Table Of ContentTHESES ZOOLOGICAE
VOLUME
20
EDITOR: RONALDFRICKE
Systematic Database ofMusca Names (Diptera)
A
catalogofnamesassociatedwiththegenus-groupnameMusca
Linnaeus, withinformationontheirclassification,distribution, and
documentation
by
CHRISTIAN THOMPSON & ADRIAN PONT
F. C.
»fe
KOENIGSTEIN
KOELTZ SCIENTIFIC BOOKS
1993
Authors'Addresses
F.ChristianThompson
SystematicEntomologyLaboratory,USDA
NHB-168SmithsonianInstitution
Washington, D.C.20560
U.S.A.
AdrianC.Pont
HopeEntomologicalCollections,UniversityMuseum
ParksRoad
OX13PW
Oxford
England
©KoeltzScientificBooks,D-6240Koenigstein
PrintedintheUnitedStatesofAmerica
Published 12.15.1993
ISBNGermany3-87429-338-6
USA1-878762-37-0
ISSN0934-8956
Contents
Introduction 2
Systematic History ofMusca 3
Earliest times to Linnaeus (1758) 3
—
From Linnaeus to Meigen (1758 1838) 10
Meigen to Stein (1838—1921) 22
TheModern Era (1921—1992) 28
The type species ofthe genusMuscaLinnaeus .... 32
The genus Musca Linnaeus 32
Workers: Theircollections and publications 33
Systematic database ofnames
Scope 38
Format 39
Cut-offdate 40
List ofabbreviations used 40
Status ofnames 43
Alphabetical listing ofnames 48
Indices
By biogeographical realm 136
By taxonomic position 141
Summary ofnomenclaturalchanges 161
Acknowledgements 164
References cited 166
Postscript 220
2 Systematic Database ofMusca Names
Introduction
ThereisnothingsodisastrousinScienceasthearrogantdogmatismwhichdespises
thePastandadmiresnothingbutthePresent.
Hoefer
Hies are importantto man. They carry diseases, sucktheblood ofman
andhisanimals, anddestroycropsandfood,butsomeareusefulaspollina-
tors, predators andparasites ofman'senemies. Inorderto dealeffectively
withthesepestsandbeneficials,manmusthaveanameforeachkindoffly.
Akindofflyiscalledaspecies,andspeciesareconcepts.Namesarecritical,
forwithoutthemcommunicationaboutconceptswouldbeimpossible.Tobe
useful, names must be unique: one and only one name for each concept.
Whilenamesforspecieshaveexistedsincethebeginningofrecordedtime,
aformalizedsystemforthesenamesisarecentinnovation.Thegradualshift
tothisnomenclaturalsystemhasleftaconfusingarrayofnames,therebeing
thesamenamefordifferentspeciesanddifferentnamesforthesamespecies.
Asacontributiontoeliminatingthisconfusion,wehavedevelopedasystem-
aticdatabaseofthenamesforflies.
Informationdoesnotbecomeobsoleteovertime.Thequalityandrelative
quantityofdatamaybediminished,butusefulinformationcanstillbefound
in oldliterature. Two examples will suffice toillustrate thispoint. Despite
its antiquity, De Geer's Mtmoires ... (1773) still contains much useful
information about flies and their biology. Vockeroth (1969: 74-76, 139)
recognized4species groups ofthegenusDasysyrphusbasedoncharacters
ofthemalegenitaliaandabdominalpattern.Inapostscript,Vockerothnoted
thathisspeciesgroupsagreedmoderatelywellwiththesubgeneraproposed
byDusekandL£ska(1967).Inadditiontoadultcharacters,DusekandLa^ka
used larval ones, which prompted Vockeroth to conclude "Further study,
especially of many currently unknown larvae, may however support the
recognitionofthesesubgenera." Atthattime,DusekandL£skaknewlarval
examples of3 ofVockeroth's 4 species groups. Unfortunately due topre-
viousauthors' disregardfortheliteratureandtheprincipleofpriority,Dusek
andL£skadidnotknowthatthelarvaofthe4thspeciesgrouphadalsobeen
described by De Geer {Musca pinastri). A quick inspection ofDe Geer's
figuresrevealsthateachspeciesgroupofVockeroth(orsubgeneraofDusek
andL£ska) iseasilyrecognizedin thelarvalstagebytheshapeandnumber
offleshyprotuberances.
A second example concerns screwworms. Some screwworms
(Cochliomyia)areendemictotheNewWorldandothers(Chrysomya)tothe
Old World. All screwworms cause great injury to livestock and humans.
Introduction 3
Hence, theintroductionofOldWorldscrewwormsintotheNewWorldand
viceversaisaphenomenonofgreatinteresttoagriculture.Theintroduction
ofOldWorldscrewwormsintotheNewWorldisgenerallythoughttobea
modernphenomenon (Gagn6, 1981, Baumgartner&Greenberg, 1984),but
it may not be so. De Geer described an Old World screwworm (Musca
erythrocephala) from Surinam more than 200 years ago, but this fact was
forgottenbysubsequentworkers(Thompson,1973).Theseworkersassumed
thatthespecimensofChrysomyamegacephalafoundinDeGeer'scollection
under the name Musca erythrocephala, while agreeing with De Geer's
description, could not have been authentic types due to the then known
distributionofthespecies.ItisnowobviousthatDeGeerdid,infact,describe
anearlierintroductionofOldWorldscrewwormsintotheNewWorld.
To access the older literature, one needs to know thatitexists and one
must have an index to it The first step in preparing such an index is
determining the meaning of the names under which the information was
recorded(thatis,developingalistofsynonyms).Ourdatabaseprovidesthis
firststepforsome2,000namesofflies.
A Systematic History ofMusca
Itisanarduoustask togivenovelty towhatisancient, authoritytowhatisnew,
interesttowhatisobsolete,lighttowhatisobscure,charmtowhatisloathsome,and
credittowhatisdubious.
Pliny:NaturalHistory
The word "fly", like the French "mouche" and German "Fliege", has
alwaysbeenageneralized,typologicalterm,andsoitwasoriginallywiththe
wordsfor"fly"inClassicalGreek("jima")andLatin("musca")whichwe
havenowcometoacceptinourscientificvocabularyastermswithaprecise
evolutionarymeaning. Thepurposeofthis chapteristotracethemetamor-
phosis and refinement of the concept "musca" from its vernacular origin
downtoitscontemporaryandverymuchmorerestrictedmeaning.
EarliesttimestoLinnaeus (1758)
Flies,andparticularlydomesticflies,havebeenman'sconstantcompan-
ionsduringtheentirecourseofhumanevolution,andbeyond.
1Forfullerdetailsoftheentomologyofthisperiod,seeBodenheimer(1928, 1929), Harpaz
(1973),Morge(1973),Beier(1973)andTuxen(1973);andforadetailedconsiderationofthe
identityoftheinsectsdiscussedbyclassicalGreekandLatinauthors,seeBeavis(1988).
4 Systematic Database ofMusca Names
The ancientcivilizations ofthe Mediterranean, ourintellectualandcul-
tural forbears, had littleprecise knowledge offlies. Withoutthe economic
resources,socialmotivation,intenseintellectualcuriosity,andopticalequip-
mentthatsubsequentlybecameavailablein 19thand20th centuryEurope,
they noted only those flies thatimpingeddirectly upon theirlives, usually
anthropomorphising them or endowing them with a numinous aura. For
example,theBible(2Kings,chapter1)reportsontheworshipofBaal-zebub
atthePhilistinetown ofEkron (inmodernIsrael). Baal-zebubmeans"lord
oftheflies", andbypunning withBaal-zebul, "lordofthedung", theJews
notonlyincludedasneeratpolytheismandtheworshipofidolsbutimplicitly
recognizedtheconnectionbetweenfliesanddung.IntheheatoftheMiddle
East, Baal-zebubwas worshippedbecausehegaveprotectionagainstflies,
andevendrovethemaway,TheancientGreeksgaveZeusandcertainother
godsthecognominanmoo&nc;(myiodes:godofflies),arcoiAmoq(apomyios:
expeller of flies) and jimaTpcx; (myiagros: hunter of flies) because these
divinitieswereabletocleartheirtemplesoftheseinsects.
During theperiodofClassicalantiquity, fromcirca 1000BC untilcirca
100 AD, theGreek "jima" andLatin "musca" were used to cover synan-
thropicfliescorrespondingtothecurrentfamiliesMuscidaeandCalliphori-
dae(Beavis, 1988).Thesewererecognizedasdistinctfromsmallergnatsand
midges,andusuallyalsofrommosquitoes,horseflies,andcattleflies(bots,
warbles) for which other terms were available: oiOTpoq, ixucoy, %covo\j/,
e|im<;,tabanus,asilus,culex,empis.Thegreatestzoologistoftheperiodwas
Aristotle (BC 384-322), whoseworkwasnotsurpassedfor2000years. He
classifiedinsectsfirstlyaccording totheirwingsandsecondlyaccording to
theirmouth-parts. He recognizedagroup ofsmallinsectswith two wings,
andalsoreportedthatthecommonhousefly{Muscadomesticd)developed
inheapedmanure.Hisaccountsofjima(e.g.inTheHistoryofAnimalsTV,
V) and those ofother authors contain a curious mixture ofsound factual
observationandpuresuperstitionandhearsay.Forexample,differentstages
ofthelife-historywereknownbuttheywerenotintegratedintoalifecycle.
Aristotlementions flies thatextrude livemaggots, yethealsorefers to the
spontaneous generation of maggots in dung and other putrefying matter,
despitethefactthat,atafarearlierperiod,Homer'sepicpoemIliadreferred
tomaggotsbeingproducedbyfliesinthecorpsesofmenslaininbattle.The
feedingbehaviourofjimawasknown: Aristotledescribedthemashaving
astrong"tongue"(%evmov)withwhichtheycoulddrawblood:hemayhave
hadthestablefly(Stomoxyscalcitrans)inmindatthispoint,orevencertain
Tabanidae. Elsewhere he classifies fima among those insects that feed
primarilyuponliquidsandjuices.
Introduction 5
In Roman times, the most significant figure was Pliny the Elder (AD
23-79)whoseHistoriaNaturalis(AD77)wasfoundedonAristotle.Thiswas
anencyclopaedic condensationofallthatwasknownatthe time, withfew
originalobservations.PlinytookupAristotle'sdistinctionbetweenflieswith
a fixedbiting proboscis andflies witha softsucking proboscis: "[muscae]
reliquorumquibusdamaculeus ...quibusdammuscis, omnibusautemhisin
oreetpro lingua" ["someoftheresthaveasting ...andsomeflies,butwith
alloftheseitisinthemouthandservesasatongue":interpretedasStomoxys
calcitrans]; and "sunthiaculei quibusdam hebetes, neque adpunctumsed
ad suctum, ut muscarum generi, in quo lingua evidensfistula esf ["with
some these stings areblunt, anddonotserve forpricking butfor suction -
forinstancewithasortoffly,inwhichthetongueisevidentlyatube":Musca
domestical.
The Medieval period, up to the close of the 15th century, was one of
intellectual regression in scientific knowledge. The temporal and spiritual
mightoftheRomanCatholicchurchwasbaseduponanauthority,tradition,
and dogma in which empirical science had no place. Progress was not
possible, nor was it desired. Whatmight have been done could not in any
casebedisseminated,sincewritingwasamonopolyofthemonasteries.The
dark cloud of ignorance and superstition was gradually dispersed by two
events:theinventionofprintingandthefallofConstantinople,therepository
ofClassicallearningforover 1100years.
Fromthisimmenselylongperiodofstagnation,afewworksandauthors
can be highlighted to illustrate how the concept of"musca" was handled.
Latin was the language oflearning, and so "musca" was usedforflies in a
vernacularratherthan a technical sense. Indeed, this continuedrightdown
to the time ofLinnaeus, with most authors using "musca" as we now use
"fly," to include flies, scorpionflies, dragonflies, caddisflies, stoneflies,
mayflies, etc.
ThePhysiologus,originatinginthe1stcenturyAD,wasthemostpopular
zoologicalbookofmedievaltimesbutistoooverlaidwithreligioussymbol-
ismtobefactuallyreliable: "muscay" thefly, isasymbolofthedevilandof
suffering. Book 12 of Origines sive Etymologiae by Isidorus of Seville
(560-636) discusses "musca" (and also "culex" "oestrus" "bibio"), and
continues thefiction ofthebeliefin spontaneous generation ofmaggots in
meat. Isidorus mentions biting flies but not house flies. De Universo by
Rhabanus Maurus (776-856) uses "muscae" to include flies, gadflies, and
probably Hippobosca. Liber de Natura Rerum by Thomas Cantipratanus
(1201-1263/93) also mentions "musca." Albertus Magnus (1193-1289), in
the lastbookofhis DeAnimalibus, discusses "musca" andasserts thatthe
6 Systematic Database ofMusca Names
fly has two wings and eight legs. It is thought that both he and Thomas
Cantipratanusmusthavehadhouseflies inmind. Petrus CandidusDecem-
brus (1399-1477) produced a Codex of animals that is one of the great
exemplarsofmedievalart.Hedealtwitha"Muscaproterva,"whichisalso
Musca domestica, and with a "Muscacypri," which is afabulous creature
addicted,—like the salamander, to living in fire ("in medio ignium impune
volant" theyflywithimpunityinthemiddleoffire).
Onebizarreaspectofmedievalman'srelationshipwithinsectswasinthe
fieldofpestcontrol.Insectswereactuallytriedbeforesecularandecclesias-
tical courts, prosecuted, and sentenced to banishment hi some cases, they
wereevenphysicallyarraignedbeforethecourtsandthenexecuted.
Amidstthismorasseofscholasticcompilation,superstition,andhearsay,
theliberatinginfluencefirstoftheRenaissanceandthenoftheReformation
stimulatedindividualstolookatinsectsafresh,tomakeoriginalobservations
withoutpreconceivednotions,andthroughagenuinelyscientificmethodto
bringaboutanadvanceinknowledge.
Acharacteristicworkofthemid16thcenturywasthatoftheEnglishman
EdwardWotton(1492-1555),publishedin 1552.Hedividedtheinsectsinto
six groups in aneo-Aristotelianfashion, usingmouth-partconformation as
his criterion. His group 4 is defined as "insectae, quaepromuscidem seu
aculeum in ore gerunt: muscae, culices, cicadae, locustae, traxales." His
chapterccxiiidealswithflies,wherehedefinestwogroups: "culex"forthe
nematocerousandbitingflies,and"musca"fortherest.
EntomologicalsciencewasconsiderablyadvancedbytheItalianUlysse
Aldrovandus (1522-1605), apioneerofpureresearch. Inbook 3 ofhis De
A
animalibusinsectis(1602)hedefinedhismajorgroups. preliminarytable
included "musca, tabanus, culex, ephemera" as a group of two-winged
insects. Inhistexthefurtherdefinedthisgroupashavingnoelytraandtwo
wings ("de anelytris, ^penn^us''). He discussed "musca" of which he
illustrated72species(pp.345-351),butalsoplacedinthesamegroupcertain
other"similar" four-wingedinsects: mayflies were groupedwith "musca,"
whilstscorpionflies, lacewings, and someruby-tailedwasps wereincluded
with "culex" His "culex" group differed from "musca" by theharderpro-
boscisandmorepainfulbite.
SimilaradvancesweremadeinEnglandbyThomasMoufet(1553-1604),
whose major work was published posthumously in Latin in 1634 (also in
EnglishinTopsel1658).Threechapters(10-12)dealwith"musca?Chapter
10 (Topsel version, 1658: 931) contains a delightful anthropomorphism
aboutthefly:"Thelight,likeTruth,hedothexceedinglyrejoycein,anddoth
Introduction 7
behave himself honestly therein and civilly". Chapter 11 "de muscorum
differentiis"isconcernedwiththedifferentkindsoffliesknowntohim. He
appearsnottodiscussthehousefly,buthis"musca"isusedinaverybroad,
almost vernacular, sense and is not confined to two-winged insects. His
illustrationsof"musca"showvariousDiptera(fleshfly,dogfly,beefly,etc),
caddisflies, scorpionflies,mayflies,parasiticHymenoptera,alderflies,drag-
onfliesanddamselflies, sawflies, andeven acockroach. His textmentions
"Muscalupus,""carnivora,""canum,""equina""bucularia,"thoughthese
are used descriptively, and also the related groups "Asilus" "Tabanus"
"Oestrus."
TheinventionofthemicroscopeinHollandattheendofthe16thcentury
openedupnewareasforinvestigation,andthefoundationofseveralacade-
mies and scientific societies encouraged dissemination of the results. The
ItalianF.Redi(1626-1698) wasthefirstentomologisttoexploitthemicro-
scope. Hedemonstratedthatinsectsdonotarisefromdecayingmatter,but
fromeggsdepositedonsuitablesubstratesbytheadultfemales(Redi,1668):
theeraofspontaneousgenerationwasfinallyover! Heillustratedtheadult,
larvaandpupaofa"Muscacerasorum"attackingcherries[Tephritidae] The
.
section ofhis work dealing with "moscas" (Redi, 1778 edition) discussed
oestridsandmosquitoes, andalsothetruefliesthatproducethemaggotsin
meatandcorpses.
Two Dutchmen, J. J. Swammerdam (1637-1680) andA. vanLeeuwen-
hoek(1632-1723)furtherexploitedthepowerofthemicroscope.Swammer-
dam (1685) classified the insects into four classes, according to their
metamorphosis, and this reflects real advances made in the acquisition of
biological knowledge. His "ordo TV, nympha-vermiformis" comprises
"musca latrinarum, asili, tabani, equina, stercoraria, acarus" The textof
hisworkdiscussesmanydifferentkindsof"musca"includingmayfliesand
scorpionflies, so it is clear that he too was using "musca" in a vernacular
sensehere. Heknewthefliesthatcomefromlarvaeinrottingmeatandthat
live as parasites in other insects, but did not mention the house fly. He
described (1685: 189, plate 11) the egg, larva, and puparium of "Musca
vulgaris latrinicolae" (in indexas: "Musca latrinicola vulgaris"), which is
clearly a species ofFannia andprobably scalaris (Fabricius) ("colorisque
est nigricantibus, sed quadam canitie temperatV). The greatmicroscopist
Leeuwenhoek (1719: 104-119) discussed "muscas domesticas" and other
"muscae."
A further advance came with the work of the Englishman John Ray
(1628-1705), who followed Swammerdam in erecting a system based on
morphological and biological facts. Ray is credited with being the first
8 Systematic Database ofMuscaNames
zoologisttorecognizethespeciesandgenusassystematicunits,definingthe
species as a collective idea for the offspring from the same or identical
progenitorsandthegenusasagroupofspeciesthatresembleeachother.His
greatwork was publishedposthumously (Ray, 1710). Flies, "muscae" are
divided into those with two wings and those with four. The section of
"muscae bipennes" (p. 266) describes 39 species. These include "musca
carnaria vulgaris" the common flesh fly (Sarcophagidae), and "musca
bombylium referens, humeris rufis, cauda albicante," abee fly (Bombylii-
dae)thatoccursinearlyspring.Onp.274thesection"muscaequadripennes"
begins,e.g."muscaquadripennis,alislongisangustisPapilionuminmodum
variegatis." A number ofparasitic "muscae" are described in the "Vespa
ichneumon" group(p.256-262).
— —
Alongside the work ofthese systembuilders the architects other
work,lessambitiousinitsaimsbutequallyworthy,wasbeingcarried—outby
the builders of the individual bricks in the edifice of knowledge the
artisans.ThesefurtherillustratejusthowbroadtheConceptof"musca"still
wasinthe 17thandearly 18thcenturies.C. Schwenckfeld(1603)described
the insects ofSilesia, illustrating aseriesofspeciesof"musca."Whilsthis
"musca" was defined as having two wings ("insectum volatile, bipenne"),
manyoftheinsectsheincludedwerefour-winged: "Muscainfectoria, eine
Schmeissfliege" (Sarcophagidae), "Musca viridis, eine grttneFliege" (Cal-
liphoridae), "Musca versicolor, eine schone bundle Fliege" (?Syrphidae),
"Musca ligniperda, eine Holzfliege/Borfliege" (Hymenoptera), "Musca
grandior oculata" and its many forms (Odonata), "Musca caudata, eine
Schwanzfliege" (Plecoptera). The Dutchman J. Goedart (1620-1668) de-
scribed 144numberedexperiments ininsectbreeding(1662a,b, c; English
translation by Lister, 1682). He used no names: the description of the
immature stages and adults were accompanied by illustrations, and he
referredsimplyto"theflyopposite"ortotheflyintheplate.Oneexperiment
(1682: 123-124,no. 130)concernedthelittlehousefly,Fanniacanicularis,
andisworthquotinginfull:
Ihadamindtotry,whatwou'dbecomeoftheputridandcorruptedUrineofaman;I
madeaFunnellofpaper,andsofoldedit,thatnoFlyorotherAnimallcou'dgetintoit:
havinginfusedintoitofttimeshumaneUrine;IfoundsomesomeWormstobebredin
thefolds,wherethefecesstayed.Ireservedoneofthem,fortheexperimentsake,which
composeditselfforchange,thefirstofMarch:andthe14th.ofthesamemoneth,ithad
theformeofaFly, depictedintheTable: itsheadwas Red,thebodyBlack, andthe
hinder-partsYellowish.
AnotherDutchman,S.Blankaart(1688)publishedadelightfullittlebook
with colouredplates andatextin Dutch, dealing with theimmature stages
ofvarious insects. No. XXm, page 81,plate 10 (oflarva, puparium, adult)
