539 The history of dipterology at the Canadian National Collection of Insects, with special reference to the ManualofNearcticDiptera Jeffrey M. Cumming, Bradley J. Sinclair, Scott E. Brooks, James E. O’Hara, Jeffrey H. Skevington Abstract*ThehistoryofDipteraresearchatthe CanadianNationalCollectionofInsects is briefly outlined. Short biographic sketches of the coordinators of the Manual of Nearctic Diptera are given and the development of the Manual project is presented to provide background on their achievements. Lists of publications by each of the coordinators and of patronyms honouring them are provided. This Festschrift honours the remarkable contributions of the coordinators, J. Frank McAlpine, Bobbie V. Peterson, Guy E. Shewell, Herbert J.Teskey,J. RichardVockeroth,and D.Monty Wood. Re´sume´*On de´crit brie`vementl’histoirede la rechercheen dipte`rea` la Collection nationale canadienne d’insectes. On pre´sente de courtes biographies des coordonnateurs du Manuel de dipte`res ne´arctiques, ainsi que le de´veloppement du projet du manuel pour fournir de l’information sur leurs re´alisations. On fourni des listes de re´fe´rences publie´es par chaque coordinateur et des noms patronymiques en leur honneur. Ce nume´ro est en l’honneur des contributions remarquables des coordinateurs J. Frank McAlpine, Bobbie V. Peterson, GuyE. Shewell, Herbert J.Teskey,J.RichardVockeroth, et D.Monty Wood. Introduction been taken from personal accounts, archived sources housed in the Diptera Unit, and Thishistoricalreviewtreatsthedevelopment publications on the history of systematic of the Diptera collection at the Canadian researchinCanada andpublicationsfrom the National Collection of Insects, Arachnids Research Branch of AAFC, including Gibson and Nematodes (CNC) and the individuals and McSwaine (1920), McDunnough (1926), responsible for its growth and curation. The Holland (1956), Spencer (1964), Vockeroth establishmentoftheDipteraUnitbyCanada’s (1981), Anstey (1986), Cody et al. (1986), Department of Agriculture (now Agriculture Dang (1992), Riegert (1999), and Huber and and Agri-Food Canada (AAFC)) in Ottawa Cumming(2008). led to numerous accomplishments, including the conception, coordination, and publica- tion of themonumental three-volume Manual History of the Diptera Unit of Nearctic Diptera (McAlpine et al. 1981a, 1987; McAlpine and Wood 1989) (Fig. 1), The CNC had its humble beginnings in one of the most significant achievements in 1886 with the donation of a personal collec- insect systematics during the last century. tion of insects and plants by James Fletcher, The information contained in this review has thefirstDominionEntomologistandBotanist. Received26January2011. Accepted 3 March 2011. J.M. Cumming,1 S.E. Brooks, J.E. O’Hara, J.H. Skevington, Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, K.W. Neatby Building, Central Experimental Farm,960Carling Avenue,Ottawa,Ontario, CanadaKIAOC6 B.J. Sinclair, Canadian National Collection of Insects, Arachnids and Nematodes, and Ottawa Plant Laboratory(cid:2)Entomology,CanadianFoodInpectionAgency,K.W.NeatbyBuilding,CentralExperimental Farm, 960CarlingAvenue, Ottawa,Ontario,CanadaKIAOC6 1Correspondingauthor(e-mail: [email protected]). doi: 10.4039/n11-029 Can.Entomol.143:539(cid:2)577(2011) #2011EntomologicalSocietyofCanada 540 Can.Entomol. Vol. 143,2011 Fig. 1. Covers of Volumes 1(cid:2)3 oftheManual ofNearcticDiptera (photographbyJ.E. O’Hara). Fig. 2. The K.W. Neatby Building on the Central Experimental Farm in Ottawa, home of the Canadian National Collection of Insects, Arachnids and Nematodes; the Diptera Unit is located on the third floor (photograph by J.E. O’Hara). This marked the inauguration of the national Department of Agriculture. In 1919, Hewitt, biological collections in Ottawa. Fletcher was who now headed the Entomology Branch, assisted by Arthur G. Gibson, starting in formed the Division of Systematic Entomol- 1899, and by the time he was succeeded as ogyandappointedJamesH.McDunnoughas DominionEntomologistbyC.GordonHewitt the Division’s first Chief. During his 28 years in1909, theentomologycollection hadgrown of service, McDunnough and his small staff to approximately 200 drawers. The national began building one of the best insect collec- insect collection continued to grow through tions in North America, along with a large the acquisition of several private collections reference library. George P. Holland replaced and the incorporation of the insect collection McDunnough as head of the Division in of the Biological Division of the Geological 1948 and ushered in a lengthy period of Survey of Canada. By 1917, 12 steel cabinets expansion in insect taxonomy. In 1949, under with 600 drawers were formally transferred Holland’s direction, the entire collection to the Entomology Branch of the national was moved to its current location in the #2011EntomologicalSocietyofCanada Cummingetal. 541 K.W. Neatby Building on the Central Experi- Fig. 3. Charles Howard Curran, New York, circa mentalFarminOttawa(Fig.2). 1955. Through the early collection activities, a considerable number of Diptera specimens were amassed. In 1922, C. Howard Curran (Fig.3)wasappointedasthefirsttaxonomistin theSystematicEntomologyDivisiontospecia- lizeontheorder.CecilR.Twinn,whostudied black flies (Simuliidae) and mosquitoes (Culi- cidae),alsojoinedtheDepartmentofAgricul- ture in Ottawa in 1922. Curran published prolifically, producing nearly 150 taxonomic papersonnumerousfamiliesthroughoutDip- terabeforeheleftin1928forapositionatthe AmericanMuseumofNaturalHistoryinNew York. He was not replaced until 1937, when GuyE.Shewell(seebelow)(Figs.4,5,12)was putinchargeofwhathasnowbecomeknown astheDipteraUnit.Guyidentified allgroups of Diptera and published on several families. He retired in 1976, but remained in the Unit for 20 more years as an Honorary Research Associate. Arthur R. Brooks joined Guy in 1938 as a biting-fly specialist, although he alsopublishedonTachinidae.In1948,Brooks Jim had already spent the summer of 1950 was transferred to Saskatchewan, his native in the Arctic at Chesterfield Inlet with Dick province, where he specialized on various Vockeroth,collectingfliesandotherinsectsfor prairie insect groups, particularly grasshop- the Northern Insect Survey. It was in the pers (Orthoptera) and beetles (Coleoptera). Arctic, during several expeditions, that he J. Richard Vockeroth (see below) (Figs. 5, 6, developedakeeninterestinFanniidae,Musci- 8, 12, 13) officially joined the Diptera Unit in dae, and Empididae. Chillcott led the Diptera 1949, collecting all groups of Diptera for the Unit during much of the 1960s and initiated extensive Northern Insect Survey (initiated the project to revise Curran’s (1934) Families in 1947) and later publishing on numerous and Genera of North American Diptera, which familiesthroughouthisproductivecareer.Dick later developed into the Manual of Nearctic retired in 1991, but contributed broadly Diptera (McAlpine 1967; McAlpine et al. to Diptera activities at the CNC as an Hono- 1981b).HealsoledaCanadianentomological rary Research Associate until 2009. J. Frank expedition to the Himalayas in 1967, where McAlpine (see below) (Figs. 5, 7, 8, 12, 13, tragically he died of a heart attack, ending 15,17)joinedtheDipteraUnitin1950aspart hisshortbutproductivecareer(Shewell1967). of a recruitment program associated with the Some of the staff at Agriculture Canada’s Northern Insect Survey. Frank published pri- MedicalandVeterinaryEntomologyResearch marily on families of acalyptrate Diptera and on insect fossils in Canadian amber during Laboratory in Guelph were transferred his prolific career. He retired in 1985 but to Ottawa starting in 1959. The first was stayed on as an Honorary Research Asso- J.AntonyDownes(Fig.13),headoftheGuelph ciate for a few years until the publication of laboratoryatthetime,whoformedtheExperi- the last volume of the Manual of Nearctic mental Biology Section at the CNC. He Diptera (McAlpine and Wood 1989). James published extensively on Diptera behaviour, G. Chillcott (Figs. 5B, 9) joined the Diptera evolution, and ecology during his career, as UnitayearafterFrank,in1951.Asastudent well as on the taxonomyof Ceratopogonidae. #2011EntomologicalSocietyofCanada 542 Can.Entomol. Vol. 143,2011 Fig. 4. Guy Shewell. (A) In Ottawa, circa 1955. (B) At the 3rd International Congress of Dipterology in Guelph, Ontario,August1994. Antony retired from the CNC in 1978 but Chironomidae untilhisretirement in1995. D. continuedworking as a Research Associate at Monty Wood (see below) (Figs. 12, 13, 14) the Canadian Museum of Nature in Ottawa joined the Diptera Unit in 1964. Monty untilhisdeathin2003attheageof89.Bobbie studied many groups of Diptera throughout V. Peterson (Figs. 10, 12, 13) and Herbert his productive career with AAFC, but specia- J.Teskey(Figs.11,12)werealsotransferredto lized primarily on Simuliidae and Tachinidae. Ottawa in the early 1960s, after the Medical He retired in 1986, but continues an active andVeterinaryEntomologyResearchLabora- research program in the Diptera Unit as an tory in Guelph closed. Bobbie (see below) Honorary Research Associate. After a pro- joined the Diptera Unit in 1963, continuing longed absence of hiring, Art Borkent was primarilywithhisresearchonblackflies,until recruited in 1982 to study systematics of 1983, when he accepted a position in the Cecidomyiidae and Ceratopogonidae. He left United States Department of Agriculture at theUnitin1989torelocateinBritishColumbia, the National Museum of Natural History where he continues to conduct independent in Washington, D.C. Herb (see below) trans- research on Diptera. Other scientists formally ferredayearlater,in1964,publishingprimar- associated with the Unit as Research Associ- ily on Tabanidae and immature Diptera until ates or contractors were Hugh C. Huckett hisretirementin1987. (1958(cid:2)1965),whoworkedonnorthernMusci- Lewis Davies, who worked on the ecology dae (Fig. 5A), William R. Thompson (1959(cid:2) andtaxonomyofblackflies,joinedtheExperi- 1965), who worked on the Tachinidae of mental Biology Section at the CNC in 1959. Trinidad, and Kenneth A. Spencer (1967), Afterpublishing afewpapersonsimuliids,he who worked on the Agromyzidae of Canada leftin1960tocontinuehisresearchinBritain. andAlaska.FenjaBrodo,aresearchassociate Don R. Oliver (Fig. 13) joined the CNC in with the Canadian Museum of Nature in 1962 and published extensively on the taxo- Ottawa, hasworked as the de facto curatorof nomy and ecology of adult and immature TipulidaeattheCNCsincetheearly1970s. #2011EntomologicalSocietyofCanada Cummingetal. 543 Fig.5. CNCdipteristsandcolleagues.(A)InFebruary1958;lefttoright:FrankMcAlpine,GuyShewell, Hugh Huckett (Department of Agriculture Research Associate and expert on Muscidae), and Dick Vockeroth.(B)InSeptember1963;back,lefttoright:DickVockeroth,FrankMcAlpine,andGuyShewell; front,lefttoright:GeorgeC.Varley(influentialinsectecologistvisitingfromtheUniversityofOxford)and Jim Chillcott. #2011EntomologicalSocietyofCanada 544 Can.Entomol. Vol. 143,2011 Fig. 6. Dick Vockeroth. (A) Collecting flies during the Northern Insect Survey in Churchill, Manitoba, 1950. (B) On a field trip in Miyazaki Prefecture, Kyushu, Japan, in 2006, following the 6th International CongressofDipterologyinFukuoka (photograph by T. Saigusa). assistants, most notably (in chronological or- Fig. 7. FrankMcAlpine, Ottawa,1965. der)LeoForster,Mary(Roussel)Dillon,Harold Walther, Bruce Cooper, Barbara (Bissett) Eamer, and Doug Kritsch. In addition, the Unitwasextremelyfortunatetohaveemployed the artistic talents of Ralph Idema (Figs. 13, 15) and Barry Flahey (Fig. 16). Ralph pre- paredmostofthe3987figurespublishedinthe Manual of Nearctic Diptera. His drawings profoundlyincreasedtheimpactandinfluence of the Manual, and they remain among the finest scientific illustrations of insects ever published. Ralph’s figures have also been extensivelyreproducedinvariouspublications, including the Manual of Central American Diptera (Brown et al. 2009, 2010). Barry contributed over 150 illustrations to Volumes 2and3oftheManualofNearcticDiptera,and continues to illustrate Diptera and other arthropodsforresearchersattheCNC. Currently (2011) the Diptera collection is considered one of the strongest elements of the CNC and is one of the largest Diptera collections in the world, reflecting a history of keen fieldwork and curation by the staff. Approximately 2.5 million fly specimens are housed in over 5000 drawers, several alcohol TheDipteraUnitwassupportedovermuch cabinets,andabout30slidecabinets(housinga oftheperiodreviewedaboveandintotheearly totalofapproximately20000slides).Although part of the 21st century by several research mostspecimensarefromtheNearcticRegion, #2011EntomologicalSocietyofCanada Cummingetal. 545 Fig.8. FrankMcAlpine(centre)andDickVockeroth(right)withvisitingWashington,D.C.,dipteristCurt Sabrosky (left),April1963. coverage is worldwide, with impressive hold- s.l.,Lauxaniidae,Lonchaeidae,Muscidae,Myce- ingsfromtheNeotropicalRegion(particularly tophilidae s.l., Pipunculidae, Sarcophagidae, Chile, southern Brazil, and Costa Rica) and Scathophagidae, Sciomyzidae, Simuliidae, Stra- thePalaearcticRegion(especiallyScandinavia, tiomyidae, Syrphidae, Tabanidae, and Tachin- Great Britain, Austria, and Spain), as well as idae. The collection also contains the most Nepal and Uganda. In addition, the Nearctic extensiveholdingsofimmatureDipterainthe materialcontainsalargenumberofspecimens world and several hundred Upper Cretaceous collectedfromthefarnorth,includingGreen- fossil flies in Canadian amber. In addition, land (Pearyland), during the Northern Insect over 4600 primary types reside in the Diptera Survey (1947(cid:2)1961; for list of localities see collection (catalogued online at http://www. Riegert 1999) and the Arctic Insect Program nadsdiptera.org/Catalogs/CNCtypes/CNChome. (1962(cid:2)1969). Approximately 65% of the Dip- htm), in part through the acquisition of the teracollectioniscuratedtothelevelofspecies Mesnil,Reinhard,Hull,W.R.Thompson,and or species group, and more than 80% of the Garrettcollections. collection is curated to genus level. In total Current (2011) staff in the Diptera Unit nearly27000describedspeciesarerepresented consists of (in order of appointment) Jeffrey inthecollection,withhalfofthesebeingfrom Cumming(1986),JamesO’Hara(1989),Jeffrey the Nearctic Region (an inventory is available Skevington(2003),andBradleySinclair(2007), onlineathttp://www.canacoll.org/Diptera/Main/ assisted by Scott Brooks (2004), Shannon diptera.htm).FamiliesofDipterathatareparti- Henderson (2006), and Scott Kelso (2006). cularlywellrepresented includeAgromyzidae, Owen Lonsdale (2009), Collection Manager Anthomyiidae, Calliphoridae, Ceratopogoni- of the CNC, is also a Diptera specialist, and dae, Chamaemyiidae, Chironomidae, Cono- Monty Wood is an Honorary Research Asso- pidae, Culicidae, Dolichopodidae, Empididae ciate (see the Laboratory profile in the #2011EntomologicalSocietyofCanada 546 Can.Entomol. Vol. 143,2011 Fig. 9. Jim Chillcott, Ottawa,1965. Fig. 10. BobbiePeterson,Ottawa,1963. of the Diptera Unit at the time, initiated the project and had it formally authorized by the Department of Agriculture in 1966. In all, 53 2008 Bulletin of the Entomological Society of authorscontributedtotheManual(McAlpine Canada,Volume40,pp.62(cid:2)65). et al. 1981b; McAlpine 1987), with each of the coordinators authoring or co-authoring multiple chapters. The coordinators of the Manualof After Chillcott’s death the project was NearcticDiptera led successively by Shewell (1967(cid:2)1969), McAlpine (1969(cid:2)1975), Peterson (1975(cid:2)1978), One of the most remarkable accomplish- and Wood (1978(cid:2)1981). Frank McAlpine was mentsoftheDipteraUnitwastheconception appointed scientific editor of the Manual pro- and completion of the three volumes of the Manual of Nearctic Diptera (McAlpine et al. ject in 1979 and took on many of the lead 1981a, 1987; McAlpine and Wood 1989) duties necessary for the completion and pub- (now available online at http://www.esc-sec. lication of each volume (Fig. 15). The listing ca/aafcmono.html). The idea, first expressed of the coordinators for the first two volumes by Guy Shewell in the early 1960s, was to was alphabetical and not based on individual update Curran’s (1934) Families and Genera contribution, although McAlpine would have of North American Diptera by collaborating beenlistedfirstin anyevent.The coordinators with, and coordinating contributions from, all had a broad understanding of Diptera the leading dipterists of the day. The project (each having studied numerous families) and was initially driven by the need to provide a strong desire to ensure an excellent product. a more authoritative basis for identification. Preparationsincludedinvitingesteemeddipter- ThepublicationofACatalogoftheDipteraof ist Willi Hennig from Stuttgart, Germany, America North of Mexico (Stone et al. 1965) to Ottawa for 3 months in the fall of 1967 provided further impetus. Jim Chillcott, head (Fig. 12) to help lay the foundations for the #2011EntomologicalSocietyofCanada Cummingetal. 547 Fig.11. HerbTeskey.(A)Collectingfliesasastudentin1950inChurchill,Manitoba,duringtheNorthern Insect Survey. (B) At the Biting Fly Conference in Ottawain 1979, with Verne Pechuman on the left and John Burgeron the right. (C) At the Biting Fly Conference in Knoxville, Tennessee, in 1986, with Sandy Fairchildonthe leftand DickRoberts inthe centre. Manual through discussions on morpho- and a rationalized family-level classifica- logy, phylogenetic relationships, and higher tion for the order. Numerous complimentary classification of Diptera. These discussions reviews of the volumes have been published, encouraged the group in Ottawa to investigate and in 1989 the Entomological Society of homologous features across the order and America presented Frank McAlpine (Fig. 17) establishauniformmorphologicalterminology and the other coordinators with the presti- attheoutset.Thebroadestablishmentofthese gous Thomas Say Award for ‘‘Excellence homologiesnaturallycontributedtothephylo- in Systematics, Morphology or Evolution’’ in genetic interpretation of Diptera relationships recognitionoftheir‘‘magnificentsynthesis’’of published in Volume 3. theDiptera(Fig.18). Submitted chapters were edited by one or In addition to their contributions to the more of the coordinators, as well as the Manual, each coordinator published exten- technical editor,Sharon Rudnitski ofAAFC’s sively on Diptera and occasionally on other Research Program Service. All figures were orders (for a complete list of references for drawn, labelled, and organized into plates in eachcoordinatorseeAppendixA).Therespect the Diptera Unit to ensure consistency. To- commanded by the coordinators within the gether the volumes provided awell-illustrated entomological community is reflected in part andeasilyinterpretableidentificationguideto by the large number of patronyms published thefamiliesandgeneraofDipteraofAmerica in honour of their scientific achievements. A north of Mexico, using a uniform morpho- total of 164 species and genera have been logicalterminologyforadultandlarvalstages, named after the coordinators prior to this #2011EntomologicalSocietyofCanada 548 Can.Entomol. Vol. 143,2011 Fig.12. TheManualofNearcticDipteracoordinatorswithGermandipteristWilliHennigduringhisvisit totheDipteraUnitinthefallof1967.Back,lefttoright:FrankMcAlpine,HerbTeskey,andGuyShewell; front, leftto right:Monty Wood,DickVockeroth, BobbiePeterson, andWilli Hennig. Festschrift (see Appendix B; this number Manual, Frank essentially became the leader would increase by 35 if names honouring of the project just prior to publication of Jim Chillcott were included). Short biogra- the first volume. He wrote two of the im- phic accounts of all coordinators follow, in portantintroductorychapters,onecomprising alphabeticalorder. a family key to adults (McAlpine 1981b) and the other reviewing adult morphology and J. Frank McAlpine. Frank was born on terminology(McAlpine1981a).Themorphol- 25 September 1922 in Maynooth, Ontario. ogy chapter became the standard adopted by He obtained his B.S.A. from the Ontario virtually all dipterists worldwide because it Agriculture College in Guelph in 1950 and provided a uniform homology-based termi- then joined the Diptera Unit later the same nology for Diptera that was clearly laid out year. In 1954 he obtained his M.Sc. from the and consistent with that for the rest of University of Illinois in Urbana and in 1962 the insect orders. In total he published received a Ph.D. from the same university for 251 new Diptera taxa (7 family-group names, research on the evolution and phylogeny of 9 genus-group names, and 235 species-group Lonchaeidae. After Frank joined the Diptera names) (Pape and Thompson 2010). Frank Unit, he began working on the classification culminated his career with the publication of and evolution of higher Diptera, particularly his phylogenetic synthesis of Cyclorrhapha offamilies in Acalyptratae. He also published (as Muscomorpha) in the last volume of the several well-received papers on insect fossils Manual(McAlpine1989),inwhichhecladist- in Canadian amber. Frank authored or co- ically analyzed the classification of this huge authored 84 scientific publications, including lineage. He retired in 1985, and except for a 18 chapters in the Manual of Nearctic Diptera fewprojectsthathecontinuedintoretirement, (Appendix A). As scientific editor of the did not further pursue research on Diptera. #2011EntomologicalSocietyofCanada
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