syrphids 1775, All syrphids to the All SYRPHUS In genus But by 1805 they were scattered among half dozen Musca genera

Gunilla, it took a Great Dane To get it Right ☺ Well, Gunilla, it started with a Swede! 2 September 2005 Overview of flower fly Systematics 12 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2 2 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2

Latreille (1797) Meigen (1803-1838) • Expansion of generic classification • A family for flower flies • 1803 => 10 new genera • 1822 => another 10 new genera

So, Gunilla, it took a Frenchman to give us our own family ☺ But, Gunilla, it is the Germans who do it Right! 2 September 2005 Overview of flower fly Systematics 13 2 September 2005 Overview of flower fly Systematics 14

Newman 1835 Edward Newman 1835

• First subfamily classification – Chrysotoxites • Long antennae (Microdon, Chrystoxum) –Syrphites • Syrphities So, Gunilla, while Alan does • Eristalities not believe it, it took an Englishman to order the ever • Volucellities increasing number of genera • Rhingities ☺

2 September 2005 Overview of flower fly Systematics 15 2 September 2005 Overview of flower fly Systematics 16

Rondani 1845, 1856 Schiner 1865 etc •Syrphinae • Volucellina => plumose arista (Linnaeus) • Volucellinae • Eristalina => R4+5 sinuate (Harris 1776) • Syrphina => everything else • Eristalinae • Microdonina => scutellum tubercles • Milesiinae => Apical r-m crossvein • Callicerina => antenna with terminal style • Chrysotoxinae • Microdoninae And an Italian to turn it into Pasta! • Cerinae Naturally, one needs an Austrian to bring Order with analytical keys, etc., BUT … 2 September 2005 Overview of flower fly Systematics 17 2 September 2005 Overview of flower fly Systematics 18 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2 3 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2

Williston 1883, 1887 Sorry, but •Syrphinae • Eristalinae • I forgot the – Pelecocerini* – Eristalini – Frenchman => Bigot – Microdonini – Volucellini • Bacchini – Bacchini –Sericomyini • Xylotini – Brachyopini* – Milesini •Etc. – Syrphini – Xylotini* – The other Italian => Lioy – Cheilosini* • Cerinae – Melanostomini* – The other Austrians => Brauer, et alia Finally an American, with more Resolution, but no VISION! But, Gunilla, I only have 20 minutes ☺ 2 September 2005 Overview of flower fly Systematics 19 2 September 2005 Overview of flower fly Systematics 20

George Verrall 1901 Shannon, 1921-23

• Nausigasterinae* • Chilosinae • Followed tradition (Schiner, Williston) • Cerioidinae – Pipizini • Microdontinae – Spheginini • But give an evolutionary scenario – Pelecocerini • Syrphinae –Paragus, as most primitive – Brachyopini – Melanostomini – Chilosini – Ceria, as most advance –Syrphini – Bacchini • Eristalinae – With , Cheilosines, Syrphines, Volucellines, – Paragini – Helophilini – Eristalini Eristalines and Chrysotoxines in between • Volucellinae • Xylotinae • Sericomyiinae –Xylotini Gunilla, apparently Darwin had effect • Milesiinae – Temnostomini* on the British! 2 September 2005 Overview of flower fly Systematics 21 2 September 2005 Overview of flower fly Systematics 22

14 subfamilies 17 tribes Shannon Hull, 1949 Developed • Calliceratinae character • New Characters • Syrphinae • Pelecoceratinae – Syrphini • Volucellinae morphoclines, – Bacchini • Sericomyiinae especially – Postpronotum, pilose or bare – Melanostomini • Xylotinae facial shape – Number of Male pre-genital segments • Chrysotoxinae – Xylotini • Microdontinae – Temnostomini – Malloch = Metafemoral basal setal patch •Eumerinae –Milesini • Nausigasterinae – Criorhinini –Etc. • Cheilosinae – Pocotini – Rhingiini – Tropidini • Syrphinae, clearly defined as well as – Pipizini •Psarinae – Chrysogasterini • Cerioidinae • Eristalinae, rat-tailed maggots – Spheginini •Eristalinae – Myioleptini – Helophilini – Cheilosini –Eristalini 2 September 2005 Overview of flower fly Systematics 23 2 September 2005 Overview of flower fly Systematics 24 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2 4 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2

Shiraki, 1949 Goffe 1952 Capt. E. Rivenhall, R.A.O.C. (retd.), F.R.E.S. • 21 Subfamilies (no tribes) • Historical Review • Return to TWO subfamilies • Splits off – Syrphinae & Sphixinae – Graptomyza & • Used LARVAL characters – Merapioides for new subfamilies – But Pipizines with Sphixinae!! • Reversed traditional morphoclines • Used characters for categorical ranking Gunillia, the Japanese took it to the MAX! • Hence, tribes & subtribes did not make sense So, Gunilla, a retired pilot brings sense back to Our classification ☺ 2 September 2005 Overview of flower fly Systematics 25 2 September 2005 Overview of flower fly Systematics 26

Slobodan Glumac, 1961 Wirth et alia 1965

• First of the Modern Regional Diptera Catalogs • Merged the Shannon-Hull classification into Goffe’s two subfamily system • Hence, set the future trend for family • Overlooked, but classifications • Use of Male Genitalia first time for higher classification So, Gunilla, new characters, better hypotheses ☺ 2 September 2005 Overview of flower fly Systematics 27 2 September 2005 Overview of flower fly Systematics 28

Thompson 1969 Vockeroth, 1969 • Global Revision Syrphin – Based in part on Dusek & Laska European work • Reviewed Syrphinae tribal classification • 5 tribes –Syrphini – Paragini – Toxomerini – Chrysotoxini • Paragodon – Bacchini => • Elevation of Microdontinae again • which he questioned status of • Hennig Argumentation Scheme • Zoogeographic analysis • Problem of Pipizini characterized. 2 September 2005 Overview of flower fly Systematics 29 2 September 2005 Overview of flower fly Systematics 30 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2 5 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2

Thompson, 1972 From then to now

• Revision Neotropical Milesiinae • Shatalkin 1975 • But from Global perspective – More characters, new analyses • Regional Diptera Catalogs • Hennigian argumentation schemes for tribes • Regional Diptera Manuals • Character x Taxon Data Matrix Notice, Gunilla, – Nearctic • Zoogeographic analysis that I am lost at So, Gunilla, usage sets the the ends of the – Palaearctic current “traditional” • Enlarged Earth! ☺ – Central America classification! ☺ – (Psilota, Nausigaster, Alipumilio, Merodon) 2 September 2005 Overview of flower fly Systematics 31 2 September 2005 Overview of flower fly Systematics 32

Get rid of “Descriptive,” Systematics Is a Science, Not Art

What’s the solution? 2 September 2005 Overview of flower fly Systematics 33 2 September 2005 Overview of flower fly Systematics 34

Solutions from the past Beware • History reveals a progression of • Drosophila is not a Barnacule! • More characters! From • Despite the best, more comprehensive • More Taxa! analysis done on the largest character • Resulting in better hypotheses! dataset from all sources, etc • Cladophenetics is not necessarily Science So, Gunilla and your team are the FUTURE ☺ We need molecular sequence data, BUT …

2 September 2005 Overview of flower fly Systematics 35 2 September 2005 Overview of flower fly Systematics 36 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2 6 , 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2

Challenges Diversification of flower flies happens along with diversification of Schizophora, that is, what we call the families of Calypterates and Acalypterates. Syrphidae must have arisen in the Upper Cretaceous, Check here for a more but first fossils are Eocene (Asarkina!) … complete Time [Hennigian Rank] and effectiveness of genes version of this for resolution of evolutionary charge are talk correlated. SO … more genes need to be explored. AND naturally, we need more immatures reared. 2 September 2005 Overview of flower fly Systematics 37 2 September 2005 Overview of flower fly Systematics 38

Come see us at: http://www.diptera.org We are working on Tomorrow Today!

2 September 2005 Overview of flower fly Systematics 39

, 3rd Syrphidae Symposium Presentation given at the nd to 5th September, 2005 Leiden, The Netherlands, 2 7