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This is to certify that the
dissertation entitled
A.Revision of the Genus Ravinia (Diptera:
Sarcophagidae)
presented by

Gregory Alan Dahlem

has been accepted towards fulfillment
of the requirements for

Dom-ova] degree in m

 

 

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MSU Is An Affirmative Action/Equal Opportunity Institution

 

A.REVISIGN'0F'BHE1GENUS REVINIA (DIPTERA: SARODPHAGIDAE)

By

Gregory Alan Dahlem

1A DISSERTATION

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

[IKHURLOF'PHILDSOPHY

Department of Errtcmology

1989

@091193

A REVISIQI OF THE GENUS m (DIPTERA: WW)

By

Gregory Alan Dahlem

'Ihe gems mini; Robineau-Desvoidy is revised. All available
tamic, biological , and distributional data for the larvae, puparia,
and adults are incorporated. A diagmstic key and illustrations of the
male and female genitalia are presented for the identification“: of the
thirty-sevenspecies ofmmgkmwnfmtheworld. 'Ihegenera
Ward—Morav' amplacedassymmsofm.
‘Bfirty-three species are redescribed and the following species are
describedasnew: Wmm, wmm, B_,_
Mi franArgentina, and W111 fran Colorado. 13. haematodes is
placedastheseniornameofthelonespecies frantheOldWorld. B,_
W1 isredefined, removed frunprevims symnymy, andplaoedas
flieseniornameofW. Wisraisedtotheseniorname
forthespecieswhidihadbeenunderthename&_l=h_emdni__1. m

mmwmmamflwwofmma- 5..
mmmmWfimWarflmplaoedasa
waffle. R. letbiamisraisedfrnnsxmspeciesto
spxiammsmmmmwmmmm.
Wisplacedasasymymofw. 'mespecies
W.m.mmmmtflwmm
firsttimeinthegemsm.

'mis dissertation is dedicated to my mther, Shirley Marie Dahlem.

'mankywhhn, foralwayspnshingmetodomore,

andacceptingmeforthepersonIam.

iv

I would like to thank the following institutions for loans of
specinens: British Museum (Natural History) (Mil-I); Biosystematic
Research Centre (BRC); California Academy of Sciences (GS); Colorado
State University (CSU); [hymn Miseum of Natural History (DIM-I): Field
141m of Nattn'al History (PM); Museum of (imperative Zoology (MCZ);
Michigan State University OBU); Ohio State University (OSU); State
University of New York (SONY); U.S. National Museum of Natural History
(USN!) -

ImildliketoexterdmypersonalgratiufletoDr. GuyBush, who
hasopa‘xedmyeyestosomanynewpossibilitiesaboutprocessesof
evolution and speciaticn; Dr. Fred Stehr, for his help, advice, and
abilitytogetthingsdme: Dr. mintyWood, forhiserrtlmsiasmarri
mraganentflimigrnrtthisstudy: Dr. GuyShewell, forallhishelp
and advice; and Dr. Richard Merritt for all the help he has offered on
shortmticeasamrberofmycarmittee.

SpecialthanksgotoDr. RolandL. Fisdierforall ofthetimehe
lusspenttalkingwithne. Itrulyconsidermrltmdrtinediswssicnsm
insect collecting experiences, systematic theory, various entcnological

twicsmflamrentaffairsasanintegralpartofmydoctomleoqyerierne.
wecialtharflrsalsogotoDr. William L. Domes, Jr. forhis invaluable
assistamettuuiglnrtthisproject. 'Ihemalthofhisperscnalhmledge
ardexperiernettnthehassharedwithneardhisgreatenumsiasmfor
misfamilyoffliesthathasspreadtomehasbeenasoureeof
inspiration.

Iwould also liketothankmy father, Valentine, forhisassistance
wimtheiifldrgoffliefiguresardhispatienthelpoverfliecmrseofmy
life as well as my brother and best friend Andrew for always being there
tolisten. Finally, Iwould liJcetothankmywife, 'Iheresa, forher
mderstandingarflassistameduringthecairseofthisstudy. Ifully
realizethatshegaveupmanyofherdreanstohelpmegainmine.

misrevisionwasaidedbyaGrant-in-Aid ofResearehfran Sigma Xi,
the Scientific Research Society, a Grant-in-Aid of Research frcm the
mnaColl Famdatim for study at the Biosystematic Research Centre, a
grant fran the Deparment of Entarnlogy and Cbllege of Natural Science at
Michigan State University for study of European types of American species
ofmaniattenianceattheFirstInteniatiamalCongressof
Diptemlogyinaadapest,mmgaryardthe8mdgrassnamrialneseardx
Award firm the Entcnological Society of America.

vi

WOFWIS

Introduction

Nmenclatural History
Biological Review

Larval Morphology

Adult mlogy

Methods and Materials

Gets m Robineau-Desvoidy
Key to species

M (Walker)
mm (Walker)
W (10988)
W (Dodge)

' mm (Walker)
Miriam (Hall)
mm (W)
W (Wise-rd)
W (Prado and Fmseca)
W 11-813-
W (Hall)

vii

12

18

19

20

23

211

31

37

HO

1+3

H6

58

61

63

68

71

714

mm 1 (tapes)
Miami} (loves)
mm 'cta (Walker)
m m1 n.sp.
Mime—aim ' W)
m' W (van der Wulp)
mm gm ig‘ (Aldrich)
mg gum (Aldrich)
MW (Meigen)

E . . l '!l . tapes

mm germ 'eri (Robineau-Desvoidy)
W n-sp-
Mischa (Blandmd)
Wis (Hall)
Miners (Aldridi)
W (Aldrich)
mm (Dodge)
W (van der Wulp)
We (Waller)
W ('Ibwnserfl)
Wit n-sp.
m use
W (van der WHIP)
W Rdoack

viii

77

80

83

91

99

98

102

105

108

11”

117

121

122

126

129

132

137

139

1H5

153

156

159

171

175

W (van der WHIP) 177
AppendixA:Acatalog ofthem 182
Appendix B: Figures of the heads of males of several rem 'a species
Bibliography 205

ix

Figure 1:

Figure 2:

Figure 3:

Figure 4:
Figure 5:

Figure 6:

Figurea:

Figure9:

LIST OF FIGURES

Genitalia of minia ag'b_a. (a) Male fifth sternite: (b) Male
cer'ci: (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Genitalia of Ravinia advena. (a) Male fifth sternite: (b) Male
cerei (lateral view); (c) Male cerci: (d) Aedeagus: (e) Female
sternites 5, 7 and 8.
Genitalia of W.
Male cerci: (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Genitalia of W.
Genitalia of mm.
cerci: (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Gmitalia of W-
Male cerci: (c) Aedeagus: ((1) Female sternites 6, 7 and 8.
Gmitalia of W.
mle cerci: (c) Aedeagus
Gerfitalia of W
(b) Hale cerei: (c) Aedeagus
Gamitalia of W.
cerei: (c) Aedeagus: (d) Fenale sternites 6+7 arri 8 (Fenale

(a) Male fifth sternite; (b)

Farale sternites 6, 7 and 8.
(a) Male fifth sternite; (b) Male

(a) Male fifth sternite; (b)

(a) Male fifth sternite: (b)

(a) mile fifth sternite;

(a) Male fifth sternite; (b) Male

Figure 10:

Pique 11:

Figure 12:

Figure 13:

Figure 14:

Figure 15:

Figure 16:

Figure 17:

Figure 18:

Figure19:

Figure 20:

Figure 21:

redrawn fran W and Gregor, 1973).

Gmitalia of W.
Hale cerei: (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Genitalia of W1.
mle cerei: (c) Aedeagus: (d) Fanale sternites 6, 7 and 8.
Genitalia of Bavinia coad_iellensis. Male gonopod and paramere
(redrawn fran Hall, 1931) .

(a) Male fifth sternite; (b)

(a) Male fifth sternite: (b)

Genitalia of Bavinia gglunbiana. (a) Male fifth sternite; (b)
Male cerei: (c) Aedeagus: (d) Fanale sternites 6, 7 and 8.
Genitalia of W.
Male cerci: (c) Aedeagus

Genitalia of W
Male cerei: (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Genitalia of W1.
Hale cerci: (c) Aedeagus: (d) Fatale sternites 6, 7 and 8.
Gerfitalia of W.
Male cerei: (c) Aedeagus: (d) Fanale sternites 6, 7 and 8.
Genitalia Of W-
Male cerei; (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Gmitalia of Wis
(b) Male cerei; (c) Aedeagus: (d) Fanale sternites 6, 7 and 9.
Genitalia of mm.
Male cerei: (c) Aedeagus: (d) Fenale sternites 6, 7 and 8.
661112113 0f W

(a) Male fifth sternite: (b)

(a) Male fifth sternite: (b)

(a) Male fifth sternite; (b)

(a) Male fifth sternite: (b)

(a) Male fifth stemite; (b)

(a) Male fifth sternite:

(a) Male fifth sternite: (b)

(a) Male fifth stemite: (b)

Figure 22:

Figure 23:

Figure 24:

Figure 25:

Figure 26:

Figure 27:

Figure 28:

Figure 29:

Figure 30:

Figure 31:

Figure 32:

Figure 33:

Male cerci: (c) Aedeagm: (d) Female sternites 6, 7 and 8.
Gmitalia of W-
Male cerci: (c) Aedeagus: (d) Fenale sternites 6, 7 and 8.
Genitalia of W
(b) Male cerei: (c) Aedeagms: (d) Female sternites 6, 7 and 8.
Gaiitalia of W1.
Male cerci: (c) Aedeagus: (d) Fanale sternites 6, 7 and 8.
Gmitalia of W.
Male cerci: (c) Aedeagus: (d) Fanale sternites 6, 7 and 8.
Genitalia Of W-
sternite: (b) Male cerei: (c) Aedeagus (redrawn fran Hall, 1928).
Genitalia of W
Male cerci: (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Genitalia of W
Hale cerei: (c) Aedeagus: (d) Fenale sternites 6, 7 and 8.
Genitalia of W.
Male cerci: (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Genitalia of W- (10)
Dale cerci: (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Genitalia of W-
Male cerei: (c) Pedeagus: (d) Falale sternites 6, 7 and 8.
Gmitalia of W 0»
Male carci: (c) Aedeagus: (d) Fanale sternites 6, 7 and 8.
Gadtalia of W1.

(a) Male fifth sternite: (b)

(a) Male fifth sternite;

(a) Male fifth sternite: (b)

(a) Male fifth sternite: (b)

(a) tale fifth

(a) Male fifth sternite: (b)

(a) tale fifth sternite: (b)

(a) Male fifth sternite: (b)

(a) Male fifth sternite:

(a) Male fifth sternite: (b)

(a) Male fifth sternite:

(a) Male fifth sternite: (b) Male

xii

Figure 34:

Figure 35:

Figure 36:

Figure 37:

Figure 38:
Figure 39:
Figure 40:
Figure 41:
Figure 42:
Figure 43:
Figure 44:
Figure 45:
' Figure 46:
Figure 47:
Figure 48:
Figure 49:

oerci: (c) Aedeagus
Genitalia of W. (a) Male fifth sternite; (b)
Hale cerei: (c) Aedeagus: (d) Fetele sternites 6, {and 8.
Genitalia of W.
oerei: (c) Aedeagus: (d) Fanale sternites 6, 7 and 8.
Genitalia of Bavinia tancim .
Male oerci; (c) Aedeagus: (d) Fenale sternites 6, 7 and 8.
Genitalia Of W.
Male oerci; (c) Aedeagus: (d) Female sternites 6, 7 and 8.
Head of W male.
Head Of W nele.
Head of mm ' barroi Hale.
Head of W male.
Head of W male.
Head of minia domesi male.
Head of Wheat; male.
Head of W unle-
Head Of W male.
Head of mm male.
Head of W mle.
Head of W male-

(a) Male fifth sternite: (b) Male

(a) Male fifth sternite: (b)

(a) Male fifth stemite; (b)

Front and lateral view.
Front and lateral view.
Front and lateral view.
Front and lateral view.
Front and lateral view.
Front and lateral view.
Front and lateral view.
Front and lateral view.
Front view.
Front view.
Front and lateral view.
Front view.

xiii

‘Ihegemsmigiarepreeentsagrwpofveryoamenflieewhidiis
oftm foundinassociation withmaninNorth America. ‘Ihosenanbers
whose life histories have been studied are primarily ooprqmagous in a
widevarietyofmamnaliandmg. 'IhenatureoftheNeotropicnlnertbers
awaits further investigation, as very little is lawn of their biologies
at present. ‘Iheonly species frantheold World, LEM (Meigen),
hasbeenmtedmnmemisoocasimsasbeingaveryoannonsynanflimpic
species in the Palearctic arri Oriental regions. This revision provides a
keythatwillseparatemalesarrimstfenalesofthisgemstospeciee,
navsynmymy, ardpills togetherthewidely seattered information onthe
genus, arablingassociatimofmidiofthepastreseardiwithwrrent
names.

Inrecentyearstherehasbeenmmmterestintheirsectfamiasof
dung,e$eciallyoowd1mg. mismteresthasbeenspawnedbythespread
oftheintromcedfaoefly,WDeGeer,acrossomtirental
Northbnericaardbyirwestigatiusomcemingtheintrecmoedmmfly,
W(W). Bothoftheseintroduoedspeciesare
inportantpestsofcattleanddairyoperationsardbothbreedinoad

mature. wafflemrkmrrentlybeirgdmeinNorthAmericamthe
faunas of cow dung involves searching for native parasitoids and
predators that may be meful in the biological control of these pest
species. hittrnsepestsarerecentinvadersofthedmrginsectcouplex.
Studies, surfinasthoseby'hmaer, m (1968) and'manasawi'ngo
(1968) , indicate that the predator/parasitoid carplex of species famd in
cavdnrgismidimre mitedanipronetocontrolthenative flies found
breedinginthedung, mainlymembersoftheflesh flygemsmvm 'a. In
order to better understand the possible predator/parasitoid relationships
wl'xioh nay have inplicatims on the control of face fly and horn fly, more
mfornatim needs to be acanmlated on the natural associations of
predator/parasitoid carplexes with m, and precise identifications
of the species involved are essential to allow repeatability of the
finiirqsfrunsmzhsuriies. aierecentstudyindicatesthatspecies
within the m may be inportant biological control agents against the
face and horn fly (Pickers, 1981) .
Anotherinportantareaofcmcernisflyproductimswreesinurban
calamities. IthasbemmtedinstmliesaxhasthosebySdioofgLaL
(1954) and Silverly & Sdioof (1955a, 1955b, 1955c) that dog dtmg is one
of the most inportant preductim media of nuscoid flies in metropolitan
areas. Fleshfliesarethemstinportantdecmposersofdogdm'gfard
in these alviruwrts and, although the previously mentimed authors were
unbletoidentifyorfiniexpertstoidentifythesareqmagidsthey
collected, species of the genus Minis are, most probably, the most

importantspecies involved. Collecting offlinjgbymyself, andmany
years of collectingbyDr. William L. Domes, Jr., has shownthatmembers
ofthisgemsdomtmrmllyoccirinsimatimsmidiwufldmakethem
pestsofmnnans(su:hasinlmses,mpordxes,oratpimics). This
carrntbesaidforotherspecies,sud1asthetnxsefly,whid1havebeen
rearedfrundogdung. Itis, therefore, inportanttolmowmreabout
thesedecarposersofdogfecesarumiwrtmsinordertopruwtethose
qaeciesofmin'gwhididotheessentialserviceofbrealdmdamthis
to aid: studies is the proper identification of the species involved.
Identificatims of undetermired Sarcqmagidae fran various large,
instimtimal collectiors indicate that one-fourth to one-fifth of all
sar'corhagids collectedinNorth America belaigtothegemsmilia. 'Ihe
lastcmprrehensive revision ofthisgemswasprovidedbyI-Iall (1928),
hrtmstofthenamsusedinthisrevisialarysudyhavesubseguently
beenplacedassynmyms. Seventy-nineweciesnameshavebeenarpliedto
whereofmisgerus,ofwhid1thirty-sevenarewnentlyrecognized.

WW

'me germs m was established by Robineau-Desvoidy (1863) for

his new species, m, m, B. Mata, and Meigen's (1826)
‘ with the latter designated as the type species. He

 

mmmmmsmmbmmfiy.u-_W
Rhineau-Desvoidy, ard Lmrtiggla Rabineau-Desvoidy, as syncnyns of 3.
mabigm).h¢didmtm\flemnmimsfimbimau-
Desvoidyinthisnewgems. Bottd1er(1913)includedmin°gasa
Woffiamardimluiedfliespeciesfigmneigaiasa
rather. NoNewWorldspecieswerecasideredasnalbersofthem
grcupuntil Parker (1914) redescribedthegemis museum
mmlettardhismweciesL—gmsselam
mammthegenus. Aldrich (1916) suggestedthatm
cmldbeplacedasasubgenusof Sam, included Parker’s (1914)
species, “Williams-1m. www.m-es
(asavariantofWParker),ard§._glflu_sasmmbemoffl1is
subgenus.
PhileParkerarriAldridiweremsydescribingtheNorthAnerican
fauna, C. H. T.‘I‘ownse1flwasworkingonthefam1aof$mthAnerica. In
BlZ,deesaibedflegamWamplacedmem
speciesELWasthetypeofthegems. In1917,'Ibwnsend
rmedseveralgereramfispeciesmataremrrentlycmsideredwithin
m. HedescribedthegenlsmmwithWParkerasme
www.mmmwithummmmmasuetype
mammammswiflafiflrimcoqufllettasflntype
specialrwwiflzssmmdasfletypemiw.
“WwithhisnavspecieSMasthetypespecies.
Inl927,1bwsaddescribedthegeniszithhisnewspecies

WEWWSPGCRS-

Hall (1928) revisedtheNewWorld wecies ofmmtcmsidered
Wasaalbgemscffiamardsymrymizedmfi'sgenera
W.W.W.W.W,
“Wwithmism. Heredescribed'lmiserd’sspecies
Emitsssasrim. MW. HeraisedAldridi's
Wm. mmmies level becauseGreene (1925)
fornrithespeciestobedistinctfranR.canmmisParkerintheinneture
stages- mwmwmmmswm
because of correspa'derce with Tinberlake. Tinberlake found only one
species of sareqi‘iagid which agreed with 'Ihmsm's description frdn
Hawaii. Halldidmtexamine'nnnsm’stype. Healsoincludedhisnew

species 51m. mm. $._orsitsijs. M. s.
W.MLW.LM. amssreuiegriinthis

subgerus. Hall (1929)d1angedthenameof§_._m__ir_mato§._m.ata,
since the name m was preoccupied by an earlier Robineau-Desvoidy
name.

Errierlein (1928) caused many rmemlatural problems thmrghcut the
Sarcqingidae,inchflirgthegereraardspeciesnuvcmsideredinthe
m. PlecasideredflegermWRbimau—Desvoidyasa
Mofmammmedflemiwsmneigm.s.
Whig, Withdrawn. W Fabricius. $.-
Wmdermmmuisvandermmarflsrnlm
Wiedarannasnarbersofthegaus. Ofthesespecies,mlythetype

species, 3W, isairrentlycaeideredinthem. Healso
Wmmmmmmwmmmneau-
Desvoidy and gamma 'Ibwnseni with Biggie Robineau-Desvoidy,
witturtanydiscussion. Heestablishedthegerustori
W van der Wulp. Townsend (1938) later found that Enderlein's g.
m was actually a misidentification of W.
Bflerlein also established the germs W for Mist;
Fabricius which was shown by lnndbeck (1927) to be a senior name for S;
W Meigen. Hall (1937) notes that Lam Hall was placed
in the gems Dim by therlein (1928) as an undescribed species.
Until 1930, the names applied to War World species of sarcofiiagids
by the early authors such as Robineau-Desvoidy, Hacguart, Walker,
'Ihcnscn, arrivanderWulp, wereignored forthemstpart, sincetheir
descriptionsweremtspecifica‘nglitoseparateonespeciesfrun
amtherandthetypesweregenerally inaccessabletoAmerican authors.
W, in 1929, the Unitai States National Husann sent J. M. Aldrich to
visitseveraloftheinportantniropeanmsamsforthepurposeof
lemminingtypesofAmericanDiptera. Inthecmrseoftheworkhewas
abletoseemostofthetypesofAnericanm. Hereportedhis
fit-dings m m in 1930 and established the following synonymy:
WW Robfleau-Dasvoidy = W Parker: stanzas
themtmmtmmti
mm:mm:wm:mm: and
Wm. MW=W

marmmm3amsrmvanderme si
mm'miflalhfiiflmmdflmwwi
mmdermlp- mmderme=semudridn
ammderMP=s£imardiimllo seesipisvanderwulp
- Wag Parker. He also fund that his 1916 descriptim of s;
mmmmmmderWulp. He
redescribedfiMWalkerarflplaceditinflaeWgrupalumg
wiflmerIixittatsMacqmt-

Frunthe1930'stothel950'sanibeyu‘d,thereappearstobea
split between the generic runenclature used by North American authors and
SthhAmerican authors. 'lheNorth Americarswereheavily influencedby
Aldridi's (1916)majormrkarrittmscu1sideredmin'gasaaibgemsof
m. 'n'ieSurthAnericanswereheavilyinfluencedby'Iulnserd,
especially his works included in his [Jewels of Myiology (1934-38), and
utilizednanyofhissmallergererathatareherecusideredsymnymuis
withm.

Hall (1931) describedtwonewspecies intheNorth American fauna in
mmmwsmsiessmm Beel(1931)
describedfimmaolivia. Prado&Fuseca(l932) described
Wmmn. Blanchard (1939) describedmmrayim
m1 frunArgentim. Intheold World, Imribeck (1927) examined
Fabricim' (1794) types of W and placed W Heigen
asasyrniymofthisoldernane. Seguy(1935)described§_,_azgafrun

Libyaard, in1938, describedmigigjisjmmm.

waver,in1939,Seguyplacedthesewoweciesassymrynsof$_.
mwabriciusb

Lmes(194la)placed§;_b§1m1Prado&chsecainthegaus
mmmdisuissedthefatale genitalia. Irpes (1946a) considered
Wammmsmplwimmymm
Mp,&mm,mgmmasmcruus
gaus,aswellashismspeciesg_._m. Healsoredesczribedgg
mmderwtflp.rawveditfrmsymmywims._sum1s§
Walker,atriplaceditintheggae_tgrg_v_igig. Inthissamepaperhe
”Bi-WW Robineau-Desvoidy, M Hall, m
vanderfiflp,ams_,MMdridaasmrsofm,ard
redescribedmvanderwulp,rmavingitfrunsyralymywithfi.
m. Icpe(1946b)describedg,_g1mig_a_iasamvspeciesinthe
WW-

Rdaack (1952) desaibedthemweciesWard, in
1954, consideredm as a valid gemswith the following synonymy:
WW.WW.WW.
WW.WW.WW.
WW,arfifigelmam1ein. Heredescribedthe
gamardincltfledthefollowing speciesasnmbers: mien,
Wm.srassmm.smm1.hw
W,§AMW,MW&W,L
www.mnm.mmmrs_erm
m.S-_emhrdavandermlp.s._flsrmuA1dridhs.mes

ll

Aldrich, m1 P311. M Parker, W Robineau-
mm.mmmmm.WMI.ssums
Hall, fiMMdridi, mama, mama
Newsweemmmssstmwaneni
aggvandermlp,migmmback, Qt, trivimta Townsend, and_S_.,
mmdermlp-
mdge(1956a)corsideredgammiaasavalidgermsardimluded
thefollowing U.S. speciesasmalbers: WW,§4
WmdeIWulp,_S_.MWalker,§.assiduaWalker,B_.
mmrfiemmLWmmem
speciesgm. FruncorresporflamewithDr.vanErdenatthe
BritishhzsunnofNatmralHistormnodgefunrlthatthetypeofg.
Mmcloselyreearbledthespecieskmparkerandg‘
mmderMpflnnfliguggi—ammfllett.arfl.flmfore.
mmmmmmmnmmmtdmctemcufld
befunflinfliefanalesttetwmldseparategmwfi.
Wfrmuieanother. HeplacedmjggWalker asthesenior
nmnefornrssdrimooquillett- IntheIflSMinis.Doder-=
sepamtedvtathadbemhmnasfiWiRobfieau-Desvoidyhrto
mm6.&mlsm).&lbsmmeri(mbfieau-Deswidy)
“WW)withmefollowimsymnymy= W=fir
MW:MW:WW:NW
m- W=mm:mm:m
Wrardmmflmsm- m=mmm<en

10

InfluissqaaratimtementiusttntflrespeciesLngrginiefiarpeared
tobeacmplexofspecies. Healsosymmizedmyglisnallwith
mutidiammacedWasamerofflemig.
Dodge(19583)describedthenavspeciesmifrmolbaarflprevided
adescriptiui ofthefanale ofnglgnngs.

tapes (1962) reviewedthegemsmmarddescribedh
WNWEMMWmds
mmmmmmia.

Dowms (1965) placed Chaetomm" asasubgenusof minis. He
miwmtmmmmwably=miguammflet
BM (AMI-"1611) probably = W (Aldrich). HE also placed §_-
Wiellasasymwmofmfmmldfidx).

nodge(1968)describeduaenevspecies_c._g§gpd_afmnoentra1
America.

Iqaes (1969) separatedthegemsminiaintothefollowimmree
ceremwimimimtedsyrmymw Mis=amasammerm
WW.WW.wWW:
mm=mmaaeriemmm=em
W- mmmmmmmmm
Hacmartasmalbersofthegausminiaarflmentiusfliatthemeofg:
W by Hattos (1919) was a misidentificatim of the species B,
31:93:1(mfimb mwmmmplwm
mwawofgmmm-

Lmes(l975a)describedthespeciesfi,_tgiMs_ifrmestaRica.

11

limes (1976) redescribedWWalker, placed it inthemm
mplwmmmmmmww-
ropes (1977) examinedtheholotype «WW, redescribed
thespeciesarriplaceditinthemin'g. Healsoenraminedthe
mlmofwmm.mmmmflmid_us
Walker. I-leraisedthospeciesstamswitlfinthe
Wardplacedsrasids.mi&.arfls_latim
Parkerassynonyms. Wenhedidnotdisuisstherelatiorshipbetween
emerge (vandermp) New laps (1978b) med
mfmfleMimsmflerditwiflimms
Dodge. Hemtesthatheorigimllyplacedthisspeciesinthefiayifion
the basis of Dodge's (1965a) figures of the fanale genitalia.

'meplacanentofWHarrisastheseniornaneoverthe
lu'ie Palearctic species W (Fabricius) , sluvly worked its
wayintctheliterature. Dodge8Seago(l954)refertomasthe
seniornameasdoesDownes (1965). Iopes,g_§l_,_ (1977),however, lists
Wasaquestiomblesymnymofitriata. 'menamemscagtriag
Fabriciusispreocuipiedbyenelin, 1790. NoHarristypeshavesurvived
toflrepresent,yetautlnrshadinterpretedtheoriginaldescriptimas
that of lame Palearctic m. Pepe (1986) examinedthetypematerial
offillmarflfunflfliatmmigansmnmwasajmior
mofm- Wknmiswiedw
Villers (1789).

mmismvisimoffleMm.mammardmmus

12

areplacedassyrurynsofmmia. 'Ihirty-threespeciesareredescribed
andthefollowingqaeciesaredescribedasrav:3,_g§§ellifrun
W.Ldzmifmm,mfmmgartm,am&
mmmlorado. Wbplacedasfllesmicrnameof
the lame species fruntheold World. am is redefined,
rmedfrmpreviuissyruiymy,andplacedastheseniornaneof&
MafterexaminatimoftheholotypeinthemseeNatimal
D’HistoireNaturelleinParis. Wismisedtotheseniornanefor
flespeciesWiQhadbeenmderthenameWaftershfiyof
theWaJJcerarrivanderWulptypesattheBritishmsamofNatural
History- Wmmmmwedasm
«W. argivivaisrerwvedfrunsynonymywithgrm
ardisplacedasasynmymofm. Wisraisedfm
abspeciestospeciesstamsarflasmfigmismvmized
tritium. Llealseiisplacedasasynmymofa-M. The
Species WW. Lauri. ands-m are included for
firefirsttimeinthegermm. 'l'heholotypeofm
'mmmnacquartwasalsosufliedarfiitwasfumtobemta
Worm.

ApperdixAprovidesacatalog—style listingofnunenclaturearri
syratymyadoptedforthisrevisim.

1310mm m

13

Rich of the biologin informatiui recorded in the literature deals
withrearirgspecies oftheBayinia frunvariusmsdia. Rsaringdata
described in the literature may be divided into two categories, rearings
frmmhnalnediatakeninfrunthefieldardrearingsaccurplishedin
thelaboratorybyplacingthefirst—instarsuiadrosenmedia. Both
types of investigatims can provide valuable informatim m the life
histories of these flies. The folluring biological reviav uses the
taunenclature adopted in this study. To associate literature references
with the names given below, corsult the individual species descriptions .

mrnivore, umivore, and herbivore durg is rurtinely utilized by
MoffllemMiQs- W: W. New
have been reared frun dog fews (Dodge, 1956a; Wilton, 1962: Coffey,
1966: personal observations). 3% has been reared frunwolf
duty collected at a zoo in Lansing, Michigan (persmal observatims) . B;
m. mm. 3m. W 81.23231. firmness. and
WMbemrearedmmmanment (Howard, 1900: Aldrich,
1916: Kirdiberg, 1954: Dodge, 1956a: Mihalyi, 1965; Coffey, 1966: Kano,
Field and Shira'aga, 1967: lopes and Kano, 1968: Sydisvskaya, 1970;
Iqaes, 1974). Cowmanure isthemostcummly studiedbreedirgnedia,
ardfleSPecieSW. RM. RM. ELM: E.
21mins. midis. EM. and W have been reared
frun it (I'bdard, 1901: Greene, 1925: niplirg, 1936: m, 1943; mffey,
1966: Saniers and Dobsm, 1966: 'mrner, My 1968: Dunes and Wingo,
1968: Valiela, 1969: Blume, 1970: 'lhulas and Morgan, 1972: Valiela, 1974;

lu

Merritt ard Andersu'l, 1977: Davis ard Turner, 1978: Phartm ard than,
1979: Pickers, 1981). m ard 3m have been reared frun
sheep dung (Wilsm, 1932: Coffey, 1966). Laurie ard BM have
been reared from horse marnlre (Coffey, 1966). W has also been
reared a: pig fees (befey, 1966) .

Smespeciesofgggim'ahavebeenreportedasparasitoids ofother
insectsardotherinvertebrates. milesuneoftheserecordsare
probably based on misidentifications, others appear to have suns
validity. ;B;_e;:§bgng§ has been reared.fromltheIMeXican.bsan beetle,
.Eeilashnaryarixsstisinnlsant (Cbledptera: Cbccinellidae) (van Enden.
1950). Ea_anria has been reared fndm.Lasnnsstsrnarhsrni Smith
(Cbleoptera: Scarabaeidae) (van Eden, 1950) ard has been fund to be a
predatorofeggsorlarvaeoftheface fly, Wmceer
(Pickens, 1981) . B. haemaw has been reared frun mm
L. (Colecptera: Scarabaeidae) (W, 1943: van Enden, 1950), as well
as frun grasshoppers ard snails (Seguy, 1932, 1941: Rodetdorf ard Verves,
1978).

Severalspecieshavebeenrearedinthelaboratorybyvariuls
researchers. Winsbeenreared frunmeat (Dodge, 1956a). B_.
m has been reared m deculposirr; hanhlrger (Knipling, 1936). B;
saris. Ea_nus191a. and Br_stiaulans have been reared on pork liver
(Sanjean, 1957). W has been inplicnted in humn mind ard
intestinal myiasis (Jones, 1947: Trofimv My 1958: W. 1965:
Roldandorf ard Verves, 1978) .

15

Suns of the data m collectim of species of mini; also provides
valuable informtiul m the life histories of these species. B_.
W (Hall) wascollected intrapswhichwerebaitedwith
decaying liver, urea, ard water (Hall, 1931). W, R_,_
W. W. Bemsiela. mam (pmbably=8.
Ma) arerecordsdasscavengermsciesmpigcarionbyPaynesrxing
(1972) . am has been collected in traps baited with decaying
beef (Hall, 1931) . 3. helm usi Lopes was collected on variuls flowers
in Costa Rica (Lopes, 1975a). R. we is noted frun onion fields
(Merrill & film, 1953), as being attracted to dog dung (Coffey, 1966) ,
ardcollectedfrunatrapbaitedwithaharemss inAlaska (Gill,
1955) . m is noted as being attracted to pig dung (Coffey, 1966) .
Wists. More. LllrM'sieri. Bans-isle. and M were
collected in traps baited with chicken entrails ard, suretimes, lumen
feess (Dodge8Seago, 1954). Wismtedasbeingattractedto
Inmlan dmlg (Coffey, 1966). Inndbeck (1927) notes that W is
cunminsardyplacesardisoftenseenonmmanexcreuent. B;
Whasbeenreportedtobeattractedtofeces, meatardfruits
(Fberhardt, 1955: Gregor ard Povolny, 1961) ard has been captured in
food mrkets in ampst, Hlmgary (Aradi ard Hihalyi, 1971). am
is attracted to hmnan, sheep, pig ard dog dlmg (Coffey, 1966) .

Sure informatiul on fanale feundity is present in the literature.
Knipling (1936) fund that fulr farales of mum yielded 20, 24,
28, ard 30 larvae, respectively. He notes that 18, 32, and 40 larvae

16

weresqnezedfrunthreefanalesoffiMars. ForWhefund
20, 27, 29, 32, ard 4O larvae in five gravid ferales. He collected 15

larvae frun a female of W. He also fund that five fanales
of M produced 21, 24, 25, 4o, ard 44 larvae. Zakharova (1966)
dissected34 femalesofhmardmtesthemininmfeundityto
be16 larvaeardthenaximmfeunditytobesz larvae. Iarvalculnts
canbemisleading ifobtainedfrunpinned specimens, asthefanales of
ministerdtoexpellmnyoftheirlarvaeastheydie (personal
observations).

Information (I: the length of the life-cyles of species of this germs
is limited. Khipling (1936) notes that the life cycle, frun first-instar
toadult, naybelessthanll days formugz. Healsonotesthe
developmrtalperiodofWasbefigMdaysammas
being 13 days. Times & Lbrgan (1972) fund that the time of pupation of
thehomfly, Liam, coimidedwithttetineofpmationof&
Wardusedgmasanirdicatorspeciestotellwhenflremm
flies were pupating.

Anmberofinsectparasitoidshavebeenrearedfrunthepdparia of
bovine dung-breeding Baginia (Blickle, 1961: Turner, M, 1968: 'Ihunas
ard Wirm, 1968: Hayes ard mm, 1971: more ard Isgner, 1973: Watts
ard Curbs, 1975: Wharton, 1979: F199, M“ 1983a: F199, $31“
198319). ‘mese include: WW Graverhorst: MM
(1.), and W 89- (Colecp’cera: Staplnlinidae): W
Saar. W (Say). “from 89- W:

17

W): Winnie (Say) and 21.91% 89- (W:
Qmipidae): W and Emile Sp. (1W: Daccilidae):
W 89- W: DiaPridac): Wis (Minced).
WGmult and Sanders. rigidities spa Meme
W Ashnead, W Latreille, ard S. m enea Curtis
(Hynencptera: Ptermalidae). In additiu'l to insect parasitoids, R_. anxia
isalsohmntobeattadcedbythensnatodewmm
Nickle (Stoffolano ard Streans, 1971: Stoffolano, 1973).

W. 932.31.; (1973) fund that W
W: Shocidae) provisiuled its nests with W aid B,_
and; adults, along with a variety of other cyclorrharhan flies. ally
Isles were used as provisions. Wharton (1979) noted that W sp.
(Cblecptera: Staphylinidae) adults captured early instars of W.
Ants are inportant natural enemies of ding—breeding Diptera, althuigh
published studies have not dealt with M specifically (Pimental,
1955: Wingo, My, 1974: mart-on, 1979).

Cbok ard Gerhardt (1977) looked at selective mrtality of insects in
mamre frun cattle fed Rama“ and DimilinR and fund that
populatiu'l levels of renters of the gems Begin]: were significantly
reduced frun Rahal, but not Dimilin.

Very little has been written (:1 male station taking ard mating
behavior of the m. Dodge & Seago (1954) fund lnlusually high
ratiosofnalestofenalesinBngipiefiardWintheir
oollectiu'ls of sarorphagids frunnnmtainummits inGeorgia. 'nleywere

18

unable to explain this phenmena as a mating behavior, since they ally
ccflectedaemtingpaircfmawmofLmsmnfisridurim
their entire stldy. Olapnan (1954) collected 41 males ard ally 1 female
ofnm inhissudyof umit-frequentinginsects inmntana.
HatitlgbehaviorofBLmLodes (EM) isdescribedbyHanmer
(1942) withthemerestatanentthattheculplespuruleeadlotherinto
theairbeforetnmblingtothegrund. LopesardKano (1968) providean
examination of the relationships of the variuls parts of the male ard
fenale genitalia of B. gforti which were killed during copulation. A
detailedaccunrtofthematingbehaviorofgm ispresented
later, inder the biology section of this species’ description.
alservatims of potential station-site locations are given in the species
descriptiui, when available.

immune!

'lheSaroophagidaearenormally ovoviviparuls, that is, thesggsare
'retainedwithinfllebipuldleduterusfordevelqnentatdfirstilstars
aredeposited. 'Ihelarvaearestill alclosedintheirdlorionic
Mbranesdmdepositedardreleaseflmselvesbyclawimvigorulsly
withtheirmrthhooks (Sanjean, 1957). 11115 inportant facet of the life
historyofsarooriagidsalloustheqlporundtytoswythefirstirstars
ofthesefliesbydissectirqadead,pirmedspecinenofafanaleard
wtainingthelarvaeheldwithintheutems. 'Ihelarvaepassthrulgh

19

tm'eeinstarsbeforewpariatim. Firstinstaxsdomthavedevelcped
anterior spiracles, seoadinstarshavemlytmoslits intheposterior
spiracles milethirdinstarshavethreeslits.

Adetailedsuflyofthelarvaeofmg'gwaswtsidemescope
ofthisrevisim,h1tp.1blishedaooamtsofthelaxvaehavebeen
inoorporated. 1011le (1936) studied the first instar of R. giola, B;
m.m(mly=WwBM).B-_m
“MEE- Sanjean (1957) shxliedthefirst, second, andthird
instarsof B,gs_iola,3._m¥andR. stinulans. 'Ihelarvae of};
WMbeendescribedbyKano8Sato (1951). Lopes (1982)
describedmwmg. almeidai first instars. Greene (1925)
mmmmhofW.&_llmihie£i.amW.
Whartm anduocn (1979) described thepuparia of W. 3;
M.W.W.W,MW.

mmmmmmrmmmmtan
extrenestateofthelabnminwhidzitappeaxsahollymatbrarmsani
haveverywell developed psaxiotrachea, which are often darkly pigmented
(Domes, 19553pexsmaldasetvations). 'Ihewell developedpsaadctradmea
alsoooansinseveraloopmghagmsspeciesofthegemsmani
myherelatedtothedm—feedimmdeof life. 'Iheirfunctimis
mflmownatthepresenttine.

WWW

20

'megemsminiamsdescribedbynobineau-Desvoidy (1863) andwas
mantedfranothergenerabythedmamcterofthefrartalbristles
diverging very little anteriorly. 'lhis character is generally true in
them,mtitcamiotbeusedtosepaxatethegetusfrmcther
saroqmagidgenerareoognizedtoday. mltsofthegerusmmngare
separatedfrunallcmersbythepresenoeofthefollwingdiaractezs:
postalarwall sem1ate:tegulaeorangish ingroundoolor, sanetimes
partly infuscated; male without apical saltellar bristles and with
reddish-orange genital capsule; fanale with frontal vitta at midpoint
miceormrewidthofparafzmtalplateatthatlevel. mu;
n.sp.isthemlyspeciesthatdoesnotoonfomtoalloftheseexternal
characters, havingablack tegulaandblack genitalia. 'Ihefom ofthe
male and female genitalia, tum-Iver, definitely associate this species
withothersinthegemsm.

'Iheterminologyused inthis revision isthat of Mpine, fig]:
(1981) exoeptforduaractersoftheaedeagm, werethetemimlogy of
lunack(1954)hasbeenused. 'nxemaleaedeagushasproventobethe
singlemstinportanttoolusedtomstseparatespeciesinthisgenzs
(andnearlyallothergeneraofSaroomaginaeaswell). Ihave
agressively searched for non-genital ic features of the adults that are
useful in the separatim of species and are nentiamedwhere applicable.

WWW

21

Nearly 12,000 specimwereaaminedfzmnmeruislccalitiesin
North, South,anica1tralAmerica. 'mewecimenswerefrunvarious
institutims, thecollecticn of Dr. W. L. Downes, Jr. (WID), andmy
personal collectim (GAD) . Damimtials and dissections of specimens
weremzmllymidertakenatamagnificatimof50Xwiththeusecfa
leitz Wetzlar dissecting microsccpe. Illustrating of the genitalia were
accomplished withtheaidofaBausdxandImbMicrcpmjector.

Dissecticns of the male and fanale genitalia were accomplished
usingthefollcwingteclmique:

Step1:l-leatupasolutimofmi(7pelletsm-It010ccH20)in
awaterbathqutjustbelwboiling. 'Ihisusuallytakesabwtzrnms.

Step2: Carefullyrawvetheentireabdanenofthespecimentobe
sufliedbygraspirgitgentlywifliforcepsardmdcirgflieabdmenupand
dammitilitbreaksofffrunthethorax.

Step3: Placetheabdaneninthelflisolutimforslninrtes.
miletheabdmenisinthesolutim,placedistilledflzoin3
depressicrs in a culture plate.

Step4:RencvetheabdarenfmthelOisolutimandplaceitin
are of thedepreasims filled with distilled H20. Unlerthe scqae, make

t“handSt‘hstemites(males)crst‘hanth‘h

anincisicnbetwemthell
sternites (fenales). 'Ihe article should be very pliable at this point,
ifmt,thelflisolutimshculdbelefttoheatdpfcrancthertn1rcr
”(gravidfenalesnayneedtosoakabitlaigerthanctherspecinens).

Step 5: Dissect off the genitalia and place in adjacent fresh 820.

22

Step6:Usinganeyedmppercrpipette,rinsetheabdxm's
interiorardexteriormrtilfmeoftxadaeaardcmermntedranairs.
DiscardoldandresupplyfreshlEOatleastZBtimes. Besuretocheck
forlaxvaewl'mdissectimfarales.

Step7:PlaceabdmIeninadjacentfreshwater. Retnrntoworkon
gaiitaliaarddissectaxtpartsthatmeddetailedexaminatim.

Step8: Placeabdcmeninavial of glacial acetic acid for (21.1
minrte. Renoveabdanenardplacemapapertowel,withthebasal
cpeningoftheabdanendcwn,todry. 'Ihiswillrestoretheabdanen's
pruinosityandtessellaticn pattern. If itdcesnctrestore, it maybe
time to change the 101 and/or acetic acid solutions.

Step9:Placeasma11ammtofgluemthethoraxwherethe
abdanenistobereplaced,usingaminztennnmtedmamatd1stick.
Underthe scope, replacetheabdanento its original positicn.

Stqn 10: Place genitalia in a depression filled with glycerine.
Swirlamxdmttilmstofthewaterismved. Placegenitaliaina
micrcvialardattadmtopinofspechmtomiditheybelmg.

Gems RAVINIA Riaineau-Desvoidy

m Rbineau-Desvoidy, 1863: 434. Type species: W
W Meigen (orig. des.).

W 'nownserri, 1915: 408-409. Type species: W
W 'Ibmsend (orig. des.).

W W, 1917: 190, 193, 195, 196. 'IYpe species:
W W (orig- 686-)-

W W, 1917: 190, 193, 195. 'IYpe species: {giccbia
W Coquillett (orig- des.) = mists (Waller)-

mjnig 'Ibwnsend, 1917: 191, 194, 195. 'Iype species: W
Parker (orig. des.) =M§ (Walker).

W W, 1917: 191, 194, 195. Type species: m
213mm Aldrich (orig. des.).

W Townsend, 1917: 191, 194, 195. Type species: m
m 'Downsend (orig. des.).

W IUmserxi, 1927: 220, 295. Type species: W
21mm (orig- dos.) =m Waller)-

Rgglim Ehderlein, 1928: 16. Type species: W
Enderlein, nec. van der mlp (orig. des.) = misida'rtificatim of
mm (W).

W Enderlein, 1928: 42-43. 'Iype species: W
Fabricius (orig. des.).

23

214

mm: Frartal setae diverging very little. Male usually without
proclina‘te orbital setae, fanale usually with two. Femle with well
develcped cuter vertical setae. Female with frontal vitta at midpoint
twice or more width of fronto-orbital plates at that level . Postalar
wall se’mlate. Tegulae orangish in gramd color, sanetimes partly
infuscated (with the exception of my, n.sp. where the tegulae is
blackish). Male without apical scutellar bristles. Mesofenur of male
with posters-ventral, apical curb of short, stout, setae. Metatibia with
apial postercventral bristle not differentiated or but slightly so.
Presutmral acrcstichal setae differentiated. R4+5 with dorsal setae, R1
with or without dorsal setae. Genital capsule crangish (with the
exception of ELM where the genital capsule is bladdsh) . Male
aedeagus with basiphallus and distiphallus fused. Juxta absent (with the

W161 of&_effism_ta (Walk-I'D-

KEY'IOSPECIESOFRAVINIA

l leithdorsalsetae ............................................. 2
1’ leithoutdorsalsetae ..... . ....... 25
2(1)Cbstawithventra1semlaeextendimpastSctoR1.. .......... 3
2’ CbstawithventralsemlaeerdirgatSc ...... 8

3 (2)R1withdorsalsetaeexterdingpastberdof8c, malewithsimms
wimlaml Vim OOOOOOOOIOOOOOOOOOOOOO0...0.0.0.0.0... ..... 7

3’R1withdcrsalsetaeerdingatberdofSc ........................ 4

25

4 (3) Male with bifid cerci (figures 2b-c), female with sternites 6, 7,
and 8with nearly parallel margins (figure 2e) 3. advena
4' Oerci of malenotbifid, fanale with sternites 6 and? nnchwider

than sternite 8 (figures 13d, 25d, 32d) .................. . ....... 5
5 (4) Palpus orange ........................................ R. rufipgs
5’ mm b1” OOOOOOOOOOOOOOOOOOO .0. ...... O OOOOOOOOOOOOOOO O O O O ..... 6

6 (5) Oerci of male with long, thin, parallel apices (figure 25b):
hillaeaedeaguswith distinctlydarkenedbase (figure 25c): sternite
8 of female with long, basal projections (figure 25d) mm;
6' Oerci of mle withstouterandmore divergent apices (figure 13b);
hillaemtdistinctlydarkenedbasally, notdarkerthanadjacent
distirhallus (figure 13c); sternite 8 of female without long, basal
projections (figure 13d) .......... . .................. 3. limbiana
7 (3)8ternite50fma1ewithlargemesalwinicwardm11tilcbed
arpearance along mesal convexity (figure 3a); sternite 8 of falale
witha flat, apical lip (figure 3d) R. Midai
7’ Steniite 5 of male withmesal margins nearly parallel (figure 29a):
sternite 8 of ferale uniformly convex, without apical lip (figure
29d) ....................... .. ........... . 3m
8 (2) Males ...... ............ . ..... 9
9 (8) Cerci with apices strongly convergent (figures 9b, 15b) ..... . 10
9' Cerci with apiess slightlycmvergent orparallel 11
10 (9) Legs reddish-orange .. ...... .............. m

26

10' legsgrey ........ . ......... . ........ ....... W
11 (9) Vesica lcng, extending nuch past hillae in lateral view, my be
bentupuardsagainstbasirhallus ..... ........ ..... 14
11' Vesica short, shorter or slightly longer than hillae in lateral
Vlad ...... ... ..... ..................... .............. . .......... 12
12 (11)Prcc1inatetmerorbitalsetaepresent..... ....... .E.dcmesii
12’Proclinateupperorbita1setaeabsent.... ........ . ..... 13

13 (12) Fifth sternite with a pair of large, ventrally produced lobes
and several strcngsetae (figure 18a) .. ....... 3. errabunda
13' Fifth sternite producedmedially, appearing as parallel brushes

(figure 14a) . ...... . ............... . ........ . ......... W
14 (11) Distigiallus scurried, vesica strongly bent upwards against
basiphalusinlateral view (figure 20c) ...... MW

14’ Vesica not strongly bent upwards against basiphalus in lateral

Via ......OOOCOOOOOOOOOOOO ........ O ........ O ..... ......OOOOOOCOO 15
15 (14) Oerci with lag, thinapices (figure 17b) LEM

15' Oerci with rather short, thick apices (figures 34b, 37b) 16
16 (15) Hillae rounded dorsally in lateral view (figure 3%)ma
16’ Hillae pointed dorsally in lateral view (figure 34c) .. am

am
17 (8) Sternites 7 and 8 at least partially fused (figures 9d, 20d).. 18
17’ Sternites 7 ard 8 mt fined ..... 19
18 (17) legs reddishorange ........ . ............. W
18'legsgrey.... ..... ................ 341M

27

19 (17) Sternite 7 cmpletely divided into two separate plates (figure
20(19)Sten11te8m1d1smallerinwidfl1thansternite7arrisimatedin
apical indmtation of sternite 7 (figures 17d, 34d, 37d) 21
20’ Sternite 8 nearlyaswide as sternite 7 (figures 4, 16d, 18d) 23
21 (20) Sternite 7 with a nearly straight anterior margin, not bi-lobed
(figure 17d) . ef ta
21' Sternite 7 incised anteriorly, appearing bi-lobed 22
22 (21) Sternite 7 with untried anterior ldaes (figure 34d) W
22' Sternite 7 with pointed anterior lobes (figure 37d) W
m
23 (20) Sternite 7 long, wide and corspicucus (figure 18d) 3. errahnria
23' Sternite7mtm1d1differerrtissizefranstemites6ard8 24
24 (23) Sternite 8 with lateral, oval depressions (figure 16mm;
24' Sternite 8 withartsuchdepressicns (figure 4) W
25 (1) With 3 postsutnral dorsocentral setae . ............... 26
.25'With4postsuuiraldorsccentralsetae .................. . ....... .31
26 (25) 'legulae andgenitalia orange 27
26’ 'Iegulae andgenitalia black 3M);
27 (26) Dale with hillae broad dorsally and nuch narrowed ventrally
(figure 21c): female with sternite 7 long and rectangular (figure
21d) (Old World) W
27' Hillae of nalemtdrasticallymrrcwedverrtrally: sternite 7 of

l!

28

female oftal triangular, if rectargularthenmtlargerthanwide
(Naonrld) 28
28 (27) Hillae broad dorsally with a distinct nctch, giving the aedeagus
an ax-like appearance (figures 22c, 36c): sternite 7 of female more
rectargularthantriargular ingeneral appearance (figures 22d, 36d)
................................................................. 29
28' Hillaewithout a distinctnctch andnctbroaddorsally (figures 1c,
30c): sternite 7 of female triangular (figures 1d, 30d) 30
29 (28) Hillae with broad, conspicuous notch (figure 36c); sternite 7 of
fanalewith nearly straight apical margin mm
29' Hillae with small mtdl (figure 22c); apical margin of sternite 7 of
ferale with triangular mesal projection (figure 22d) .. B. neithausi
30 (28) Hillae nuch thinner dorsally than ventrally (figure 1c);
sternite 7 of feuale broadly triangular (figure 1d) B. cerba
30’ Hillae nearly equally thick dorsally and ventrally (figure 30c):
sternite7offalalemtasbroadasabove (figure30d) -B:_EA§191§
31 (25) Palpus and legs orange W
31' PalpLB black 32
32 (31) Mesoarrimetathoraciccoxaeanitrodlantersorange m
32’ Cbaearritrcd‘lantersblack ........ ........................ ..... 33
33 (32) Males 34
33' Females 45
34 (33) Genagoldmpruinose 35
34'Genagreypruimse,withatmstyellcwpruimsityrestrictedto

29

echeofgeraatgmalgroove ................ . ............ 41
35 (34) Corpus abrubrtly exparried dorsally, posterior margin of aedeagus
in lateral vial distinctlynotched (figure 6c) W

35' Posterior margin of aedeagus in lateral view anoothly rounded 36
36 (35) Aedeagus with a distinct plate between hillae, extending

anteriorly.............. ....... . ....... ............. WW
36’Aedeaguswitlnrtadistirctplatebetweenhillae........ ..... 37

37 (36) Fifth sternite with long lateral ans (figure 23a)R. lherminieri

37' Fifth sternite larger than wide (e.g., figure 8a) .......... 38
38 (37) Frontal vitta distinctly narrowed posteriorly ...... Km

38’ Frantal vitta with parallel nargins or slightly narrowed

posteriorly ...... ................ .. ............ 39
39 (38) Hillaemmdedarribulbals at tip (figure 10c) Mi
39' Hillaenotbulbous, rather triangular in appearance 40
40 (39) Corps smoothlyr'om'riedposteroverrtrally (figure 26c)

eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeoeoeeeeeeeeeeeee W
40' Corplswithdistimtnedial exparsicn posteroventrally (figure 8c)

.............. . ...... . ........ W
41 (34)Corplsnld1redu3edandthirmedatapex(figure28c) ........

............................................ W
41' Cornisbroadly rcurdedat apex 42
42 (41) Ground color of posteroventral area of tergum 5 orangish, hillae

amearing rather triangular (figure 11c) W
42'Gramicoloroftergm59reythrulglwt,hillaemtdistinctly

30

narrowingdorsally . ........ 43
43 (42) Hillae clear umbranous dorsally, corpus not exterding nuch past

ventralmargin of hillae ....... 3. fiinata
43' Hillae at met with thin clear malbrancus strip dorsally, corpus

amearingratherhllbalsbelowventral margin of hillae 44

44 (43) Aedeaguswith distinct anteriorlyprcducedmedian plate between

thehillae ..... ............ W
44' Aedeaguswitlnltsuchaplatebetweenhillae ............ .. m
45 (33) Sternite 7wit1'mtdensegoldenpruinosity ............ 46
45' Sternite7withdensegoldenpruinosity. .......... .. ..... 49

46 (45) Sternite 7 very broad, not distinctly narrowing posteriorly

(figure 31d) ........................ 3M
46’Sternite7narrowingposteriorly. ..... . ...... 47

47 (46) lateral margins of sternite 7 convergent, nearly frun basal
margin (figure 5d) .................... . ................... W
47’ lateral margins of sternite7 nearly parallel overbasal half 48
48 (47) Sternite 6 with medial imaginatim on anterior margin (figure
27d) W
48’ Sternite 6witha straight anterior mrgin (figure 28d)
49 (45) Sternite 7 withbasal lobes (figure 6d) Law
49' Sternite 7 with lateral nargirs smothly rourried 50
50 (49) Sternite7nearlyequalinsizetosternite6,sternites6ani7

together amearing as a broad oval (figure 35d) W

31

50'Sternite7la1gerthansternite6, sternites6and7notforminga
broadoval ..... ........ ...... 51

51 (50) Genagreypruinose, atmostwith tinge of yellow at nargin with
genalgrcove ......... ..................... BM

51' Gena golden pruinose .......... . ........... . .......... .. R. auriger_la

minia acerba (Walker)
(Figures. 1a - d)

W Walker, 1849: 824-825. Type-locality: Nova Scotia
(fanale). Aldrich, 1930: 15 (symnymy).

W Parker, 1914: 58-59, figure 45. Type-locality:
massadmsetts (male). Metz, 1916: 218, 235, figures 98, 99
(description of dlrunoscmes). Greene, 1925: 6, 21, figure 39
(description of puparium). Aldrich, 1930: 33 (synmymy).

W (Parker). Aldrich, 1916: 114, 121-123, figure 49
(key, redescription). Hall, 1928: 334, 336, figure 3 (key,
redescripticn). Winn and Beauliel, 1932: 81 (locality records).
Bram, 1934: 250 (biology).

W191; authors, nec. van der Wulp. Sanjean, 1957: 55—56,

figures 11411-122 (description of larva).

32

W authors, nec. Rtineau-Desvoidy. Gill, 1955: 651
(biology).

Miriam: Mike—r)- Dodge. 1956a: 188. 189 (W.
redescripticn) . Coffey, 1966: 214, 217 (biology). Pedcham, $314,
1973: 652 (biology).

mm: myth: 6 - 10 um. Panto-orbital plate gray pruinose,
oftenwithcqperishtingeventrally. Genagraypruinosewithabit ofa
yellowishtingeatborderwithgenalgroove. ‘motothreewell develqaed
presutural acrostichal setae; 3 postal‘blral dorsocentrals. 'moracic
plelrae gray pruinose.

HAIR: Gerci of moderate size with apices parallel to slightly
calvergent (figure 1b). Fifth sternite as in figure 1a. Hillae of the
aedeagts well develcped, noticably renewing toward apex (figure 1c) .

FEMALE: Sternite7nd11argerthansternites6andBandrather
triangular in shape (figure 1d). Sternite 6 with sane basal, dark,
infuscaticn.

IARVAE: ‘Ihe larvae may have been described by Sanjean (1957) .

PUPARIIM: Adescriptim ofthepupariummaybeprovidedbyGreene

(1925) .

0189:0515: 'nlisspeciesmaybeseparatedfrunmanig,
Whythepresenceof3stru'gpostarteraldorsocentralsetae. It
iseasilyseparatedfrmfihmnbyitsorangetegulaeaniorange

33

        

none. .... . enoouuu .-
o... \uo. I.¢~o.ooaao.ocoo .A’o 0... no . .
u n .0 n I I o n a 00.... I. I. on... .u h.‘ a.confine-“fiehfi-w-U..fin.’h..~t*h\e.$ ‘0
. one. o a one. I 0 one o n a.“ ...:- ..0u..cuooo nu~¢..~.-.N55.. o. «I... o
.o on. o o H on o . onflmoucuoruuoaouu...~-uu..o..... .. nos. I. u
I I O O .Ieo'Ai-nl-a-Ieoeeeeeou~eo. .

  

 

(a) Male fifth sternite; 0:) Male

Figure 1: Genitalia'of W.
cerci: (c) Aedeagus; (d) Female sternites 6, 7 and 8.

3H

genitalia. Itcantsuallybeseparatedfranthespeciesmlaand
Whythepresaneofatleastzmlgpresuunalacmstidrals
anddarkgrey color. ‘Ihe genitalia are diagnostic (especiallythemale’s
aedeagus) arriwillseparatethisspeciesfranotherspecieswith3
postsuteraldorsooentrals.

TYPES: Female holotype, Nova Scotia (EN-I). mined in 1986, in
good condition.

MATERIAL m: 88 males and 87 females (sac, am, as, (so,
FMNH, K22, 160, (EU, SUNY, WLD), including 5 paratypes of mimlata
(K22). HIGH. Charming, Dickinson Co., 4 June 1983, M. Arduser (male-DSU;
used for figures 1a-c). MICH. Charming, Dickinson 0o., 12 Aug. 1987,
R.L. Fischer (farale-Ifl], used for figure 1d).

DISIRIEJTICN: Canada (Alberta, British Colunbia, Nev BnmSWiGR,
Northwest 'IeHitory, Nova Scotia, Ontario, erec), northern United
States (Alasla, California, Colorado, Maine, Massachusetts, Michigan,
Minnesota, New I-Ianpshire, Nev Jersey, New York, mic, Oregon,
Pansylvania, Vermont, Washirgtm). Distrihrtim frcm material examined.
'Ihe species appears to be relatively Wm collected an: widely
distributed.

BIOLOGY: Little is kmwn of the biology. Coffey (1966) may have

35

rearedmmmnarfidogfeoesardcollecteditmminedrmg.
Ithasbemcollectedinancpenmitepirestaniardmflowersin
Timgami, altario (Bram, 1934). Itmayhavebeen collected in a trap
baited with a bare carcass in Circle, Alaska (Gill, 1955). A series of 4
mlesandeanaleswascollectedinNaanmswickmaramcookHill
sunmit, 637’ by G.E. Sheavell (RIC). One female was collected in New
Bnmswickmflowers offingyG.E. Sheaell(H1C). &
m has been collected in Malaise traps fran the localities of
Gatineau Oo., Quebec (am: 1 male): Dickenscn (13., Michigan (DSU: 8
males, 1 female): and Kenora Oo., Ontario (as: 3 females). A
descriptimandmtesmthebetaviorofdrrumsmnsnaybegivenbyMetz
(1916) . 'Ihis species may have been famd as prey/provision of larvae in
flemstofflsmmumglimemmsay (W:
Sphecidae) by Peckham, nlrczewski, and Psokham (1973) in New York.

REMARKS: misspeciesnaybefonriincollectimsunderthenames
m.lmmieri.memardn§iela- Ineadtofflmecases.
specimensneybemixedwiththoseofgigla,tangiflrp,aswellas
severalotherspecies. Determinatimsofmwnodgeafternwseem
tobereliable, othersshmldberegardedascpesticnable.

Aldrich (1930) errmeously placed m as synmym of 1mm
“Placedmimassyrurymofggigla. 'Ihishascausedsane
prdalals with informatim about this species in the literature. Althcngh

Hayes (1963) statesflesymmyofmellatawithnsielam

36

disassingtheworkoflbtz (1916), I believe that the species in
qresticn may be m since the initial identifications were performed
byParkerandIhavesemaspecimmofggigladeteminedbyParkeras
me.mtmedetemimtimalsounluaedastatexentuntthismay
beageographicalrace. 'Nelvespecimsofflwereshfliedthat
meidentifiedasgfigllagabymrker,imluiim5paratypes. Itcan
mtbemledart,lnaever,thattheqaeciesinquestimwasactually
E5193- Sixspecinensofmsiggvereswiedthatwereidentifiedas
Whyparker. Although Gill (1955) listsarecord ofm
(thisspeciesismvplacedtmderthenanemia),Ibelievethistobea
recordofmmetothemremrthemrargeofthisspeciesard
becauseaspecimenofagflhasbeensembymyself,collectedin
Alaska,b.1thlaskanspecinersofagi_awereimludedinmaterial
examined. 'Ihelistirr;ofac_erb_aasapreyspeciesbyPech1am,§t_a;_,_
(1973) iscorsideredtobeqresticnableandmay applytoagerbaor
M,asbothspeciesarefamdinNevYork. Specimensborrowedfrun
the State University of New York collection did not imlude m
specimens attributable tothis study. 'Ihe figure of themale genitalia
of 35191; byI-Iallcck (1940a, 1940b) matches that of m, rather than
m.mwmmsynmymywiflwmiealee- fledescriptim
ofthelarvaeofggipjabySanjean (1957)arritheplpariumbyGreene
(1925)neya;plytoag:§orgl_s_icl. 'medescripticnheprovidedof
the first irstar is not sufficient to definitively separate these two
species. Ihavenotexaminedseoaricrthird-instarsoftheseuo

37

species and sud: examination may clear up the identity of Sanjean's
species. ‘medwcriptimofthefirstilstarofngigljbymipling
(1936) isbelievedtobethat ofggigla (ratherthanM) duetothe
notatim that the specimens were collected fran Illinois and Iowa. 'Ihe
speciesglsiqlaisverycmflycollectedinlwaaninlinoisanim
specinensothavebeenseenfranthosestates. 'Inespeciesreared
arrl collected by Coffey (1966) my be 921—135 or M.

maven (Walker)
(Figures 2a - d)
W Walker, 1852: 324. 'lype-locality: Brazil (male).
W Walker, 1852: 327. Type-locality: Brazil (female).
ropes, 1976: 629-630 (notes on type).
WM ‘Ibwnserri, 1927: 220, 295. Type-locality:
Brazil, Sao Paulo, Itaquaqncetuba (male and fanale). lwes, 1932:
46, pl. 3 (paratype examined).
WW Reel. 1931: 149. figure 18- Type-
locnlity: Bolivia, Northern Chiquitos (male).
mm (Walker). lopes, 1976: 629-630, figures 1-3
(redescription of type).

arms: length: 5 - 7 nm. Pronto-orbital plate yellow pruinose.
Frurtal vitta usually bicolored, reddish brown anteriorly and black
posteriorly. Ventral semlaeofcostaexterdingpastSctoRI. Dorsal

38

setaepresentle,extel'ldi.rlgtoberriofSc.
MAIE: Mesothoracic tibia with small anteroventral seta. Fifth

sternite (figure 2a) with an oval median hole. Cerci bifid (figures 2b-
c). Aedeagus salewhat c.1rved (figure 2d), hillae mstly marbranous.
FEMALE: Sternites 6, 7, and8nearlyeq1al inwidthandwell

develqled (figure 2e) .

0mm: 'merangesofMandmigaioverlapinmnhof
Brazfl,h1tbofl1malesarrifanalesof§1enacanbeeasilyseparatedfrm
WhythedorsalsetaemplexterdirgmlytotheberdofSc
ratherthancartimirgtonearthewingmargin. 'Ihischaracterwillalso
separate this species frun the Central Anerican/nortlmestern South
Anerican species We (Dodge). 'Ihe bifid cerci of themale is
conspianlsardmiquetothespeciesMalnlgthem. Fenales
mnbeseparatedfrmmlmhiem.aetflseiardmbymemofflie

TYPES: Male holotype, Brazil, Ex. coll. Saunders, 68-4. "M"
(8N!) . Redescriptim given by lcpes, 1976.

MATERIAL W: 8 males, 8 fanales. Paraguay, Villarica, VIII,
F. made collector (rm-male; used for figures 2a-c). Brazil, Nova
We, 27" 11' 8. 52° 23' L., 300—500 111, 11 1973, Fritz Plaumann

(Hm-male: used for figure 2d). Brazil, Nova remade, 27° 11' B. 52°

39

 

 

 

Figure 2: Genitalia of mm. (a) Male fifth sternite: 0)) Male
cerci (lateral view): (c) Male cerci: (d) Aedeagus; (e) Female sternites
6, 7 and 8.

149

23’ L., 300-500 m, VI 1950, Fritz Plaunam (nun-5 males, 6 fenales; one
farale used for figure 2e). Brazil, Nova Teutonia, 27° 11' B. 52° 23' L.,
300-500 m, III 1957, Fritz Plaumann (Fm-male, fenele) . Paraguay,

antera, Hapua, March 1954 (IBIJ-fatele).

DIS'mIEII'Iw: Argentina, Bolivia, Brazil (1th Grosso, Minas Gerais,
Sao Paulo, Santa Catarina), Paraguay. Distribution frun material
examined and lopes (1969) .

BIOIOGY: Very little is kmwn of the biology. Iopes (1976)
irriicates that fanales fran an existing culture were used for catparative
purposes. No further details on biology or larval substrate were given.
'No dissected fanales yielded 15 ard 17 larvae, respectively.

W: Determinedspecinenswilllsuallybefamdurrierthename
Wincollectims.

WV '3 ° ' (1099-5)
(Figures 3a - d)

W; lanes, 1946b: 227-230, figurs 1-5. 'IYpe-
lcmlity: Brazil, Mato Grosso, Salobra (Hale). lopes, 1974: 272-273
(distributim and rearing). Lopes, 1982: 315, figures 72, 75, 82
(descripticn of first instar) .

L11

mm: length: 5 - 8 m. Frurto-orbital plate pale yellow
pruinose. Ventral setulaeofcostaexterdingpastSctoRl. Dorsal
setae present an R1 extending past bad of So.

MAIE: Mesothoracic tibia with anteroventral seta. Fifth sternite
(figure 3a) with a large median hole and several long conspicuous setae.
Cersi sinus in lateral view arrl with apices convergent in posterior
view (figure 3b) . Aedeagus sanadhat alrved (figure 3c) , hillae mostly
mallaranous.

FEMALE: Sternites 6, 7, and8nearlyequal inwidth. Sternite8
flattered alcng apical margin (figure 3d) .

lARVAE: ropes (1982) described the first instar.

DIAGNOSIS: Verysimilartom. 'Ihespeciesappeartobe
geographically isolatedbytheAndesMamtains. Malesarennsteasily
separatedbytheshapeofthefifthsterniteandbytheformofthe
aedeagus. 'Iheshapecfthegenitalsterniteswilldistinguishfanalesof
thesebnweciesfrunalearnther. Bothmalesandfanalescanbe
separatedfranmmalker)byfl1epreserneofdorsalsetaee1¢arling
pastthebendinSc. mlesofMareeasilyseparatedfranm
bythesinmscerciwhidlaremtbifid.

TYPES: Male holotype: Brazil, Mato Grosso, Salcbra, Canissao do
Instituto Oswaldo Cruz, I 1941 (ICC). Female allotype: Brazil, Mato
Grosso, Salcbra, Cmissao do Instituto Oswaldo Cruz, I 1941 (ICC).

H2

 

Figure 3: Genitalia of W. (a) Male fifth sternite: (b)
Male cerci: (c) Aedeagus; (d) Ferale sternites 6, 7 and 8.

H3

‘menty-fcur paratypes (ropes, 1946b).

MATERIAL EXAlENED: 1 male an! 2 fenales. Paratype, Brazil, Mato
Grosso, Saldora, Can. I.O.C., Jan. 941 (rm-male; used for figures 3a-c).
Paratype, same data (rm-fanale; used for figure 3d) . Both qaecilnens
salamat tareral. Argentina, Team, Teaman, Jaruary 1953, S.P.
Colalao (FBU-fenale: in poor conditim) .

DISIRIHJTIQJ: Argentina, Brazil (Ceara, Mato Grosso, Minas Gerais,
Parana, Sao Paulo). Distribution frun neterial examined and Lopes

(1946b, 1969, 1974) .

BIOLOGY: Iarvaeneybecoprqhagms. Lopes (1974) reports thatone
fanale collected fr'an Brazil, Ceara, mama on 23 VII 1973 yielded 21
larvaewl'lidiwererearedmtnmanfeces. 'Nentymaturelarvaewere
ovservedm26VIIarritheadultsanergedm5VIIIl973(14daysfran
firstinstartoadult). Bearedadultswereverymall. Firstinstars
describedbylfpes(1982)vereamarentlywtairedfrunanexisting
allure ofthisspecies, butnodetailsmrearing/biologyweregiven.

W (00696)
(Figm 4)
mm Dodge, 1956a: 187, figure 15. Type-locality:
United States, Georgia, Cobb Camty, Kennesaw Mt. (fanale). ropes,

MU

1961: 419, 422, figures 1a-d (key, locality records). Wiltm, 1963:

.313 (biology). Rdfiendorf & Gregor, 1973: 8 (key). Hardy, 1981:
381, figure 164 (redescripticn) .

mm: length: 4-7m. Frurto—orbitalplategreypruinose
with pale yellowish tinge. Sternite6usually dark, 7and8crange.
Sternites 6-8 nearlyequal inwidth (figure 4). leithdorsal setae
exterriingtobendofSc.

0111910615: Generally a snall species with sternites 6-8 very
similartothoseofdggggi. Itmaybeseparatedfranthisspeciesby
theformofthelateralnargirsofsternite8,whidldomthavethe
distinctovaldepressionsseeninm.

TYPES: Female holotype, United States, Georgia, Cdlb County,
Kamesaw Mt., June 18, 1952, or: fly trap (type nunber 62337, USNM).
Paratype fenales are widely distributed in many collections. Dodge
(1956a) lists locality data for 198 paratype specimens and 1473 tramed

specimens.

MM‘ERIAL EXAMINED: 78 fanales (GS, MCZ, 160, $0, WID) incluiirg 5
paratypes (2m: 2-osu; l-QS). m: Highlands Co., Archbold Biol.
Sta., 11—x-1964, P.H. Arnaud, Jr. (QS-fenale; used for figure 4).

145

 

Figure 4: Genitalia of W. Fanale sternites 6, 7 and 8.

MB

DISIRIEJTICN: Canada (Ontario), United States (Alabama, Arkansas,
Delaware, Florida, Georgia, Hawaii, Illinois, Indiana, Kansas, Kerltucky,
Lmisiana, mrylarri, Mid'ligan, Missalri, Nebraska, Nev Jersey, New York,
North Carolina, mic, Pelnsylvania, Salth Carolina, Tennessee, Texas,
Virginia, West Virginia, Wiscmsin) . Distributim frun material
examined, Dodge (1956a) and ropes (1961).

BIOLOGY: Larvaeareccprqahagwsmcamivoreanimnivore dung.
Dodge (1956a) reportscnrearing records of 165 specimens. Mostwere
rearedfrmdcgardmmanfeoesarrisatelarvaewererearedmneat.
Wiltm (1963) reports smallmmbersof specimensrearedfrandcgfecesin
Hamil. misspeciesisarparentlyparthenogeletic. Malesareunknown
althcnghthefenalesarefairly comm. Bearings resultedinall
fenales. 'nleqlecieshasmt,asyet,beenrearedthrulghtuo

generatims to prove its parthenoge'letic nature.

Emma's (Wants-r) duplex
(Figures 5a - d)

W Walker, 1849: 818. Type-locality: N. America (fenale).
Aldrich, 1930: 15 (synonymy).

W Walker, 1852: 323. Type-locality: U.S.A. (mle).
Aldrich, 1930: 17 (synmymy).

WW 'Ihmscn, 1869: 535-536. Type-locality: rewaii,
Banlulu. Jdlrstm and Tiegs, 1922: 184, figure 6 (biology,

U7

redescriptim). Hardy, 1927: 458, figure 11 (redescriptim). Hall,
1928: 334, 339, figure 9 (key, redescripticn). Ball, 1929b: 87
(locality'data).

W Parker, in part. Metz, 1916: 218, 234 (description of
dlmnosanes).

W (Parker, in part). Aldrich, 1916: 253-255, figure
120 (redescripticn). Greene, 1925: 3, 7-8, figure 3 (description of
puparia). wulson, 1932: 84, 89 (biology). ‘Winn and Beaulieu, 1932:
81 (locality data).

W authors, nec. Robineau-Desvoidy. Aldrich, 1930: 13
(W)-

W authors, nec. Rabineau-Desvoidy. Knipling, 1936:
418, 420, 421, 425, 426, 429, 434-435, figum 6, 34, 51, 65, 81
(description of first instar, biology). Hallock, 1940a: figures 29-
31 (male genitalia). Hallock 1940b: 207, 210, 215, 221-222, figures
116-118 (key, redescription). James, 1947: 48 (biology). IMerrill
ard Hutsa'l, 1953: 678 (biology). Sanjean, 1957: 17, 25-26, 39, 40,
41, 57-58, tables 1-7 , 9-15, figures 123-131 (biology, description
of larva). Blickle, 1961: 802 (parasitoid records) .

W; authors, nec. Robineau-Desvoidy. Lopes, 1946a: 137
(locality data). Lopes, 1948: 567 (locality data). van Emden,
1950: 195, 198 (host information) . Boback, 1954: 27, 73, figures
242-244 (description of male genitalia). Gill, 1955: 651 (biology).

Dodge, 1956a: 188-189 (synonymy) . ches, 1961: 420, 422-424 (key,

U8

locality data). Boyes, 1963: 1201-1202 (description of
dmosanes). Coffey, 1966: 213, 217 (biology). Maser and Wirgc,
1967: 731-732 (biology, parasitoid records). Poonlaugh, 3314,
1968: 22, 35, figures 1-2 (biology). Valiela, 1969: 213 (biology).
nmel and Imapp, 1970: 167-169 (biology). Bayes and Turner, 1971:
444-447 (parasitoid records). Poorbaugh and Linsdale, 1971: 52, 55
(biology). Stoffolano ani Streams, 1971: 195-196, 198, 200-201, 207
(parasitoid information) . Pedcham, my 1973: 652 (biology).
Stoffolano, 1973: 263, 270 (parasitoid informticn) . Valiela, 1974:
375, 378 (biology). Merritt and Anderson, 1977: 43 (biology).
Davis and Turner, 1978: 113, 119-124, 127, tables 1-7, 12-13,
figures 2, 3, 6 (biology). Wharton and Moon, 1979: 81, 87-88,
figure 1 (description of puparium). Peterson, ML, 1981: 512,
514 (biology). Pickers, 1981: 523-526 (biology). Figg, steal...
1982: 476 (biology, parasitoid records). Figg, fl, 1983a: 962,
963 (biology, parasitoid records) . Figg, fl, 1983b: 994-995
(biology, parasitoid record).

W authors, nec. Robineau-Desvoidy (misspelling of
W). Hardy, 1981: 383-385, figure 165.

AflJIII':Le1gth:6-12nrn. Mostspecinensaroundmm. HEAD:
Franto-orbitalplateardgeralightgreypmirnse. Genalgroovenot
pruimse,ratherdarkbrownincolor. ‘nmAX:4postsrmraldorsocentral
setae, increasing in length posteriorly. Subapical saltellar setae

H9

really and: strager than adjacent lateral scutellar setae.

m: Tergite 4 with well develcped, reclinate nedian marginal
setae. Inner margin of apices of cerci smoothly rounded (figure 5b) .
Lateral arm of sternite 5 not stragly produced, giving a trapezoidal
shapetothesterniteasawhole (figureSa). Aedeagusmtstrcngly
bulbousattipardhillaemttaperirgdorsallybutmxhthimler
ventrally.

FEMALE: Sternite 7 broadly triangular in shape (figure 5d).
Genital sternites without golden pruinosity.

LARVA: A descriptim of the first-instar nay be provided by
Knipling (1936). Sanjean (1957) provides detailed descriptions of all
three instars.

PUPARIIM: A description of the puparium is provided by Greene (1925)
and Rharton and Moon (1979) .

WIS: Chnbeseparatedfrunagrfioveritsmrthernrangeby
thepresenoeof4postsub1raldorsocentralsetae. Canbeseparatedfran
Wardggtabythegreycoloratimofitsheadardabdmen.
mlescanbeuslallybeseparatedfrmWardgjsseuibyflelesser
developed lateral sartellar setae, ascmparedtothesubapical sartellar
setae. 'Ihetriangularappeararneofsternite7ofthefe1aleseparates
thisspeciesfruntherectangularsternite7ofme_rgla. ‘Ihelackof
galdel pruinosity a: the genital sternites easily separates the fenales

frmgigeui, milieu andgga. Usually noticeably larger in size

50

 

 

(a) Male fifth sternite; 6)) Male

Figure 5: Genitalia of Mange.

(d)Fena1esternites 6, 7 ands.

e
I

cerci: (c) Aedeagus

51

mmmm. Siapeofgenitaliaprovidesdiagnostic
diaracterstoseparatethisqeciesfranothers.

TYPE: Fenale holotype, ?N. Amrica (EN-i). Examined in 1986,
amears to be sanevhat tener'al.

MATERIAL EXAMINED: 1632 male ard 1296 females (32C, arm, (ES,
EU, FMNH, GAD, HZ, DBU, OSU, WLD), including male holotype of m,
casidered to be specimen frun U.S.A. frun series of 3 males. Specimens
daservedfranflawaii calfirmplacenentofmlljmjsassyrurym. Mason
(30., MIG-1., 'I’ZON, R17W, S 27, 23 July 1968, Norman '1'. Baker (male-EU;
used for figures 5a-c) . Hickory Corners, Barry Co., MIG-1., 10 July 1964,
R. Mathevs collector (fenale-DBU: used for figure 5d) .

DISIRLEJI'ICN: Canada (Alberta, British Colunbia, mtario, Qlebec,
Saskatchewan), United States (Arizara, California, Colorado, Hawaii,
Idaho, Illinois, Irdiana, Iowa, Kansas, Kentucky, Maryland,

' Massadnsetts, Michigan, Minnesota, kissciri, Montana, Nebraska, Nevada,
Nev Mexico, Nev York, North carolina, North mkota, (mic, Oklahana,
Oregm, Erode Islard, South mkota, Texas, Utah, Virginia, Washington,
Wisca'sin, Wyuning), Mexico (Baja California Norte, mrango, Hidalgo,
Jalisco, mico, Mexico D.F., Puebla, Salora, Zacatecas). This is the
mstcammlymllectednmberoffiflniaMofMexico, withthe
ecceptimoftheSartheasternstatesoftheU.S. Itappearstobecane

52

soarcerprcoeedirgsouthinbbxico.

BIOLOGY: Averycannmcoprqhagous species. 'misspecies ismost
may found breeding in cow durg (Jdmstcn and Tiegs, 1922: Blickle,
1961: Coffey, 1966; laser and Wingo, 1967: Poorbalgh, gal“ 1968;
Valiela, 1969; Hayes and Turner, 1971; Stoffolano and Streams, 1971:
Stoffolano, 1973; Valiela, 1974: Merritt and Anderson, 1977: mvis and
Turner, 1978: mm, 1981: F193, fl, 1982; Fig, _e_t__§_l;, 1983a;
Figg, m, 1983b). Merritt and Anderson (1977) found m imabiting
ard colonizing cattle droppings in four different pasture and rargeland
ecosystens at the Sierra Foothills large Field Station in California.
misspecieshasbeencollectedardrearedfrunbomemecposedcowats
and shade exposed cowpats by Poorbaugh, ;e_t_a]_.,_ (1968). Davis and 11m
(1978) nenticnthat itwasmtmmelaltocomrtmOormre larvaeina
single cowpat. ‘Lhey found this species at all 9 research plots. One
halfoftlesarcqmagidstheycollectedverereareseltedbythisspecies
atrium. 'meynotethattheadditicncfcattletotheplots
affectedtheahndarceoffiayinjamrethananyotherfactorardthey
report rearings frun cow nanure. In additim to the literature citations
ofrearings, manyofthespecinensexaminedwererearedfrmcowmamre
ardIhaverearedthisspeciesfrmcownemremseveraloocasims. E;
whasalsobeelrearedfrmsheepdmginNevJerseybyWilsm (1932)

and in Washington by Coffey (1966) . Coffey (1966) also rqlorts
collectingandrearingthisspecies franhorse, swine, ardhumandlmg.

53

Coffeymentiorsthat itisatu'actedtodogfecesardtvospecinenswere
reared, monecccasicn, frandogfecesinElCerrito, Californiaby
Pcorbaugh and Linsdale (1971) .

Sanjean (1957) fund that the average time needed for this queciee
tomature fromfirst instartoadultonpork liver at 27°Cwas 12.6 days.
'Ihetineduratimincreasedtol3.4 daysat25°C. I-lealsofamdamean
of 20.4 larvae per gravid fenale, with a sanple size of 47 gravid
fenales.

Richardson (1916) may have collected a few specimens of this species
dln'irghissufiyoftheattractimofDipteratoanmnia. Adultsmay
havebeencaptured inmicn fields inMichiganbyMerrill ardmrtson
(1953) . Gill (1955) reports capturing this species with a trap baited
with a hare amass. It seems likely that this record actually applies
tothecaptureofghm, butitneyarplytoamgia. Forfurther
discussion, look \mder "Renarlcs" section of m. Sanjean (1957)
collected this species at traps baited with: horse dung, cow dung, human
feces, chicken entrails, bird feces, sheep liver, hog liver, bull brain,
freshly killed American cockrcadres, cow udder, hog vicera, deer heart,
woodchuck, rabbit, and fish cadavers. He collected this species very
mlyasitrestedmastcmwallintheIthaca, NYarea. Healso
collected specimens on field stones, sticks on the grand, on pond
lillies, at pitcher plants, and a1 vegetation in fields. Rmmel and

Knapp (1970) collectedthisspecies frunJmethmmSeptalberin
Winu'apsbaitedeitherwith liver, banana, ormaltmixmre.

51+

Invisarrlfimler(1978)givege1eraldatamahndarneanidatesofadult
activity. Specimens were collected in a Malaise trap, a baited Malaise
trap, a baited Bishqp trap, and with a net. Petersen, eth. (1981)
notethatthisspecieswascamaflyattractedtopelletsofSWASS, abait
toxicant systen for screwworms.

'mreeparasitoids ofmwerefonndtoenergefrantheplparia
by Blickle (1961) frun specimens collected in New Hanpshire: meeta
mum (Say) (Emulate—Ia: Bracerfidae): WM (Say)
(Hymencptera: Figitidae): and mu; sp. (Hymencptera: cynipidae) .
Kaiser and Wingo (1967) provide information on field parasitizaticn of
W (Say) (Hymencptera: Brawfidae) fmm cow mamre in
centralMissouri. 'nlisspecieswasgroupedtogetherwitth
minim. Belem. Esme. and W weer the
WM sp. byHayesand'Iurner (1971) intheirstudyof
parasitoids of coprqilagous Diptera in Virginia. They note that this
group of species was Parasitized by W (Say)
(1W: Bram): W (W: qniPidae);
WW (Hyman: Figitidae): and sleaze SPP-
(Colecptera: Staphylinidae). Fig, M; (1982) reared this species
franbovinedlnng. mirteelspeciesofDipterawererearedfrunthisdlmg
and a list of 12 parasitoids is give: without nention of host
specificity. Fig, $3.13. (1983a) note a pqaulatim peak during early
sprirgeddeclireofahndelcethmlglnrttherelairdercftheseasmin
central Missouri. They report that the parasitoid species Mam

55

M (Say) W: 3130011638); mm Sp. (Hynelcptera:
Diapr'iidae); and mu; sp. (Hymencptera: Eucoilidae) accounted for at
least 2.8% mortality. Fig, My (1983b) give further informticn on
the host/parasitoid relatielships between W and W
(Say) (Hynencptera: Braomidae). Stoffolano ard Streans (1971) discuss
tnstreactiesofgmg'mtofllepansiticneetodemm
Wis Nickle. Stoffolano (1973) mtims that the factors resulting
in host specificity of the nematode W in &
m are encapsulation and melanization. Peddlam, £11,, (1973) report
misspeciesaspartofflrerestprovisiasofmmm
W Say W3 3131391338)-

A questia'nable record of m as a parasitoid of Emmi
Sm. (apleoptera: Scarabaeidae) is given by Van Enden (1950). Pickens
(1981) fund that, in laboratory tests, m larvae were significant
facultative predators of face fly larvae in cow manure. This species has
alsobeelrecordedfrunfllreecasesofenposedmmanintestinalmyiasis

in 'lexas (Janes, 1947).

REMARE: 'nrernnerclatnreinvolvedwiththisspecieshasbeen
rathercmfusedinthepastardmdnofthecmfisimdealswiththe
sepandmoffldsspeciesfrmfiflaafiflemistflcenidertityof
W1. 'mefirstnamethatwascamu'nlyappliedtothisspecies
mkflcer'sfimmjs. Quecimsofmmmweredetemined
\nderthisnamegeerallybetweell914ard1928. ‘Iherevisimprovided

58

byfhll(1928)placedmi§asasyru1ymofmlnmi§. Forseveral
yearsspecimensofgmfiaanimlavereplacedmderthisname.
Aldrich (1930) examinedtypesof Anerican Sarcophagidae inEurcpean
nlsemardheprovidedthemistakelsymnymyofgnxiaardgmwith
Wed. mmmmmidertifiedasmmgflmtil
Dodge(1956a)wasabletoprovided1aracterstoseparateglgr_gl_afran
m. 'nlerefore,refere1essbafore1956cmceminga1aofthe
earlier nanes are considered as potentially refering to this species,
mlessctherwiseindicatedbytheauthor's figures or ccmnents.
Unfortunately, Dodge (1956a) continled the inprcpar usage of
Wadesefiornaneofm. DisclssiorswithDr.W. L.
names,Jr.haveprovidedsaneirsightintohevthismistakeocalrred.
Amareltly,bothmeseriDodgeirdqaedeltlydiscoveeddlaractersto
separatevmatwascesideredltgminieriintotmspecies. 'Ihetypeof
Wmmtacmallyseenbyeitheroffllesespecialists,hrt
specimersrepresentingthetwospecieswereirdepeldentlyserttoE.
SeguyattheNaticnalmsennofNaunalHistoryinParisforcanpariscn
withthetypa. fiecamarismfldicatedflatmmgni=m.
Unfortunately, neither thought to include a third species (actually a
carplex of species), m, which was originally considered just a
variation of misbyAldridl. In 1986, the authorwas ableto
eminethetypaoflheminieriardestablishthatflraholotypadidmt
mtdlthespeciesthathadbeenplacedtmderitsnama. Drrirgthissama
yeartheauthorkasabletoecamirethetypesofWalkerattham'itish

57

Mlsemn (Namrall-Iistory). Directecaminatimofthetypesreentedin
theindicatedsyralymy. 'nms,gmgj§becanesthene:tavailablenamefor
this came: species.

Actually, mi: represents a calplex of closely related sibling
species. Closesuldyofthisspeciesoverthawidegeogramical range
that it occupies indicates that the variatim cbserved applies to more
than are species. Two larval forns are easily recogrnized fran puparia
pinned with reared flies and is mentioned by Dodge (1956a). 'Ihe first
form, mlid'nisverycamm ineasternNorthAmerica, hasverydarkly
pigmented dorsal tubercles. The second form lacks these conspicuous
tubercles. mnydissecticrs of specimens franpivotal areas, sud'las
Arizma, Oregon, aniMexico, leadtheauthortobalieve thatmorethantwo
gecies are involved in this canplex, but no definite d'laracters have been
fourd that will allow cmsistert separation of this species into snaller
tnlits. It is believed that rearing studies are needed, especially fran
western localities, to determine the range of infraspecific versus
intraspecific variation in this cmplex.

'me listing of W fran Bolivia by angel (1931) and fran
WindbyBlandlard (1939) areerrors, asthisnameappliedto
WmmthistineedmliviamldArgeltinaarevellbeyed
the range of either of these species. Ehgel’s description and figures
are not sufficielt to definitively place his species under any currently
recognizedname. 'n'nelistingobedem (1895) fran
Floridaiscesideredanerrorasthisqaeciesnamahadmtbeeuacoepted

58

intouseageinNorthAmaricaatthistimeardtheauthorhasseelno
specimensofanxiafranFlcrida.
Specinersofagfigneybafandincollecticnslmderthenares
mallepsllimis. «nee-mien- Alldetemimdspecimensstmld
bereoheckedfor separation ofgggflafrmmixedseries.

W (Hall)
(Figures 6a - d)
W Hall, 1928: 339-340, figure 10. Type-locality:
Chile, Argol. Hall, 1937: 351, 373-374 (key, locality records).
W (Hall). Ortiz, 1946: 170 (lomlity record). Roback,
1954: 74 (synonymy). Lopes, 1969: 21 (locality records).

ALIJIII‘: Length: 8 - 11 um. Gena ard parafacials golden pruinose.
Gelaabnlptlydlargestogreypmirnseatlineextedingfrmhirdmargin
of eye. Presutural acrostichals reduced, enly slightly differeltiated.
Usually with mly 2 katepisternal bristles.

MALE: Sternite 5 almost forming an equilateral triangle (figure
6a). Apices of cerci very slightly sinnus alelg inner margins (figure
6b). Cbrpus abruptly expanded dorsally, hillae broadly rotnded (figure
6c).

FEMALE: Sternites 6, 7, and8goldenpruinose. Sternite6nnch
wider than long. Sternite 7 with anterolateral projectims (figure 6d) .
Sternite 8 with broad, apical invaginaticn.

59

 

 

 

 

Figure 6: Genitalia of W. (a) Male fifth sternite; (b)
Male cerci: (c) Aedeagus; (d) Female sternites 6, 7 and 8.

60

WIS: 'Blegreyccnraeardtredlantarseasilyseparatethis
speciesfrunm. 'nleabruptlyecpardedcorplsseparatesnalesof
thisspeciesfrunall others. 'Iheanterolateral projections of sternite
7servestosq>aratethefenalesofthisspeciesfranallothers

TYPE: Male holotypa, Chile, Angol (IBNM).

m m: 29 males and 25 females (are, m, 160, 9111))
including 24 specimens fran Angol, Chile. ARG., Santa Fe, Lagtnna Paiva,
S. Bolle, Jan. 1974 (male-BBC: used for figures 6a-c) . Angol-(hile, 15
Oct. 1962, E. Bostidas (fenale-FBU: used for figure 6d).

DISIRIHJI‘ICN: Argentina (Corddla, Bntre Rios, Santa Fe), Brazil
(Rio Grarde do Sul), Chile (La Araucania, Maule, Santiago). Distribution
frun material examined and Lopes (1969).

BIOLOGY: Unhrnvn.

REMARKS: Hall (1937)mistakellysym1ymizedthisspecieswith3,_
mm. Hanctesthatthesyralymywasquestimableandthathehad
mtsttfliedthetypaoflglfgrti. Ortiz (1946) liststhisspeciesinhis
mtalogofthaDipteraofo'nilewittmtcmmelt.

61

W (W)
(Figures 7a - o)
W 'Iowrseld, 1912: 357-358. Type-locality: Paru, Piura.
Hall, 1928: 334, 341, figure 12 (redescription).
W (leased). Townsend, 1917: 191, 194, 195
(W placenent, k131?) -
W (W). Roback, 1954: 73, figures 233-234 (new
canbination, male genitalia). ches, 1969: 21 (locality records).
Lopes and Tibana, 1982a: 136 (locality records).

Awm‘ MALE: Length: 10 - 13 m. Gela and parafacials golden
pruinose. Gelawithabmptdlangetogreypruinoseatlineecteding
fran hind margin of eye. antal vitta narrowing posteriorly.
Presutural acrcstichals enly slightly differentiated. Sternite 5 very
long and rather roln'nded basally, giving the appearance of an isoceles
triangle (figure 7a) . Apices of cerci sinus along inner margins
(figure 7b). Parameres broader than those of other Neotrcpical species.
Hillae broadly rended dorsally (figure 7c) .

FEMALE: Not separable frun several other SouthAmerican species.

DIAQDSIS: This species can be differentiated fran other synpatric
species by its relatively large size, carvergent nergins of the frontal
vitta, broad parameres, ard shape of the genitalia.

62

 

Figure 7: Genitalia of W. (a) Male fifth sternite; (b)
Male cerci; (c) Aedeagls

63

TYPE: Female holotypa, Piura, Paru, March 29, 1911 (USWI).

mm mm: 6 males (as). Rio Palelque, m., 0°35's,
79°22’W, 22-26.II.1976, G.& M. Wood, 150m (2 nales; one specimen used
for figures 7a-c). m., Rio Palenque, 0°35's, 70°22'w, 22-26 Feb.

1976, 150 m., G.E. Shevell (4 males).

DISIRIEII‘ICN: Chile, Costa Rica, Ecuador, Panama, Peru.
Distribution fran material examined, Hall (1928), and Lopes and Tibana

(1982a) .

REMARE: Msed(1911)notesthatthisspecieshasbee1dissected
arddrawn,brtdidmtpublielthefiguresinthispaparardmfigures
acocnpanied his description of this species in 1912. Although fenales
carnutbeseparatedfrunseveralotharSarthAnericanspeciesatthis
timeardtheholotypeisfenale,thisspeciesisdescribedfrunmales

which match Tomsed’s (1917) calchththegensWand
thadescriptienofnalesdetermirsdtobethisspeciesby'l‘owrsed.

W (W)
(Figures 8a - c)

614

W 'Ibwrseri, 1915: 409-410. Type-locality:
Peru, Croya.

W ('Ibwnseri). Hall, 1928: 334, 342-343, figure 14
(redescriptim).

mm MALE: Legth: 10 m. Gena and parafacials light golden
pruinose. Gelawithabnptdlalgatogreypnlinoseatlineectedirg
frun hind nargin of eye. Presutural acrostidlal setae only slightly
differeltiated. Sternite 5 with nearly a straight anterior nergin, not
deeply cleft (figure 8a) . Carti deeply cleft, apices sinus along inner
margin (figure 8b) . Hillae alnost triargular in appearance, corpus
broadlyrundedwithposteranesaltnmperteriirgfrunapexashort
distance upward (figure 8c) .

FEMALE: Not separable frail several other Neotropical species.

WIS: 'n'nenearly straight anterior margin of sternite 5 and
shapaoftheaedeagusseparatethisspeciesfrunallothersimilar

species.

TYPE: Male holotypa. Croya, Paru, over 12,000 feet, Rio Mantaro
valley batten, March 7, 1913 (IBM).

mm: 2111316 (1w). more, QJitO, Pid'lirtha, 15

January 1958, R.W. Hodges (used for figures 8a-c). Calderm, m,

65

 

Figure 8: Genitalia of W. (a) Male fifth sternite;
(1:) Male cerci: (c) Aedeagus

66

Pidlincha Prov., 1 my 1958, R.W. Hoges, 8500 feet.

DISIRIETI'IQJ: Ecuador, Peru.

BIOLOGY: Unknown.

W (Dodge)
(Figures 9a - d)
W Dodge, 1956b: 97-99, figures 1-5. 'Iype-locolity:
ciba, Habana, lomas de Camoa. ropes, 1969: 22 (locality records).
Rdldendorf and Gregor, 1973: 8, figures 15a-c (key, male and female

genitalia) .

AllJLT: Legth 6.5 - 8 run. Pronto-orbital plate grey pruinose with
yellow tirge. Legs with tibia ard femora reddish-orange in grand color.
leithdorsalsetaeexteriingtobeniofSc.

MALE: Fifth sternite Y-shaped, the sides divergent and straight
‘ (figure 9a). Carci with apices calvargent (figure 9b) . Aedeagus with
tell develqaed vesica and large, bowed, rather memoraluls hillae (figure
9c) .

FEMALE: Sternite6nud'nwidarthan5. Sternites7and8fused.
Sternite 8 narrow with a praninent pair of lateral pits (figure 9d).
Note: description of fenale and figure 9d fran Dodge (1956b) as no
fenales of this species were ecamined by the author.

67

 

 

(a) Male fifth sternite; (b) Male

Gemdtalia of Barinia_bsrrei-

Figure 9

(d) Fenale sternites 6+7 ani 8(Fena1e redraw: fran

e
I

cerci: (c) Aedeagus

lbhdeldorf and tiger,

1973).

68

DIAGNEIS: misspeciesappearstobecmfinadtotheislandof
Cuba. 'Bsredlegswilleasilyseparateflmifrantheothertvn
speciesthathavebeencollectedmoma,_e_fmgardgm. 'Ihe
maleandfenale gelitaliaarelmiquearddiagnostic.

TYPE: Male holotype, cuba, Habana, Lanes de (:anoa, March 14, 1952,
Dodge and Seago; net (type amber 62778, USNM). Fenale allotype, Clba,
Oriente, Sierra MaEtra numtains, Lana del Gato, June, 1952, Fernardo de
Zayas (USNM). ‘Ihreeneleparatypes, samedataasholotype, butsane
takenbytrap. Gremleparatype franolba, Beliato.

MAL EXAbflNED: 1 male. Paratype, Clba, Boniato, 11-25-04, H.S.

Parish (EU; lsed for figures 9a-c) .

DISIRIHJTICN: CLIba.

BIOLOGY: Unlcuvn. Identified as a synanthrcpic species by
Rdldendorf ard Gregor (1973) .

W1 (Prado and Fcnseca)
(Figures 10a - d)
W Prado and Fenseca, 1932: 39, figure 7. Type-locality:
Brazil, Sao Paulo. Hall, 1937: 373 (synonymy).

69

W (Prado and Pam) (miSSPelli-rq 0f 31:91:21)- I096.
1941a: 221, figures 17-18 (desaiption of spermathecae and accessory
glands). Lopes, 1974: 272 (locality records). Ups, 1982b: 313-
315, figures 69-71 (descriptim of larva).

W1 (Prado and Fonseca). knack, 1954: 74 (synonymy).
m and Kano, 1968: 295-296, 299-301, figures 8-12 (mle alfi
female genitalia, biology). Iqaes, 1969: 21 (locality records).
Lopes and Tibana, 1982b: 288 (locality records).

mm: Length: 11 - 14 urn. Gena and parafacials golden pruinose.
Gerewithabnptdlangetogreypmirnseatlineexterdingfranhim
margin of eye. Presutural acrostichal setae only slightly
differentiated.

ME: Sternite 5 with shallow anterior invagination arr! with
posteriorarmsextelfiingelnlghtogive lateralnarginsaalrved
appearance rather than straight (figure 10a). Apices of cerci stout and
sinus along interior margin (figure 10b) . Hillae salewhat bulbous at
tip (figure 10c).

m2: Sternite 6, 7, anngoldenpnn’ncse. Stemiteémldl
widerthanlalg. Sternite7with lateralnarginsbroadlyconvergent,
apex with shallow invaginatim (figure 10d) . Sternite 8 large,
minately 1/2 length of sternite 7 ard with anterolateral lobes.
Imes (1941a) describes the spermatherze and accessory glanis. Not
separable frun several other Nectrtpical species.

 

(b)

(a) Male fifth sternite;

mm-

Figure 10: Gelitalia of

(d) Fanale sternites 6, 7 and8.

Male cerci; (c) Aedeagus:

71

IARVA: 'nlefirstinstarisdescribedbylopes (1982b).

WIS: 'Ihenearly parallel mrgins cfthe frcntalvittaandthe
hflbousappearameofthetipofthehillaeservetosqaaratenalesof
misspeciesfrunctherNectropicalm.

TYPES: Male holotype, Sao Paulo, Brazil (IOC?).

MATERIAL EXAMINED: 1 male, 2 females (la—used for figures 10a-d)) .

All quecimens frun S.Paulo, Brazil, Brune Pchl collector, and determined
by 11.8. Iqaes.

DISIRIEJTICN: Argentina, Brazil (Ceara, Mato Grosso, Goias, Minas
Gerais, Rio de Janeiro, Guanabara, Sao Paulo, Parana), Oolunbia,
Paraguay, Trinidad. Distribution fran Lopes (1969, 1974).

BIOLOGY: Lopes and Kano (1968) provide a' discussion of the
relatiaship of the various parts of the male and female genitalia of
this species during cqaulation. 'Ihe specimens that they used for this
shady were reared on lumen feces.

W map-
(Figures 11a - d)
W11 n.sp. Type-locality: Kenmdcy, Maria: 00., Man's

72

mm: length: 8-12nm. Genaandparafacialsgreypruinose.
Genamayhave atirgeofyellctvpruixnsityrearmrgin atgenal groove.
Anterior acrostichal setae differentiated, usually with 4 pairs. 'Iergite
Sgreypnlinose, atnnstwithorangegrumdcolorrestrictedto
postercventral margin.

MAIE: Sternite 5 with long, thin posterior ants (figure 11a) .
Apices of cerci with nearly straight interior margins, not sinuous
(figure 11b) . Hillae similar in amearance to a right triangle (figure
11c).

FEMAIE: Sternites 6, 7, anngoldenpruinose. Sterniteémxzh
wider than long. Sternite 7 with lateral nargins broadly convergent
(figure 11d) . Sternite 8 with anterolateral projections.

DIAGNOSIS: The large size, greypminose gena, and reddishgrcmd

coloroftergiteSrestrictedtopostemrentralmarginseparatethis
ispeciesfrunotherNeareticspecies.

TYPES: Male holctype and allotype, KY: Karim 00., Rman’s Imob, 19
my 1985, G.A. mhlan (me-used for figures Ila-d) . 11 male arr] 15
fanale paratypes: KY: Marion 0)., Rowan's Knob, 19 my 1985, 6.1:. Dahlen
(1 male-me, l male-GAD); KY: Holy Cross, Karim 00., 20 May 1985 (1

male-GAD): KY: Nelsm 00., nr. Holy Cross, 25 July 1985, G.A. mhlem (1

73

 

 

 

 

 

 

Figure 11: Genitalia of MILL (a) Male fifth sternite; (b)
Male cerci; (c) Aedeagus; (d) Fanale sternites 6, 7 and 8.

7M

femle-EIC, 1 female-183, 6 fenales-GAD); Imood Hill Park, New York,

N.Y., 8-IX-1962, P.H. mam, Jr. (1 mle—C'AS): New York, NY, mp 24-Ix-
1961, P.H. Arnatd, Jr. (1 mle-QS): (Bltral Park, Near York, NY, l-VII-
1962, an. Arnaud, Jr. (2 males-<25): Oertml Park, New York, NY, 30-VI-
1962, p.11. mm, Jr. (1 male-C35): Omtral Park, Nev York, NY, s-vrn-
1961, an. Arnatrl, Jr. (3 males, 2 females-<25): central Park, New York,
NY, 2-VII-1962, P.H. Arnaui, Jr. (1 tale-CBS): Cantral Park, New York,
NY, 6-VIII-l961, P.H. Arnaud, Jr. (1 fmale-QS): NEW YORK, Riverhead,

L.I., 9.IX.1961, P.H. Amaud, Jr. (1 female-(ES): Maspeth, L.I., Sep. 4,
1927, Schott (1 fanale-FMNH): Tar Hollow St. Eon, Vintcn OO., Chic, 4-

D(-74, Collector G.A. Oocvert (1 female-m1).

BIOLOGY: Urflmam. Several fenales collectedmfreshccwmanure in
Waybytheauthor.

REMARKS: Fatalesofthisspecieswillualallybefcurdmfierthe
namefingorggta. mlesmaybefanflmderm,m,or
m. misqaeciesisnamedintniorcfmymtherardherbrgthersard
sisters,mstofutnnarecurrentlylivinginl<entlnky. Specialthanks
are giventomyamt, M. ale Cissell, whocollected and froze daily
MalaisetrapsanplesfornefrunNelsmOo.,Kmecky.

Wis (Hall)
(Figure 12)

75

W Hall, 1931: 182, figure 1c. Type-locality:
United States, California, Ooadlella (nele). Dodge, 1956a: 186, 188
(key) . Poorbaugh and Limdale, 1971: 55 (biology).

AIIJIII': Length: 6-8nm. leithdorsalsetaeextendingtoberdof
Sc. ‘Iergite 5 orangish along posterior margin.

MALE: Apices of cerci nearly parallel, not distinctly convergent.
Paramere distinctly broadened apically (figure 12) .

FEMAIE: Sternite6, 7, and8verysimi1arinappearancetothatof
mm (figure 37d)-

DIAGNOSIS: 'Ihis species is very similar to M.

TYPE: Male holotype, United States, California, Coachella, smmer
of 1929 (type umber 43265, USIM). Nine paratypes, same locality.

mm:1heauthorrasmtexamiredspecinenscfthis

species.
DISIRIHJTIQY: Appearstcbeveryrestricted. Alllmcwnspecimens
have been collected in California’s Ooadlella ard Inperial Valleys.

Distrihrtim frun material examined and Dodge (1956a).

BIOIDGY: 'lype series collected in traps which were baited with

76

 

Figure 12: Genitalia of 2am 'a coaQfllensis. Male gonopod and paramere
(redrawn fran Hall, 1931).

77

decaying liver, urea, arr! water. Poorbaugh and Linsdale (1971) report
rearings of this species fran dog dun; in Riverside, (hlifornia.

REMARKS: 'merecordofthisspeciesrearedfrundogdtmggivenby
Poorbaugh and Linsdale (1971) is casidered highly questicnable.
Daminatimof largemnbersofmjnia frcnRiversideandother
localities in this general area Of mlifornia leads the author to believe
that this citation is based on a misidentification of 3m.

Altknlghtheauthordidmthave specinensofthisspeciesto
dissectandfigure, paratypeswereexaminedduringatriptothe
BiosystexaticReseardlcentreardflieirgerfitaliaweremtedasdistimt.
The description provided canes frun information provided in Hall (1931)
and Dodge (1956a). The figure Of the paramere is frun Hall (1931).

W (loves)
(Figures 13a - d)
Wiggles Lopes, 1962: 170, figures 19-22. Type-
loality: Oolunbia. Irpes, 1969: 22 (locality records).

awm': Iength:6-9nm. mlpiblack. Gala,genalgroove,
parafacials and frurto-orbital plate gold pruinose. Gena abruptly
dumhgtogreypruimsealalglimexterflirgfrmposteriormrginof
eye. Iegsdarkgreyingromdcolor. Ventralsetulaeofcostaextendirg
pastSctch. leithdorsalsetaeextendirgtcbendofSc.

78

mm: Posterior arne Of sternite 5 lug ard laterally produced
(figure 13a). Apices Of cerci lag and with sinlcus inner margins (figure
13b). Gorplswell developedand minded. Hillae large anibulbous at
tip (figure 13c).

FENAIE: Sternite 6 smoothly carved anteriorly and with a straight
posterior margin, almost senicircllar in appearance. lateral margins of
stemite7divergentinbasalthirdthenconvergenttoapicalmargin
(figure 13d) . Sternite 8 with convex, ovoid, lateral margins.

DIAGNOSIS: 'Ihe cmbination of the wing setation mentioned above,
dark palpi and legs, and distictive genitalia separates this species from
all others.

TYPES: Male holotype. Pasto, Departanento de deinamarca,
Colmbia, 3500 m, A. Unigarro, 29.x.1958 (IOC?). Allotype, Bogota,
Departamento de Ordinanerm, Oolunbia, 14.111.1937. 7 male and 10

female paratypes designated by Lopes (1962) .

mm: 3ma1esand8 females (EC). ParamodePurace,
emu, CDIIMBIA, 26.11.1970, ca. 3400 m., 0.1!. Wood (1 male; used for
figures 13a-c): 16 km. e. Silvia, Cauca, CDHMBIA, 22.II.1970, ca. 2800
m., 0.14. Wood (1 male, 2 females): mm, Pich., 20 km. W. Aloag,
250m. 4.III.76, G.E. Shevell (1 male, 1 fanale): mm, Pid1., 10 km.

W. Aloag, 3000 m., 17.III.76, G.E. Shewell (1 fenale): mm, Napo,

79

    

v/ ’///// I

 

 

 

Figure 13: Genitalia Of W. (a) Male fifth sternite: (b)
Male cerci: (c) Aedeagls: (d) Female sternites 6, 7 ard 8.

80

301cm. W. Baeza, 250011., 5.III.76, G.E. Shavell (1 fenale; used for figure
13d): mom, Napo, 43 km. w. Baeza, 3500111., 4-7.III.76, G.E. Shewell (1
femle): VENEZ., Merida, mi, 10km. E. 'nobay, 2000m., 28.IV.81, H.

Tonnes (2 fenales) .

DISIRIEJTICN: Colunbia, Ecuador, Venezuela. Distribution frcm
material examined.

BIOLOGY: thflcwn.

Mums (1096)
(Figures 14a - c)
W Limes, 19463: 135-137, figures 36-40. Type-
locality: Mexico, Chapultepec. ropes, 1948: 567 (locality record).

AIIIIE‘: length: 7 - 9 m. Pronto-orbital plate golden pruinose. R1
with dorsal setae extending to bend Of Sc.

ma: Mesotibia with anteroventral seta. Fifth sternite with a
basalmedianholeandmesal areasofposteriorarnswithnanysetae,
givirgtleamearameofhuoparallelbrushesinpimed, run-dissected
specimens (figure 14a). Ceroi with apices straight and parallel apiwa
(figure 14b). Aedeagus with large hillae and rounded corpus (figure
14c) .

W:Notfigmedbypreviwsautlnrsaninotseenbytheauthor.

81

   

   

(b)

(a) Male fifth sternite;

Figure 14: Genitalia Of W.

Male cerci; (c) Aedeagus

8’2

Descripticnofallotypedoesmtdismssanydlaractersthatare
sufficient to separate m fenales fran those Of other species.

WIS: 'meappearanceofthefifthsternitecanbeusedto
separatefllisspeciesfrunthesynpatricm,gmm
m. 'melackofproclinateugperor‘bitalsetaeinthemalewill
separatefigmfifrunm. 'Ihestructureoftheaedeagussuggests
affinities with m, which may mean that the fanale genitalia mild
mstresenblethatspecies.

TYPES: Male holotype and female allotype, Mexico, Mexico D.F.,
Chapntepec, May 30, 1941-June 28, 1942 (ICC) . 'Ihree mle and one female
paratypes, Mexico, Morelos, Cuernavaca, W l-Decatber 5, 1941.

mm: 3 nales. Mexico, Dirango, 5mi. W. Diramo,
6500', Aug. 11, 1964, J.F. McAlpine (BIC) (used for figures 14a-c).
Mexico, Gliapas, 25 mi. so. Tinctla, elev. 8-9,000 ft, km 1133 s. DEX.
City, Rte. 190, 10-VIII-62, H.E. Millira'l (EC). Mice, lbrelos,
'Iepoztlan, 8-20-56, R.& K. Dreisback (MSU).

DISIRIHJTICN: Mexico (Chiapas, DirangO, Hidalgo, lbxico D.F.,
mrelos). Distributim frun material examined and Icpes (1946a, 1948) .

BIOIOGY: Urflcnom.

83

Wigs (Walker)
(Figures 15a - d)

Wig; Walker, 1852: 322. Type-locality: United States
(male). Berscn and Wingo, 1963: 254. Sanders and Dobscn, 1966: 957
(biology).

W Coquillett, 1901: 17-18. Type-locality: United
States, Washington, o.c. (male). Howard, 1900: 568-569 (biology).
Board, 1901: 44 (biology). Baker, 1904 (nec (bquillett): 3
(locality records based (:1 misidentificatim?) . Jdmson, 1913: 75
(locality record) .

WW (00915-11337- Greene, 1911: 334
(locality record). Prat, 1912: 181 (locality record and biology).

W (Coquillett). Parker, 1914: 60—62, figures 27, 46
(description of adult, distribution).

W (Ooguillett) . Aldrich, 1916: 249, 296-298,
figure 143 (key, description Of adult, distribution). Hall, 1928:
334, 346, figure 18 (key, distribution, male genitalia). Hall,
1929: 88 (locality record).

WW (Coq1illett). Tomsend, 1917: 195
(Genotype designation). Greene, 1925: 5, 14, figure 20 (biology,
descriptim Of puparium) .

W Aldrich (nec. Walker). Aldrich, 1930: 15, 17

8H

(synonymy). Hall, 1931: 182, figure 1A (cmparism of gcnOpod and
paramere). Knipling, 1936: 420, 421, 432-433, figures 4, 31, 63
(biology, descriptim of larva). Hallock, 1940a: figures 35-37
(male genitalia). Hallcck, 1940b: 208, 210, 222-223, figures 122-
124 (key, distribution, male genitalia). Hohr, 1943: 285-286
(biology).

W Aldrich (nec. Walker). Lopes, 1946a: figure 32
(male genitalia). ropes, 1948: 566 (locality record).

W Aldrich (nec. Walker). Rcback, 1954: 14, 15, 27,

42, 73, 74, figures 215-219 (description Of adult, male genitalia).

William (Walker). 00399 and 59390: 19543 53: 55
(biology). Dodge, 1956a: 185-186, 188, figure 16 (key, description
Of fenale). Reed, 1958: 241 (biology). ropes, 1977: 43, 44,
figures 1-6 (redescription) .

W (Walker). I-Icnser and Wingo, 1967: 731-732. Blume,
1970: 1023 (biology). mm and Knapp, 1970: 167-168 (biology).
Hayes and Turner, 1971: 444-445 (parasitoid record). Payne and
King, 1972: 160 (biology). Moore and Isgner, 1973: 250 (parasitoid
record). Watts and (limbs, 1975: 564 (parasitoid record). marten
ard than, 1979: 81, 87 (key and descriptim of puparium, biology).
Peterson, fl, 1981 (biology). Fig, My 1982: 476
(parasitoid records). Figg, fl, 1983a: 962-965 (parasitoid
records). Figg, My 1983b (parasitoid record).

85

NIJLT: length 6 - 8.5 nm. Promo-orbital plate grey pruimse with
yellowishtinge. leithdcrsalsetaeexterdingtoberdofSc.

m: Occasional specimens will well developed cuter vertical seta.
Fifth sternite U-shaped, with a small, densly setulate, ventrally
projectirgplatenearapexoftheirmermarginoftheposteriorarms
(figure 15a). Oerci with dark, cmvergent apices, resenbling a crab's
claw (figure 15b) . Aedeagus long, with darkly sclerotized and angular
hillae (figure 15¢) .

FEMAIE: Sternite6verylargeandconspialms. Sternite?
completely divided with the lobes situated laterally. Sternite 8 and:
smaller than the preceeding genital sternites (figure 15d).

IARVAE: Knipling (1936) provides a description Of the first instar.

HIPARIUM: A description Of the puparium is provided by Greene
(1925) and marton and Moon (1979) .

DIME: 'Ihe claw-like cerci is the easiest character to use to
separatethemalesofthisspecies franallothersoveritsrange. 'Ihe
aedeagusisalsoverydistinctive. 'nlelargesixthsterniteofthe
fenale is distinctive in pinned specimens and will allow identification
without dissectim. In dissected specimens, the cmpletely divided
seventh sternite is unique to M.

TYPE: Male lectotype. U.S.A., ex coll. Saunders, 68.4: =
m cog” det D.J. Clark, 1953 (without head).

86

 

Figure 15: Genitalia Of W. (a) Male fifth sternite: (b)
Hale cerci: (c) Aedeagus: (d) Female sternites 6, 7 and 8.

87

MATERIAL EXAMINED: 533 mls ard 642 fenals (52C, (38, (SU, INNH,
GAD, H2, 160, m, (BU). FLA: Highlands Oo., Archbold Biol. Sta. 9-X-
1964, P.H. Arnaud, Jr. (ms-male; used for figures 15a-c). Laguna
Atascosa Ntl. Wildlf. Ref., m 0)., m., 18 March 1972, R.K. Zaidel

OHJ—fsale: used for figure 15d).

DIS'LRIEJTICN: Canada (Q1tario), United Stats (Alabam, Arkarsas,
Colorado, Connecticut, Deleware, Florida, Georgia, Illinois, Indiana,
miss, Way, Louisiana, Harylard, Massachusetts, Michigan,
Minnesota, Mississippi, Missouri, Nebraska, New York, North Chrolina,
Chic, Oklahata, Penrsylvania, South Carolina, South mkota, Tennessee,
Texas, Virginia, West Virginia, Wisconsin), Mexico (Michoacan, Nayarit,

Tabasco, Veracruz) .

BIOLOGY: A cmum coprqmagws specis. BM has been
reared m numerous occasicrs fron cow mamre (I-Icward, 1901: Pratt, 1912;
Greene, 1925: Knipling, 1936: m, 1943: Bensm and Ningo, 1963:
Sanders and Dobscn, 1966: Kaiser and Wingo, 1967: Blume, 1970: marten
andlbm, 1979). Itisalsoreportedbychard (1900) asaieofthemost
abundant flis bred and captured frun mnnan malt. Rhipling (1936)
wasunsuccssful inhisattetptstorearthisspecismdecmposing
mats. Ithasmtbeenrearedfrondogdlmg.

'Ihe life cycle, frcn freshly deposited, first instar to adult was

88

recordedbymiplirg (1936)totakelssthan11daysmccwmn1re.I-le
fanflthatthelarvaewerebegirmirgtotrarsformtothethirdinstar
after21 hours. Ripariationbegan lssthan4 daysafterlarviposition
ardachltsbeganenerging6daysafterplpariatim. Howard (1900)
reportsthatthisspecisdevelmedfrmfirstirstartoadultin6days
mmnnanexcrenent. Henctsthattheeeatherwasverywarmtheaverage
tsperature being estimated at 90° F.
'memaximnnrnmberoffirstinstarsthatwasdissectedfrouasingle
fanale is 30 (Knipling, 1936), withmst gravid fenals havingnuch
fewer. Estimationsof feonditybasedmnmbersoflarvaecmtairedin
pinned secimens of this specis is difficult, due to the emulsim Of
larvaefrongravidfenalsastheydieinakillirgvial (personal
daservaticn).
'I‘ramingrecordsofadultsincludeDodgeardSeago(1954),w:O
collected 100 nels and 316 ferals at several sits in Georgia in traps
baitedwithd1ickenentrailsand,sanetiues,rnmanfecs. Emmeland
Knapp(1970) collected largenmbersof secimensthrargtnrtthemrths
omee,July,N.:gustardSepte1beratseveralsitsinKentuckyin
mdified USDAtrapsmidlwerebaited with liver, banana, beer, ormalt
mixture. Reed (1958) collectedthisspecisfrondogcarcasss. Payne
ardKirg(1972)listsB,_mligtaasascavergerspecismbabypig
carrimwhidiwasplacedmthescilsrrface,intres,ardinwater.
'mestudysitswerenotdscribedhltareassmedtobelocatedinScuth
arolina. Adultshavealsobeencollectedinanalaisetrapina

89

grassy/stmlbby field ((11, Hartgcmery OO., Huber Heights, Jule-July, 1975,
G.A. mhlen). Orespecinenfronaesuriyareabaitedwithmss
pellets, a bait toxicant for W, was collected by Peterson, e_t
a1, (1981).

Mals have been noticed exhibiting station-taking behavior at two
general sits, at snall animal carcass and at amivore durg. Station-
takirgtaksplaceintheafternomintheam. Informatimonmale
staticn-taking behavior cans fron biological nots taken by Dr. William
L. Downs, Jr. during the sunlner Of 1964, mainly at IL, Mason Co., Mason
State Forest (EF) . Male station-taking behavior is highly deperriant on
exact site and time of day. Therefore, the following information is
provided for future, corparative studis. Times given are Central
Daylight Tine, # refers to W.L.D. Bio. Note timber, and locality is DEF,
unlss otherwise noted. (:1 June 8: nine mals, 2 fenals were collected
at a fresh fieasant carcass (6:45-7:00 P.M, #86). On June 9: the male on
racoon dung (11:45-12:30 P.H., #99): cne male at pheasant carcass (1:25-
1:40 P.M., #102): (me uale at pheasant carcass (2:40-2:55 P.H., #109):
two nels, one fenale at pleasant carcass (5:45-6:00 P.M., #111). On
June 23: are male at ground squirrel carcass (1:30-1:45 P.M., #176); two
mals, 1 female at fresh human fecs (1:45-2:00 P.M., #177); five mals
at lumen fecs (4:45-5:00 P.M., #183). 0: June 24: three mals, 1 female
at day-old lumen fecs (2:37-2:52 P.H., #205); one male at olrtle carcass
(Spring lake State Park, IL 6:40-6:55 9.14., #213). On June 25: three
mals at 2-day Old lumen fess (8:45-9:00 14.14., #218): we male at 2-day

90

Old lumen feces (3:45-4:00 P.M., #231). 'Lhese specinens are located in
Dr. Dames' persa'el collectim.

Wmtobemitesecretiveduringnetirg. Ofthe
1175 specimens examined, no correctly associated mating pairs were
iniicated. DodgeardSeago (1954) nentionstheumsualcaptureofa
netedpairinalighttrapandproposstletthediurrel copulationwas
mtmmletedbynigtrtfallardttetrpmbeirgdishm‘bedtlecwple flew
to the light. A nele Of dereligta, amarently attelpting copulation with
afanaleofmm, wascollectedintallgrassadjacenttofresh
lumen fecs m a flat rock in a grassy field (KY, mrion Co., Holy Cross,
19 Hay 1985, 2:00-3:00 P.M., G.A. mhlen). 'Lhe specimers quarated in the
killing jar.

Potential parasitoid records (records for m or carplexs of
aerial: species which Mode Resellers) include: inseam Say
(F199: Eli—3L: 1982: 1983a) and met—Am (Say) (Benson and
Wirqo, 1963: Homer and Wingo, 1967: Hays and Tuner, 1971: Fig, g
a]... 1982, 1983a, 1983b) (Hymempter-a: Bramlidae): Emile Sp. (Fig, 521:
' a]... 1982) and W (say) (Hayes and m. 1971)
W: CYn-ipidae): limits 59- (F199. steal“ 1982) and Missing
muse (Say) (Hayes and 'I'tmner. 1971) (W: Figitidae):
m sp. (Figg, 95.9.11. 1982, 1983a) (Hymenoptera: Diapriidae):
W (m) (F199. £5.31." 1983a). my: BP-
(Fiqg. stall. 1982). mm 89- (F199. steals. 1983a). Hemmer
Girault ard Sarders (Fig, 93.51“ 1982),

 

91

(P199, 341,, 1982, 1983a), m Iatr. (Figg, $.81.” 1982, 1983a)
and W (mus (Figg, 93414, 1982, 1983a) (1mm:
Ptemlidae); and 31m sp. (Hayes and m, 1971; F199, §t_a;_._,
1982, 1983a) and W L. (Moore and Iegner, 1973)
(Cblecptera: Staphylinidae).

Rum: misspeciesmayhefanriinccllectionsmnderttemmes
M. 99mg. or____stimlarIS. deperflingmmenthe
determination was nade and which systanatist trade the identification.
'Ihere are records in the literature that indicate that this species
ocmr'sinvesternstatessudlasIdaho, Oregon, Utah, Nevada, and
mliforniaasvellasseveralmrribean islands, hittheauthorhasseen
noevidence foraxhclainsanditisbelievedttataxhreccrdsare
based m misidentificatims.

W n-SP-
(Figures 16a - d)
W1. ma). Type-locality: United States, Texas, 23 mi. W.
Ft. Davis, 5000' (male).

mm: Isngth:5.5-6.5 m. Erato-orbital plate yellcwpruinose.
leithdorealsetaeexterdingtobaflinSc.

m:1wpreclinateu;perorbitalsetaepmesentardwelldevelqaed
wtervertical setae. Mesotlbiawithanterwentralseta. Fiftl'lstenfite

92

wiflmtbasalnesallnlehltwitlxmsal lobes setulate, amearirgasa
pair of parallel brushes in non-dissected specimens (figure 16a). Oerci
small with apices parallel in posterior view (figure 1&3) arri apices
slightly curved anteriorly in lateral View. Aedeagus very distinctive
for a species having dorsal setae (11 R1 (figure 16c).

ms: Sternitess, 7, andsnearlyequal inwidtharrlstemites
with lateral, ovaldepressionsmidigivetheraisednnsal areaan

hourglass appearance (figure 16d) .

DIAGNOSIS: The proclinate upper orbital setae will easily separate
themalesofthisspeciesfranall others. Fanaleverysimilartc
m but with lateral oval depressions on the eighth sternite giving
themsalareaanl'mrglassappeararce. 'misdaaracterisverymticable
even in pinned, I'm-dissected specimens.

TYPES: I-blotype male - 23mi.W. Pbmvis, 1.VI.1959 TEXAS, 5000’
FJrAlpine (32C; used for figures 16a-c) . Allctype fatale - Santa Cruz
Co., ARIZ., 13 mi SE Chitirental Nadera Canyon, elev. 5500 feet, 20 June
1965, R. a J. Matthews (IBU; used for figure 16d). Fair male and four
fanale paratypes: s'ta. Rita uts., fangs Res. Ariz., Aug. 18, 1930,
Ar.L.P. Wehrle (male-52C); Big Bend N.P. m, mgout Wells, 3000’, May
13, 1959, J.F. malpim (me-me): USA 'l‘exas Big Bend N.P., panther
mum, 103°12'w 29°l9'N, 3.VIII.1982 1130111., 3.15:. O'Hara (me-mo):

mm, GralmOc., 0.9 mi. alongroadtouarijilda mnym franHwy.

93

 

Figure 16: Genitalia of W. (a) Male fifth sternite; (b)
Male cerci; (o) Aedeagus: (d) Female sternites 6, 7 and 8.

9L)

666, Alt. 3860', 3.VIII.1965, collects}: 11.8. Iesd'l (male-(ES). Sane
locality as allotype (2 fanales-MJ). ARIZ.: Portal, Cbchise Co., 3-10
July 1978, mane Flynn (2 females-960).

BIOLOGY: Unknown. Allotype farele contained 16 first instars which

wereverysimilarinamearametoMlarvae.

W: 'mefanaleofthisspecieshasmt,asyet,beendirectly
associatedwimthenele. 'Ihisspeciesisnamedinhonorofnr. William
L.Dmmes,Jr.wtnrq3eatedlyhasranirdedmeofexceptimstothemles
in the Sarccphagidae. 'Ihis species certainly exhibits diaracteristics
thatwerenctexpectedtobefamdinspecieswithdorsalsetaemnl.

W (Walker)
(Figures 17a - d)

W Walker, 1861: 309-310. Type-lowlity: Mexico
(male). Aldrich, 1930: 20 (synonymy). Hall, 1931: 181 (key).
W Wulp, 1895: 272. 'Iype-lomlity: Mexico (nele and

fanale).
W Wulp, 1895: 272. 'IYpe—locality: lie-(ice (male).
W Hall, 1928: 345-346, pl. 22, figure 17. 'Iype-
locality: Janaica (male/fanale, maiting pair).
WW (Walker). Icpes, 1946a: 131 (locality record).
ches, 1948: 566 (locality records). Dodge, 1956a: 188, figure 18

95

(fenele genitalia). Dodge, 1965b: 475, 484 (key, lomlity record).
Dodge, 1965c: 498, 503 (key, locality records). Rdiderdorf &
Gregor, 1973: 8 (key). Lopes, 1975a: 485 (loca1ity record).

WM (Walker). Rcback, 1954: 11, 13, 26, 73-74, figures
227-229 (male genitalia).

mm: length: 6 - 9 mu. Harte-orbital plate grey pruinose with
yellowishtinge. leithdcrsalsetaeexterriingtobenlofSc.

MALE: Fifth sternite U-shaped, with ventrally produced, setulate,
medial ldaes m posterior arms (figure 17a). Oerci with lcng, thin,
parallel apices (figme 17b) . Aedeagus with juxta well develcped and
very laig vesica (figure 17c) .

W: sternites 6, 7, and8verydifferentinshapefrunone
anther. Sternite 7 with apical irdentatim ard sternite 8 maller in
width than preceeding sternites and situated in indentation of sternite
7 (figure 17d).

WIS: 'nielaigapicesofthecerciandthestnictureofthe
male genitalia (especially noticeable is the lag vesica) serve to
separatemales ofthisspeciesfranall others. ‘Iheangle ofthe fronto-
orbital plate as seen in lateral view is usefull for separating this
species frcm mm (see figure 15a) collected in Madden localities.
'Iheshapeofthegmital sternitesisuniquetothisspecia, but
dissecticm is often necessary for determinations.

96

 

Figure 17: Genitalia of W. (a) Male fifth sternite: (b)
Male cerci: (c) Aedeagus: (d) Fanale sternites 6, 7 ard 8.

97

TYPES: Male holctype, Mexico (EN-I).

mm EXAMINED: 122 males and 76 females (sac, (as, ucz, MSU,
CE], WID). FLA: Highlarrls Co., Archibcld Biol. Sta., 8-X-1964, P.H.
Armand, Jr. (male-ms: used for figures 17a-c). MEX.: Sonora, Alamos,

25-II-1963, P.H. Arnaud, Jr. (female-C38: used for figure 17d).

DIS'IRIEJ'I‘ION: United States (Arizona, California, Florida, IDams) ,
Mexico (Baja California Norte, Baja California air, Chihuahua, wrango,
Guanajuato, Guerrero, Jalisco, Mexico D.F., Morelos, Nayarit, Oaxaca, San
Inis Potosi, Sinaloa, Sonora, Veracruz, Yllcatan), Guatanala, Honduras,
Costa Rica, Panama, Cblunbia, olba, Jamaica, Puerto Rico, Barbados,
Trinidad, Bahamas.

BIOIDGY: (be male swept fran flowers W (USA: ARIZ.:
Pima Co., Organ pipe draws Nat. Mcn., Williams spring, 15-II-197o,
A. Gray, (15) . (be male collected at flower W (MEXICI):
W, 17 road miles W. arrango, 10-IX-1966, 6600’, D.E. Breedlcve,
(ES). are fatale collected at fir W (MEX.: Sonora,
Alamos, 21-II-1963, P.H. Arnaud, Jr., GS). Haidmm umber of first
instarsdissectedfrunonefanalewas 13. Firstinstarwithdarkly

pignerrted psaflctradiea.

98

W: Altlnlghthereareseveralsyrrmynnisnames,virtuallyall
determinedspecimersstmldbefandmflerthenamem.

mm (mlp)
(Figures 18a - d)

W thlp, 1896: 278. Type-locality: Mexico, Tabasco
(4 miles). Aldrich, 1930: 33 (synmymy).

W Hall, 1928: 345-347, pl. 22, figure 20. Type-
locality: United States, Texas, College Station (male). Van Brien,
1950: 199 (parasitoid record).

William (mlPL 10936: 19458: 131 (locality record).
Lapse, 1948: 566 (locality records). Dodge, 1956a: 186, 187, 188,
figure 19 (key, female genitalia).

mm (Wulp). Rcback, 1954: 74, figures 230-232 (male

genitalia) .

mm: length: 5 - 11 m., usually 9 - 10 m. Pronto-orbital plate
bend of So.

MALE: Mesotibia usually with anteroventral seta. Fifth sternite
with cmspicuous mesal lobes with several long setae (figure 18a). Oerci
large with apices slightly cawergent (figure 18b) . Aedeagus large, with
sell developed corpls, short vessica, and partially hidden hillae (figure

18¢) .

99

ms: StemitesGarri7mar1yeq1alinwidth, sternite8
slightlynarrower (figure 18d). Sternite7nld11argerthanmrmmding
sternites andcaspicuous, even inpirmed, non-dissectedspecimens.

DIAGNIBIS: The male genitalia are very distinctive and
determinationscanusuallybemadeeveninrm-spreadspecinensby
ermininaticn of the large setulate lanes of the fifth sternite. 'Ihe
genital sternites ofthe fenaleareusuallyveryapparentand
distinctive, thus allowing determinations of non-dissected specimns.
'meaedeagtsappearstohavenanyaffinitieswiththatofm,
particularly in the structure and amearance of the hillae.

TYPES: Eourmale syntypes, Mexico, Tabasco (HM).

MAL ram: 148 males and 68 feneles (mac, as, m, ncz,
MSU, osu, WLD). Fairbanks Sprs., Ash Meadows, Nye Co., Nev., 20(vi)51,
IaR. and ‘Ied Frantz (male-GB; used for figures 18a-c). Aguanga, Cal.,
4/28 '33, A.J. Basil'qer Collector (fenale-(ES; used for figure 18d).

DISIRIEJTIQJ: United States (Ariza'ia, California, Kansa8,
Louisiana, Nevada, New Mexico, Texas), Hexicc (Baja California Norte,
Chiapas, arrange, Hidalgo, Jalisco, mica, Mexico D.F., Horelos, Oaxaca,
Riebla, San Inis Potosi, Sabra).

100

 

 

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Figure 18: Genitalia of W.

(a) Male fifth sternite; (b)
Male cerci: (c) Aedeagus; (d) Petals sternites 6, 7 and 8.

101

BIOLOGY: Van Brian (1950) lists W as a parasitoid of
W mls. (Colecptera: Ooccinellidae). Sixteen males
ard seven females were collected byAJ. Basinger at human went
((3/2) San Jacinto, Cal., 5/18 '33: (0/1) Aguanga, (21., 4/28 '33: (2/0)
Riverside, Cal. 6/20 '33: (0/1) Riverside, (21., 11/23 '33; (1/2)
Riverside, Cal. III-8-34, Oottcndood can: (7/0) San Andreas Can., Riv.
0)., (21., 1.2/20 ’33: (0/1) San Ardreas Chm, H-21-33, Riv. Co., (31.:
(2/0) Indio, (21., 11/24 '33; (1/0) Chino, (21., rv-20-34). mo males
and five females were collected by A.J. Basinger at an orange dtmp
(Riverside, Gal. 3/27 ’33) . are nele and four females were collected by
A.J. Basinger at fresh dlicken guts (case Grands, Ariz. III-25-34). me
male and one fanale was collected by A.J. Basinger at arr-lids (Dcwney,
Cal. 11/27 '33) . Five males and one fanale was collected by A.J.
Basinger at fish bait ((1/1) Riverside, on. 5/19 '33; (2/0) Riverside,
(21., 12/20 '33: (1/0) Riverside, (21. 5/4 '33; (1/0) San Andreas Can.,
Riv. Co., Cal., 12/21 ’33 ). One rule was collected by A.J. Basinger at
fish (Riverside, (21., ll/26 '33). me male was swept from flowers of
WA. Gray (USA: ARIz: PimaCo., OrganPipeCactusNat.
Morn, Williams Spring, 17-11—1970, P.H. Arnaud, Jr. ). one male was
collected at flowers of mm (MEXICI): Orange, 17 road miles
W. mrarrgo, lO-IX-1966, 6600', D.E. Breedlove). One rule was mllected
in a light trap (CAL. Mom Co., 'Ihe Rot Spgs., 2.5 mi. 8. of Bridgeport,
15-VIII-l963, 11.8. leech). All of the above menticmed specim'ns can be
found in the collectim of the Chlifornia Acadany of Sciences.

102

REMARIG: Virulally all determined specimens in collections will be
found mrler the name m.

W (Aldrich)
(Figures 19a - d)

W Aldrich, 1916: 249, figure 117. Type-locality:
Miami, Florida. Hall, 1928: 334, 336-337, figure 4 (key,
redescription).

Wig (Aldrich). Roback, 1954: 74, figures 235-236
(syncnymy, male genitalia). Downes, 1965: 954 (locality records).
Dodge, 1965b: 491 (locality data). Hades, 1969: 21 (locality
records). Rohderriorf and Gregor, 1973: 7 (key, locality records).

Arum: Length7-llnm. Ganagreypruinose. Pedicelandpalpus

bright orange. Presutural acrostiohal setae well differentiated. legs
orange in ground color.

‘ MALE: lateral nargins of sternite 5 divergent and slightly octave.
Sternite 5 length anaroxinately equal to width at apex (figure 19a).
Apices of cerci stout, with nearly straight interior nargins (figure
19b) . Hillae rourded apically, exterrling fran corpus at nearly a right
angle anteriorly (figure 19c) .

FEMALE: Sternite 6, 7, and 8 with scattered golden pruinosity.
Sternite 6 mad: wider than 1mg. Sternite 7 with small anterolateral

103

processes (figure 19d) .

DIAME:'n1eorangepalpiardlegscmbinedwiflltheladcof
d3rsalsetaemR1,allowseasyseparationofthisspeciesfrmall
others.

TYPES: Male holotype. Miami, Florida, C.I-I.'r. Townserd collector

(USNM). Allotype fran same locality (USNM). Six paratysz frauvarious
localities (Aldrich, 1916) .

MATERIAL mm: 6 males and 5 females. Tifton, Ga., Oct. 16 ’96
(rum-male paratype: used for figures 19a-c). St. Petersburg, Fla.,
Snell Isle, Nov. 8 1967, Peter J. mrtirlat (MSU-1 fenale: used for figure
19d). Mayport, Fla., June 16, 1954, R.L. Fischer (rm-1 female).

Tiftcn, Ga., June 13 '96 (FMNH-nale paratype and 1 fanale). St.
Mustine, Fla. (FUNK-male paratype). Tifton, Ga., Oct. ’96 (FUNK-1
female). Orlarric, m., 22 June 1981, m. Domes, Jr. (Win—2 males).
Sarasota Co., Fla., 6-30—1952, col. L.R. Remer (WID-l female).
Gainesville, Fla., 17.VII.55, R.A. Morse ccll., at dung of man (WID-l
fanale).

DIS'IRIEI'I‘ICN: Bahama Islands, United States (Florida, Georgia).
Distrihrtim fran material examined ard Dodge (1965b).

101+

  

fin.
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Figure 19: Genitalia of W. (a) Male fifth sternite:
(b) Male cerci: (c) Aedeagus: (d) Fenale sternites 6, 7 and 8.

105

BIOIDGY: Uruomn. One fanale examinedwas collected at humandurg

(see above for locality data).

REMARKS: Virtually all determined specinens in ccllecticns will be

fcundurrierthenameflgriglgis.

W (Aldrich)
(Figures 20a - d)

W Aldrich, 1916: 249, 299-301, figure 145. Type-
locality: Cuba, Havana (Hale). Hall, 1928: 334, 346, figure 19
(key, male genitalia).

W (Aldrich). Roback, 1954: 74 (synaiymy).

WM (Aldrich). Dodge, 1956b: 99, figure 13
(descripticn of fanale). Icpes, 1969: 23 (locality records).
Rohdendorf and Gregor, 1973: 8, figure 18 (key, male and female
genitalia).

AIIJIII‘: Length: 6 - 9 m. Fronto-orbital plate grey pruinose with
yellowishtinge. leithdorsalsetaeextendimtoberdofSc.

ME: motibia with small anteroventral seta. Fifth sternite
without anterior nasal hole, generally Y-shaped, and with blunt mesal
lobes m posterior anrs (figure 20a). Cerci with nearly parallel apices,
my be slightly cmvergent at tip (figure 20b). Tip of aedeagus very
cmpact, vesica strongly bent upwards (figure 20c) .

106

m: Fifth tergite orange, barely visible frun above, and folded
roof-like nedianly ammd genitalia. Sternites 7 and 8 apparently fused,
some irriication of lire of fusion visible. Sternite 6 rearly equal in
size to sternite 7+8. Sternite 8 slightly smaller in width than
preceeding genital sternites (figure 20d) . Posterior margins of
abdaninaltergauaiallywithathinbardofbrcmpnrimsity.

DIAGNOSIS: 'ne dark legs of this species easily separate it frun
Marimba. 'Itecerciarristnicmreoftleaedeaguswillseparate
this species fran tle synpatric gm. 'ne folded fifth tergite,
similartottetseeninmnynmbersoftlegemng,easily
separates the female fran all other species. mile females ofhgmi
alsoshcwafusimcfsternits7arfl8,ctlerd1aracterscffleadults
do not indicate a close relationship.

TYPES: Male holotype, Havana, Olba (type no. 20575, USNM). Fanale

allotype, Havana, Cuba (USNM). 'Iwerrty-a'e paratypes frun sane locality.

m m: 5 mles ard 3 females. Chlabazar, Qiba, IV-14-
23, J.S. Hire collector (OSU-male; used for figures 20a-c). (alabazar,
QJba, IV-30-23, J.S. Hire collector (CEO-2 females; ae specimen used for
figure 20d). Rep. Daninicana, Ia cmbre, 300m, 22-23.III.1978, L. Masner
(me-2 males). Havana, Cuba, Baker (rm-paratype nale; damaged specinen,
mly thorax mine (:1 pin). Havana, Cuba, Baker, H.E. Snith Coll. (MCZ-

107

 

Figure 20: Genitalia of W. (a) Male fifth sternite; (b)
Hale cerci; (c) Aedeagus; (d) Femle sternites 6, 7 and 8.

108

male). Harianao, Cuba, V-13-23, J.S. Hire collector (CSU-fenale).

DIS'IRIHJTIGJ: CUba, Hispaniola, Puerto Rico.
BIOLOGY: m.

minia trematodes (beigen)
(Figures 21a - d)

W Heigen, 1826: 29. 'Iype-lccality: ?Germany, Aaden.
W, 1834: 225. W, 1838: 650. Walker, 1849: 820
(synonymy). Sdiiner, 1862: 572. Pardelle, 1896: 176. Villeneuve,
1900: 364 (type information). Bezzi, 1907: 481. Bottder, 1912:
707 (redescripticn). Ehderlein, 1928: 24 (systematics). Hardy,
1936: 90 (locality records).

W antlers, nec. Harris, 1780: 84.

W Fabricius, 1794: 315 (preoc. by Gnelin, 1790). Type-
lccality: Demark.

Wig Fallen, 1817: 237 (preoc. by Villers, 1789). 'Iype-
locality: Sweden.

MW Robireau-Desvoidy, 1830: 346. Type-locality:
France.

W Robireau-Daevoidy, 1830: 346. Type-locality:
France.

109

W Robireau-Desvoidy, 1830: 346. Type-locality: Frame.

W Robireau-Desvoidy, 1830: 346. Type-locality: France.

W Rdaireau-Dasvoidy, 1863: 427. Type-locality: France.

W (Meigm). Robireau-Desvoidy, 1863: 435-435
(redeecripticn, genotype placement).

W Robireau-Desvoidy, 1863: 436. 'Iype-lomlity: France.

W Robireau-Desvoidy, 1863: 437. Type-locality: France.

W Siebke, 1877: 94. Type-locality: Norway.

mm (Fabricius) . lundbeck, 1927: 169, 171-173
(redeecripticn, locality records, key). Patton 8 Wainwright, 1935:
346. Salan, 1935: 225. Seguy, 1939: 58 (synonymy). Seguy, 1941:
153 (biology). Harmer, 1942: (biology). Kano, 1950: 825, figures
8-10. Ztmpt, 1965: 108 (biology).

WM Enderlein, rec. Fabricius. Enderlein, 1928: 42
(W Film) .

Wig Seguy, 1935: 5. Type-locality: Chad.
merrier; (Fabricius). meander-f, 1937: 391. Roback, 1954: 74

(synonymy). Rohdendorf, 1960: 790. lehrer, 1961: 534-535
(redescriptim, locality records). Zakharcva, 1966: 161, 162
(biology). Kano, M14, 1967: 5, 130-132, figure 74
(redescription, locality records, biology). Slaneckcva, 1969: 225
(locality records). Iopos, fl, 1977: 563 (locality records).
Rdfleniorf and Verves, 1978 (biology). Verves, 1980: 355-356

(locality data, biology). Sugiyama and Kano, 1984: 347, 348,

110

figure 8-11 (syteratics, male genitalia). Verve, 1986: 133-134
(M. lowlity records).
WM Seguy, 1938: 43. Type-locality: Morocco.
mm; antlers, rec. Harris. lbgne, 1986: 16 (locality records,
biology). Pape, 1986: 303, 304-305 (reexandnation of type). Pape,
1987a: 45 (locality records). Pope, 1987b: 99-100, figures 206-209
(redecripticn, locality records).

ADJIII‘: length: 5-8nm. Genagreypruinose. 'Ihreewelldevelcped
postsumral dorsocentral seta.

MAIE: Sternite 5 nuch wider than long (figure 21a). Apice of
cerci convergent, inter margins noticeably bent (figure 21b) . Hillae
very broadly rounded anteriorly (figure 21c) .

FEMAIE: Sternite6veryrumdedinappeararee, mostlycoveredwith
grey pruinosity. Sternite 7 very distinctly elaagated, appearing as a
lcng rectangle (figure 21d) .

Dmm:misistlemlyspecieofm1mwntooccurin
fleOldWorldardithasnctbeenrecordedfrunanylccalityinNorthcr
SouthAnerie. 'negenitaliaaredistinctivearriwculd alloweasy
separatim frunctlerspecie if it shouldbecollected intleNaonrld.

TYPES:Ma1etelctypearriallotyperepresentedbymatimpairm
sirgle pin. No locality data acompanie speoinens, but they are

111

 

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Figure 21: Genitalia of W. (a) Male fifth'sternite; (b)
Male cerci; (c) Aedeagus; (d) Fenale sternite 6, 7 and 8.

112

alspectedofbeirgfrancemny,Aadm. Bothspecinensaremissing
tleirabdanens,malemissinglwing. mseparatepinbeidethespecinen
isawingardabdmen. Botheecinerearedefinitelym. ‘Iype
vereeuamiredin1986atmsemnNatimal D’HistoireNa‘hnelleinParis.

mm W: s nale, 4 fenales. INDIA: mesoorie, 0.13., 15
May 1962, J.P. Donahue, elev. 5400 feet (rm-male: used for figure 21a-
c). Sane locality, 16 May 1962 (lBU-l male). Cypern, Ayia Napa, 10 km
w. Capo Greco, 13-23.vi.l983, B. Petersen leg. (wm-l male, 2 females; a
farale used for figure 21d) . Morocco, Wener coll., 13.V.30 (Hz-mle) .
MAND DI': Warden-Corie m., 26.vii.54, (LA. 5: D.J. Clark (WID-l Hale).
Iyrrtm, 16.VIII.41 (WID-l female). Moccas, 9.VIII.56 (mo-1 fanale).

DIS'IRIEJI'ICN: Afghanistan, Albania, Algeria, Austria, Azores,
Belgium, Bulgaria, canary Islands, Chad, China, cypns, Czechoslovakia,
Dermark, Egypt, Finland, France, German Federal anblic, Germn
Danocratic Republic, Greece, mmgary, India, Iran, Iraq, Ireland, Israel,
Italy, Japan, Korea, Iebamn, Libya, )hdeira, Malta, mlgolia, Morocco,
Nepal, Norway, Pakistan, Poland, Formgal, Imania, Saudi Arabia, Soviet
Union, Swain, Sweden, Switzerland, Syria, 'mnisia, United Kingdan, Yemen,
Ylgoslavia. Distribution from Kano, eLaJ... (1967), ropes, egg,

(1977), and Verve (1986).

BIOLOGY: Depite tie large volume of literature that has disassed

113

this specie, little is lawn of its life history. Kano, M, (1967)
reportsthatthisspeciebreedsinhmanfecearddeadanimalsin
nature. Apossiblecaseofintetimlnyiasis causedbythisspecie is
mm by Zunpt (1965). Verve (1980) nentlcns that the larvae
develop in feoe, ooesialally develcpirg in dead animals and that it is
recorded as a parasitoid of Gastrqloda. Seguy (1941) and Ridendorf and
Verve (1978) report this specie being reared frun snails, grasshoppers
and a beetle. Zakharcva (1966) observed that fanale lay larvae in
ovarian marbrane wten a suitable substrate is available. 'Be larvae
imediatelydistardtheemalbrane, whidlrenaincntrealrfaceoftle
substrateandburrwintotlealbstrate. Hefcundapctentialfertility
ofupto52 larvaeandthat fanale laidamaximnnofthreebatdleof
larvae. Harmer (1942) describe tte mating behavior of this specie
withtleobservatimthatcoupleplraeeachctterintotleairbefore
‘bnblirg to tie ground.

REMARKS: 'misspeciehasbeencitedurriertlenaneofgtniatg
since tie publication of Inndbeck's (1927) synonymy of tie Fabricius rare
with Heigen’s M. It was discovered fairly recently that m
was preoccupied by Gnalin's (1790) W. Pape (1986) reenamired
thetypecf Fallenarrifan'dthatmmifiarpliedtofllis
specie. 'nlisnanehasbeenusedextensively fortiespeciencwlmown
eW. Fbrtmately, ttenaneisuravailableasitis
preoccupied by Villers' (1789) W. Several authors

11H

begantouseHar-ris' (1780)nanemmixforthisspecie. NoHarris
typerenainarrisuldyoftledecriptimteprcvidegivemindication
ttetmnfizisauberofm. nwermre,mstPalearcticspeoie
of Sarcoplaginae have black genital capsme ard Harris specifically
nentiaetleredgenitaliaofamtlerspecieofSarcqhagidaethatte
decribed,hltdidnotnentimredgalitaliainhisdecriptimof
m. FortlesereascnsIbelievethatttenanepgerstmldmtbe
applied to this specie and tie rext available nane, M (Meigen)
shouldbeused. maturation of the typedefinitely associatethis nane
withthisspecie.

Mostdetermiredspecimensincollectimswillbefamdmmdertle
nanegtriata.

E . . l 'I! i 153385
(Figure 22a - d)
W ropes, 1975a: 485-486, figures 1-4. type locality:
Costa Rica, Oerro de la nerte, 6 km W. Villa Hils, Inter-Am. H'wy,

San Jose Pvnce, el.: 3340 10.

mm: Iength:6-9nln. Anteriormarginofgena,parafacialsard
frame-orbital plate yellow pruinose. Gena dlanging to gray pruinose
antericrtolirefrunposteriormrginofeye. ‘Ihreepostsumral
dcrsocentralsetae. Presuturalacrcstidlalsetaewithuepair
differentiatedmrethanctlers.

115

ME: Sternite 5 nuch wider than 1mg (figure 22a). Apice of
cerci slightly calvergent, with sirucus imer margins (figure 22b) .
Hillae giving aedeagus a ratter alt-like appearance ard with mall ventral
nctdl (figure 22c).

rams: Sternite 7 with, at most, a scattering of golden pruinosity
and with a curved meal projectim at apex (figure 22d).

DIAGNOSIS: Thisspeciemaybeseparatedfrunallctl'ersbytl'e3
postsuulraldorsocentralsetaeanddistinctivegenitalia. Ihaveseenno
ctlereeciewithmly3postarturaldorsooentralsfranmstanica.

TYPES: Holctype male, allotype, and 17 rule arrl 43 female paratype
all fran: Oerro de la merte, 6 km W Villa Mils, Inter-Am. H’wy, San Jose
Pwee, Cost Rica, El. 3340 11:, Doll. E.R. Heihaus, (11 various flowers,
fran 22.x.1971 to 24.11.1972. 'ne location of tie type are not given
(106?).

mm: 27 males ard 51 fenale (WID). MRIQ,
Cent) e la Matte, 23 HI 1987, Wm. L. W, Jr., 3000 m. (MD-8
male, 15 female: 1 male and 1 ferale used for figure 22a-d). 43 of
tiesespecinereverecollectediolenIrazuinCostaRie at 200ml:cn
13.11.1988 by W.L. Dowres Jr. Otter specimere fran various localitie in
Cbsta Rica.

116

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Figure 22: Genitalia of W. (a) Male fifth sternite: (b)

Hale cerci; (c) Aedeagus: (d) Female sternite 6, 7 and 8

117

DISIRIHJI'ICR: Costa Rica.

BIOIDGY: Urncredn. Adults have been collected at flowers. Host
specinens have been collected at elevatials at or above 2000 In.

W (Robimau-DesvoidY) duplex
(Figures 23a - d)

We Robireau-Devoidy, 1830: 339. Type-locality:
U.S.A., (hmlina.

W Aldrich, 1916: 255-256, figure 121 (as m
variant). Type-locality: Harem}, Hississimi. Greere, 1925: 7,
figure 2 (description of pnparium, biology). Hall, 1928: 340, fig.
11 (redecripticn, key). Aldrich, 1930: 34 (synmymy).

W antlers, rec. van der Wulp. Knipling, 1936: 418, 426,
429, 435-436, figures 7, 33, 52, 66 (description of larva). Mchr,
1943: 283, 284 (biology).

W (Aldrich). rope, 1946a: 137-139, figures 44-46
(redecriprtion). Dodge and Seago, 1954: 52, 53, 55 (biology).
Reed, 1958: 241 (biology). Dome, 1965: 954 (locality records).
lope, 1969: 22 (locality records). Robdendorf and Gregor, 1973: 7
(key, locality records).

W antlers, rec. van der Wulp. Roback, 1954: 74, figures 237-
239 (synmymy, description of male genitalia). Poorbalgll, $31.,
1968: 22 (biology) Poorbaugh and Linsdale, 1971: 52, 53, 55

118

(biology) .

mm: length: 10 - 14 m. Gama golden to grayish-yellow pruinose.
Fulrpostsumralacrostidlal setae, anterior2nx:hreducedinsize.
lateral scrtellar setae well developed, pair adjacart to subapical
scutellar setae extendirg well over half its length.

m: 'nergite 5 usually quite orange in ground color. Sternite 5
shallowly concave along anterior mrgin. Not distinctly layer than wide
(figure 23a). Apices of cerci stout with inner nargins nearly straight
(figure 23b). Hillae resembling a right triangle (figure 23c) .

m3: Sternite7go1denpruinose. Stemite6mx§1widerthan
lcng. Sternite 7 with broadly rounded, convergent lateral margins
(figure 23d).

IARVA: The first irstar is described by Knipling (1936).

HIPARILH: Greene (1925) provides a descriptim of the puparium.
DIAGNOSIS: The golden pruinose gena and large size easily separate
thisspecies frunallotherNear'otic species, excqatgga. Itcanbe

separatedfrunthisspeciesmthebasisoftbegenitalia.

TYPES: Fanale holotype, Chroline (BNH) . In good cmditim when
aramined in 1986.

MATERIAL mm: 122 males an: 186 fenales (m, 52C, GS, CSU,

119

 

 

 

Figure 23: Genitalia of W. (a) Male fifth sternite;
(b) Male cerci: (c) Aedeagus; ((1) Female sternites 6, 7 and 8.

120

m, GAD, HE, 160, (EU, SUNY, MD). U.S.A., N. (hmlina, 1918, Add.
Manse (am-1 male; used for figures 23a-c). Highlands, N.C.,
13.VIII.1957, W.R. Richards (BRC-l female: used for figure 23d).

mm: United States (Arkansas, California, Florida,
Georgia, Illinois, Louisiana, North carolina, Chic, Oregon, W,
Texas, Virginia), Mexico (Baja California Norte, Baja Chlifornia Sur,
Dlrango, Morelcs), Guatatala, Costa Rica.

BIOIOGY: Acaunonccprqiagarsspeciesmrerits range. This
quecies has been reared frcn an! mature (Aldrich, 1916: Greene, 1925;
Knipling, 1936; Mollr, 1943). It was found to be attracted to cow manure
inChliforniabyPoorbaugh, fig; (1968)h.1twasnotrearedfranthe
mnure. misspecieshasalsobemcamaflyrearedfrandogdmgin
California by Poorbaugh and Linsdale (1971). Dbhr (1943) provides
detailed informatim m the habits of this species in associatim with

cow manure and the habits of a parasitoid, WW Say
' (Hynencptera: Figitidae) . Four females (one) were collected in Riverside
Co., California at human feces.

Wmusobemnotedtobeattractedtocarim. Reed
(1958) collected this species caning to dog carcasses. Payne and King
(1972) report this species in association with decomposing baby pigs.

littleislownofthenatinghabitsofthiswecies, thDodgeand
Seago (1954) found high calcentratims of males (:1 Georgia matain

121

emits, especially after 11:00 A.M., perhaps indiatirg that emits my
play a role as a statim site for this species.

REMARKS: misspeciesislabeledasacalplexbemuseofthe
extramdiversity of themaleand female genitalia. the species is
eitherverypolytypicinthestmchneofitsgaiitaliaoritiscmposed
of nany synpatric siblirg species. After many dissectims it was
difficult to find threemale specimens whidl exactly matched one another
inthefinestncmreoftheaedeagus,evenvmenthespecimenswere
collectedaspartofaseriesfranthesanelocatimmthesanedate.
For further explanation of the application of this name to this species,
seethereiarkssectionofm.

Mostdeteminedspecinensincollectimswillbefomdmiderthe
mmmm. liter-aulrecitatimscaicenlimMOr
mareconsideredmflerthisspecies,aslfamdwtobealmost
entirelyrestrictedtonexico.

W n-SP-
(Figures 24a - d)
W n.sa. Type-locality: Argentina, Catamaran, E1 Arenal.
2600 m., w. cord Accnquija.

mm: Length: 6-9m. Gena, genalgroove, andparafacials
golden pruinose. Harte-orbital plate also golden pruinose, but with a

122

brownish tinge. Presutural acrostichal setae well differentiated. Com
anitrcdlanterbrightorangeingramicolor. Fermrarritibiagrey
pruinose.
m:msteriorarmofstemite5nldllagerthanim1ernesal
ldaes (figure 24a). Apin of cerci stout, inner margins smoothly
roamed (figure 24b). Hillae broadly married apically (figure 24c) .
FEMAIE: Sternite 6withaca1vexanteriornargin. Sternite?

triangular in appearance (figure 24d).

unravels: 'nieorangecoocaeandtrcdianters, incmjunctimwith
thedarkgreyfaulrsardtibia, servetoeasilyseparatethisspecies
frunallothers.

TYPES: Male holotype, allotype, and 4 rule and 2 farale paratypes
all frun: ms. Catamarca, E1 Arenal. 2600 111., W. cordAcamija.,
3-4.X.68, Pena (52C). (he paratype fanale fran WT. J'ujuy, Ia Quiaca,

23.x.68, 3500 m., L.E. Pena (52C).

810%: Unknown.

mm (film)
(Figures 25a - d)
W Blanchard, 1939: 835, figure 12. 'Iype-lccality:
Argentina, Ia Falda, Cordoba. ropes, 1962: 166, 167, 172-173 (key,

123

1.

4,1/ «1. n

St “a?

.uL. Be?

Me...

 

 

 

(a) Male fifth sternite; (b)

Figure 24: Genitalia of W.
Male cerci; (c) Aedeagus; (d) Fanale sternites 6, 7 and 8.

1214

descriptim of female). ropes, 1969: 22 (locality records).
W (Blarmaxdh m. 1954: 74 (m)-

mm: length: 9 - 10 m. Gee, geal groove, parafacials, and
franc—orbital plate golden pruinose. Geld abruptly changing to gray
pruinoseat lineextending frunposteriormarginof eyes. Palpus dark.
alwithdorsalsetaeetteidixgtobedofSc. Vertral setmlaeofcosta
extending past Sc to R1.

MALE: Posterior arms of sternite 5 produced laterally, midi wider
than laig (figure 25a). Apices of cerci very lung and thin (figure 25b).
Hillae with apical menbranous tip, darkly pigmented and potato-like in
appearance basally. Vesica strongly bent near apex (figure 25c).

W: Sternite 6 with convex anterior margin. Sternite 7 with
apical nasal irwaginatim. Sternite 8 with leg posterolateral processes

(figure 25d) .

DIAGNCBIS: The wing setatim, dark palpi and golden gerae separate
thisspeciesfranall others, WW. 'Ihedistinctive formof

the genitalia easily separates m1 frcm this species.

TYPES: 'mreemaleandeefelele cotypes. laFalda, Corddoa,
Argeltina, III-1933, M. Meimke. Deposited in the Blandlard's personal
collectim. olrrelt location unlmown.

125

 

 

Figure 25: Genitalia of mm. (a) Male fifth sternite: (b)
Male cerci: (c) Aedeagus: (d) Fenale sternites 6, 7 and 8.

126

mm. mm: mm, Carr-asco, Dept. Godiabanioa, Siberia,
1650 14., October 1963, F.H. Walz (151-1 male and 1 fenale: used for

figures 25a-d) .

DISIRIHJTICN: Arge'ttina, Bolivia.

BIOIDGY: mflomn.

REMARIG: Blanchard’s figure of the male genitalia of this specie
clearly differentiates it. fran all others. I believe that lope
correctly associated the female of this specie and his figure match
those provided here.

Ram ’a 611mm: (Hall)
(Figures 26a - d)
W Hall, 1928: 341, figure 13. Type-locality:
Peru, Ollantaytanbo.
We (Hall). Robadc. 1954: 74 (em-rm)- Lope.
1969: 22 (locality records). lope and Tibana, 1982a: 135 (locality
records).

mm: Gena golden pollinose. Parafacials grey pollinose, tinged
with yellow. Presutural acrcstiohal setae well differentiated.
MALE: Sternite 5 larger than wide, medial lobe nearly parallel

127

(figure 26a). Apice of cerci sligrtly cmvergert, inner margins sinlous
(figure 26b). Corpus not cmspiouously widening at attadmelt point with
hillae and bulbous apically (figure 26c)

m:Agaeararceoffl1ege1italsternitesmtdistimtenlghto
allow separatim fran several other Neotropical specie.

DIAC'NOSIS: 'Ihe structure of the aedeagus allows separation of this
specie frcn all others.

TYPES: Hale holotype, Ollantaytanbo, Peru, 9000 feet, 20 vii, 1911,

Yale Peruvian Dcpedition (USNM) . Allotype fran same locality (USNM) .

mm:mspecinexsofthisspeciewereefamined.

DISIRIHJI'IQJ: Chile, Pen. Distributim fran lope and Tibana
(1982a) .

BIOLOGY: IJnlcrm.

m:1tavemtseenrepresentativesofthisspecies, but
mll's (1928) figure of the geiitalia and discussions with Guy Sherell
attheBicsysteraticReeardiceitre iniicatethatthis isavalid

specie. 'Ihe figures preerted here are modified fran Hall (1928) .

128

 

Figure 26: Genitalia of W. (a) Male fifth
sternite: (13) mile cerci: (c) Pedeagus (redrawn fran Hall, 1928).

129

W (Aldrich)
(Figure 27a - d)

Wiggle Aldrich, 1916: 251-253, figure 119. Type-locality:
Indiana, is Fayette. Hall, 1928: 339, figure 8 (redescription).
Imipling, 1936: 421 (biology).

W Hall, 1928: 337-338, figure 6. Type-locality:
Colorado, Paonia (as Colunbia, ermr). Hall, 1929: 71 (locality
record).

W (Aldrich). Rfloack, 1954: 74, figures 247-250
(symnymy, decription of male genitalia). Dodge, 1956a: 189
(redecription, synonymy). Downe, 1965: 954 (locality records).
Turner, My 1968: 1013 (biology, parasitoid records). Haye ard

Turner, 1971: 444 (biology, parasitoid records).

mm: length: 5-7m. Gelagraypruinose. Subapicalsoutellar
setae well developed, without adjacent lateral soutellar setae. Plelra
gray pruinose in gmnd color. Four postartural dorsoceltrals.

MALE: Frontal vitta very wide, sane quecimes with proclinate
orbital setae. Geeral appearance of head resembling fanale head.
Anterior margin of sternite 5 straight, lateral margins divergert ard
nearly straight (figure 27a). Apice of cerci slightly convergent, with
redial bed along inner margins (figure 27b). Hillae clear mebranous
dorsally, corpenotectedingnldlpastvettralmrginofhillae (figure

27c) .

130

FEMALE: Sternite six with antercnesal cmcavity. Sternite 7 with

lateral margins slightly convergext, anterior margin carve: (figure 27d) .

WIS: Arathersmall speciemidlmaybecmfusedwith
Mmflebasisofectemalfeature. mleareeasilyseparated
frunthisqaeciemthebasisofthestrucmreoftheaedeagus. Fenales
canbe separated, with sane difficulty, bythe shape of the genital
sternite.

TYPES: Holotype male, Ia Fayette, Ird., Aldrich (USIM). Allotype,
Arizala (IBNM). 5 male paratype.

MA‘IERIALEDGMINED: 6 male ard 21 female (BIC, as, m, GAD,
MSU, osu, wm) imluding 4 paratype: Tifton, Ga., Oct. ’96 (mm-3
male) and Alab. (FMNH-l male). HEXICD: Hidalgo, 13 mi. SE mohuca,
thorn (pmtia scrub, VI-22-82, 8000’, Greg A. Dahlen, col. (GAD-l male:
used for figure 27a-c). ,Ane, Icwa, 22 VI 1952, W.L. names (WID-l

fenale: used for figure 27d) .

orsmrwnov: Canada (British colunbia) , United States (Alabam,
Arizcna, Colorado, Georgia, Illinois, Irdiana, Iowa, Kansas, Kentucky,
Louisiana, Michigan, Minnesota, (hi0, Utah), Mexico (Hidalgo).
Distributicn fran material examined. Wideranging specie but very
:mcannonly collected.

131

 

 

Figure 27: Genitalia of W. (a) Male fifth sternite; (b)
Male cerci; (c) Aedeagus; (d) Fanale sternites 6, 7 and 8.

132

BIOLOGY: Rhipling (1936) notes that this species averwinters as
p.1paria in Iowa (diapause). 'mrner, M (1968) reports rearing this
species fran oow nature and lists the followirg parasitoids: mm
W (San!) W: Braomidae). W (Say)
W Cynipidae). and MW (SaY)
alymenqartera: Figitidae). Hayes and Turner (1971) report rearing
mammadm’ginvnginiahxtmteflaattheyoafldmtseparate
itspzpariafranllotherooprofimagmssaroofiiagids. 'Iheyincludedthis
species under the heading m sp. in their diswssim of parasitoids,

which include the species mentioned above and a staphylinid beetle,
mm 8m-

REMARKS: Examination of thousands of specimns of N indicates
that this species is very mm. 'Iherefore, literature citations of
tlfisspeciesasaocprqhageinoowmmmareomsidemdqmtimable,
astheotherbovinedxmg—bmedirqmmigamtsuallyvexyomnmly
collected.

Mam (Aldrich)
(Figures 28a - d)
W Aldrich, 1916: 249-251, figure 118. 'Iype-locality:
NewMexioo, Alamogordo. Hall, 1923: 334, 337, figure 5
(redescriptim) . Hall, 1929b: 8‘7-88 (locality data).

133

W (Aldridi)- W: 1917: 195 (W
designation).

W121: Hall, 1928: 338, figure 7. ‘Iype-locality: New
Mexico, State College. Downes, 1965: 954 (synonymy).

W (Hall). Roback. 1954: 74 (W)-

W (Aldrich). Rnback, 1954: 74, figures 245-246
(synonymy, Hale genitalia). Dcwnes, 1965: 954 (synonymy, locality
records). Coffey, 1966: 213, 214, 217 (biology). Poorbaugh, gel:
31;, 1968: 22, 35, figures 1-2 (biology). Merritt and Anderson,
1977: 44, table 4 (biology). Davis and Mar, 1978: 120, 122,
table 1, 3, 5, 6, 7, 12 (biology). mar-ton and Moon, 1979: 81, 88,
figure 3 (description of puparium) . Wharton, 1979: 183 (parasitoid
record). Peterson, gal“ 1981: 512 (biology).

mm: length: 4 -7m. Genagreypruinose. Lightgreyto
greyish—yellw pruinose in ground color. 4 postsutural dorsocentral
setae. Preapicnl scutellar setae well develqaed with no adjacent lateral
sartellar setae.

m2: Frontal vitta very wide, head female-like in arpearance.
SterniteSsl'nrt, posteriorarnsnotextetflingmmPastnedianlobes
(figure 28a). Apices of cerci slightly convergent, inner margin siunous
in appearance (figure 28b). Corpus highly reduced ventrally to a thin
posterior lobe. Vesicn umbrarnls and very small (figure 28c) .

m2: Sternite 6 with nearly straight anterior margin. Sternite

131+

 

Figure 28: Genitalia of W. (a) Male fifth sternite; (b)
Male cerci; (c) Aedeagus; (d) Fanale sternites 6, 7 and 8.

135

7 nearly triangular in amearance overapical third (figure 28d).
PUPARILM: 'mepiparitmisdescribedbyimartmarrluom (1979).

DIAGNCBIS: 'Iheanall size ofthisspeciesarri4postartural
dorsocartralsetaeseparatesitfrunallomers,exceptp§§§jgt_a, over
itsrange. Itcaneasilybeseparatedfrungt'gflmthebasisofthe
genitalia.

TYPES: Male holotype, Alamogordo, N.M., May 5, 1902 (USNM).
Allotype, Morton Oamty, Kansas, 3200 feet altitude, F.H. Snow collector

(IBM). 8mlearrl6fenaleparatypes.

MATERIAL mm: 636 males and 482 remiss (am, BBC, as, CSU,
mm, K2, FBU, (BU, WID). (ELIE: Sierra (13., 211133 Pass, V-27-80,
Malaise trap, Stanley C. Kuba (Gs-mle: used for figures 28a-c) .
Rillito, Pima 06., Ariz., III-26-34, A.J. Basinger coll. (ms-feels;

used for figure 28d) .

DIS'IRIBJI‘IQJ: Canada (British Columbia), United States (Arizona,
Califoniia, Colorado, Idaho, Illinois, Iowa, Kansas, Mrtana, Nebraska,
Nevada, New mico, Oklahana, Oregon, South Dakota, Texas, Utah,
Washington, Wyaning), Mexico (Baja Calir'cnia Norte, Baja California Sur,
Chihuahua, mrango, Zacatecas). Distributim fran material enamined.

136

BIOIDGY: misspecieshasbeencollectedardrearedfrancowdlmg
(Coffey, 1966: Poorbaugh, $31,, 1968: Merritt and Ardersm, 1977) and
steep an; (Coffey, 1966). Coffey (1966) also fcmd that it was
attractedtohorse, swine, anddogdminsaltheasternWashingtm.
mvis ard Turner (1978) found that it was attracted to cow, deer, coyote,
arrircdentdtmhltweretmabletorearthisspeciesfrananyofthese
media. MerrittarriArdersa1(1977) rearedplamim francowdungat
all of their study sites at the Sierra Foothills Range Field Station in
Chlifornia except their irrigated pasmre site. mis and Turner (1978)
collected this species at all 9 of their surly sites in Wallowa-mitnnan
National ForestinmrtheasternOregonandthcylistthisspeciesascne
of the two most cannmly collected sarcqhagick at these sites. ‘Ihis
specie was collected using Malaise traps, baited Malaise traps, baited
Bishcpp traps (mvis and 'nn'ner, 1978). Peterson, M (1981) found
thatthisspeciesmsattractedtoSWASSpellets, abaittoxicantsystem

for screwworm cartrol.
'Ihe only parasitoid record for this species is Wt;
(Coleqatera: Staphylinidae), provided by Wharton (1979).

m: Determinedspecimmsincollecticnsmybefandmderthe
mmwm. Faralescanbeseparatedintotwo
Wypesmthebasisoftheshapeofthegenitalstenla,hrtm
dlaracterscmldbefandinthenalegenitaliatosupporttlnseseenin
thefenales.

137

W (W)
(Figures 29a - d)
W Dodge, 1968: 423, 426-428, figures 4-5. 'lype-
locality: lbxico, S.L.P., 12 mi. W Nuevo Morelos, Tanps., 1225 ft.
(male).

mm: length 6 - 9 rm. Panto-orbital plate golden pnlinose.
CostawithventralsetulaeexterdingpastSctoRl. leithdorsal setae
extending past bend of Sc.

m: Mesotibia usually with anteroventral seta. Fifth sternite
withdeepmediancleftwhidlismtcomlectedapically, andmedialpads
with lag setae (figure 29a). Oerci with convergent apices when viewed
posteriorly (figure 29b) ard strongly sinlous is lateral aspect.

Aedeagus curved with hillae mostly membrarnls (figure 29c) .

FEMAIE: Sternite: 6, 7, ard8near1yequa1inwidth. Sternite7

slightly larger than mung sternites. Sternite 8 without apin

”lip" (figure 29d) .

DIAGNOSIS: Very closely related to M1, both species having
uxevmtralseuuaecfureodstaectedingpastSctcnlarddorsalsetae
oleexteniingpasttheberdofSc. 'Ihesetwospeciesappeartobe
allqaatric,butmybeseparatedinborderregia'smthebasismthe
stnlcuneofthemaleaedeagusardfanalestenlitee.

138

 

 

Figure 29: Genitalia of W. (a) Male fifth sternite: (b)
Male cerci: (c) Aedeagus: (d) Fanale sternites 6, 7 and 8.

139

TYPES: Male holotype, Mexico: S.L.P., 12 mi W Nuevo Morelos,
'I‘anps., Sept 4, 1962, 1225 ft, Malaise trap (USMI). Fanale allotype,
Parana: Barro Colorado Island, 27.VI.1956, Rettenneyer's (RU). Paratypes:
cne fanale, same data as allotype: two rules, Turrialba, Costa Rica, Aug.
6, 1952, W.W. Neel, trap: one fetale, (brrizal, Guerno, Mex” Aug. 10,
1950, W.G. Dams: ale fanale, Mocz, Gualatcn, Guatenala, J. Bequaert; one

male, Actenango, Cuimaltenango, Guat., 1S.VI.1950, PBS.

MM‘ERIALW: 4 males and 21 females (BBC, GAS, mm, 1K2, 050,
Win), includingafenale, labeledasaparatypebyncdgebrtmtlisted
in Dodge's publication, fran Bartica, Br. Guiana, June 6, 1901, 11.3.
Parish collector (CBU). (IBM RIG: San Pedro, Mate de la Oca, 6 XII 1987,
Wm.L. Domes, Jr. (male-Win; used for figures 13a-c). EnJAD. Rio
Palenque, 0°35's, 70° 22m, 22-26 Feb. 1976, 150 m., G.E. Shewell

(female-ERG; used for figure 13d) .

DIS'IRIEJ'I'IQJ: Mexico (alerr'o, San Iuis Potcsi, Scnora, Veracruz),
antenala, Honduras, Costa Rica, Parana, Cblunbia, Ecuador, British
Oriana, Trinidad. Distrihrtim frun mterial eaamined and Dodge (1968) .

BIOLOGY: Unknown.

Miriam (van der WHIP)

1M0

(Figures 30a - d)

W191; van der Wulp, 1896: 278. Type-locality: lbxico,
Mexico city. Aldrich, 1930: 33 (syna'lymy). mipling, 1936: 425,
432, figures 3, 30, 49, 62, 79 (descripticn of larva, biology).
Hallock, 1940a: figures 32-34 (male genitalia). Hallock, 1940b:
205, 222, figures 119-121 (key, redescriptim). Mohr, 1943: 285
(biology). Sanjean, 1957: l6, 17, 39, 40, 41, 55-56, table 1-7, 9—
15, figures 114a-122 (description of larva, biology).

W authors, nec. Parker. Metz, 1916: 218, 235, figures
98-99 (description of dlranosanes). Aldrich, 1930: 33 (synonymy).

W Hall, 1928: 333, 335, figure 1 (preoc. by Robineau-
Desvoidy, 1863) . Type-locality: alebla, Mexico.

W Hall, 1928: 334, 335—336, figure 2. Type-locality:
Sierra Madre, Chihi, Mexico.

W Hall, 1929a: 71. Type-locality: Rléala, mico (new
nane for M) .

W (Hall). Iopes, 1946a: 137 (locality records). Lopes,
1948: 567 (locality records). Rdoack, 1954: 74 (synonymy).
Peterson, M, 1981: 512, 515 (biology).

W (Hall)- Roback. 1954: 74 (symnymy).

Wig]; (van der Wulp). Roback, 1954: 74, figures 251-253
(synonymy, male genitalia). Dodge an! Seago, 1954: 53 (locality
data). Reed, 1958: 241 (biology). Boyes, 1963: 1201 (dlrunosanes).
Downes, 1965: 955 (locality records). Pcorbaugh and Iinsdale, 1971:

1141

55 (biology). Payne and King, 1972: 160 (biology). Vbartm and
Mom, 1979: 81, 88 (descriptim of plparium).

AIIJIII': Length: 6-9nm. Genagraypminose. Presutural
acrostiohal setae differentiated, me pair mre than others. 'lhree
postsutmral dorsocentral setae.

MAIE: Sternite 5 with concave anterior margin, just a bit wider
than long (figure 30a). Apices of cerci parallel to slightly convergent,
with sinuous inner margins (figure 30b) . Hillae mlbals dorsally and
remaining quite wide ventrally (figure 30c) .

FEMALE: Stemite6ratherrectangularinshape. Sternite7with
lateral margins convergent almst fran base (figure 30d).

IARVA: The first instar is described by Knipling (1936) . Sanjean
(1957) provides detailed descriptions of all three larval instars.

PUPARIUM: Adescriptionoftheplparimnisprovidedbyvnlartonard

Moon (1979) .

DIAQDSIS: 'nlenorual,thinfmntalvittaofthisspeciesservesto
separatemalesfrunmgardplanflm. Malesareverysimilarin
actenalappeararnetotangjmardsmemmtareeasflysqaarated
althebasisofthemalegenitalia. Femlescanbeseparatedfrunthe

sexism. mm. mm. ardrereiglrpmthehasisofflle
genitalsternites.

1M2

 

 

Figure 30: Genitalia of Mo (a) Male fifth sternite; (b)
Male cerci: (c) Aedeagus: (d) Female sternites 6, 7 and 8.

1M3

TYPES: Dale holotype, fenale allotype, Mexico, Mexico City (RINK).

MA‘IERIAL EXAMINED: 696 neles and 708 fanales (32C, CBS, (BU, FMNH,
GAD, HE, 160, SUNY, WID). Amamlcn, m., S-V-SO, Cbll. W.G. Dams
(rm-male: med for figures 30a-c). Ames, Iowa, 22.VI.1952, W.L. m

(WID-fenale: used for figure 30d) .

DIS'IRIHJ'I‘ICN: Canada (Alberta, British Colunbia, Chtario, Quebec),
United States (Arizona, Chlifomia, Colorado, Illinois, Iowa,
Massachusetts, Michigan, Minnesota, Martana, Nebraska, New Mexico, New
York, North Dakota, Oregon, South Dakota, Utah, Wiscmsin), Mexico
(Chiapas, Chihuahua, Darango, Mexico, Mexico D.F., Morelos, Puebla,
Tlaxcala).

BIOLOGY: Despite the many literaulre references to this species,
very little is known of the prefered larval media. Knipling (1936)
rearedthisspeciesmbothdecmposirghanhngerneatardcavmamrein
the laboratory. Sanjean (1957) reared this species (I: deccnposing pork
liver in the laboratory. Molar (1943) reared this species frun cow dung
but indicatesthatthis isprobablynotthepreferedbreedingnedimn.
lthr (1943) mtesthatthiswasthemlysarcqhagidrearedfrmcattle
droppings located in dense shade. Pcorbaugh and Linsdale (1971) list
sarer'alrearihgrecordsofthisspeciesfrandogdlmg. Ihaveexamined

over 400 specimens of this species fran Iansing, Michigan which were

luu

collected by M. mynolch in 1955. Although no rearirg data appears in
caljunctim with these specimens it is believed that these rqnesent
specineIBrearedfrandogdlmgthatmymldswas investigatirgaspart
ofastuiymflyproductimsalrcesinlansirg, Michigan. I'm, many
rearirgsofdogdmgfrcmlamirgovertheslmnersof 1985-1988bythe
author have yeilded no specimens of this species. I cannonly collected
adultsoleaintheIansimareadurirgthistime.

Many collections of this species by myself and W.L. Downes, Jr.
indicate a close association of the neles of this species with carrion.
Malesmnbeeasilycollectedduringtheamnerbyseardlingabit
Wfrmasmalltonedim-sizedmanmalorbirdmss (e.g. a
road-kill). 'nlemales amearvery alert and it is believed that this
represents the station site for this species. It has been collected in
associatimwithdogcarcasseebyneed (1958) andbabypigcarcassesby
Payne and King (1972) . Sanjean collected nmbers of this species at the
following baits: horse dung, human feces, sheep liver, freshly killed
American cockroad'les, cow udder, dog viscera, dog mscle, deer heart,
woodohuck, chicken, and rabbit carcasses. It was also cmlnmly collected
sunning m a stale wall. Petersm, m1, (1981) report this species
beingattractedtoSWASSpellets, abaittoxicantsystenforscrewmrm
cartrol.

Metz (1916) myhavedescribedthedlrmnsanesofthisspeciesard
Hayes (1963) lists Metz's mm specimens urrier this name.

1&5

REMARKS: 'IheidentityofthespeciesusedbyMetz (1916) underthe
meyapplytothisspeciesorm. Specimens
determiredbyParkerhavebeenfandtorepresentboUlofthesespecies.
Aldrich (1930) mistakenlysymnymizedthis species with W.

mstdeterminedspecimensoffllisspecieswillbefomrimderthe
namenfiigla, altlnlghspecimenscancaunonlybefcurrlmderthenames
m.m.ardmellate.

Miriam (Walker)
(Figures 31a - d)

W Walker, 1849: 821-822. Type-locality: ?N. America.
Aldrich, 1930: 13, 15 (synonymy). Blickle, 1961: 802 (parasitoid
records).

W Walker, 1849: 822-823. Type-locality: Nova Scotia.
Aldridl, 1930: 13, 15 (synonymy).

W Walker, 1849: 823. Type-locality: Canada, Chtario.
Aldrich, 1930: 13, 15 (symnymy).

M Walker, 1849: 823-824. Type-locality: Nova Scotia.
Aldrich, 1930: 13, 15 (synalymy).

W Walker, 1849: 825. Type-locality: ?N. America.
Aldrich, 1930: 13, 15 (symnymy).

M Parker, 1914: 55-57, pls. 1-5. Type-lomlity:
Massadmsetts (?). Richardson, 1916: 411 (biology). wtz, 1916:
218 (chrmnsanes). Aldrich, 1930: 13, 15 (synmymy).

1U6

W (Parker). Aldrich, 1916: 253-255, figure 120
(redescriptim, biology). Wilson, 1932: 84, 89 (biology). Winn and
Beaulieu, 1932: 81 (locality records).

WM (Parker). 'Ibwnsari, 1917: 191, 194, 195 (genotype
designatim) .

Wig authors, nec. W. Hall, 1928: 334, 339,
figure 9 (redescriptim, key). Hall, 1929b: 87 (locality record).

mm authors, nec. Rebirleau-Desvoidy. Knipling, 1936:
418, 420, 421, 425, 426, 429, 434-435, figures 6, 34, 51, 65, 81
(descriptim of larva, biology). Hallock, 1940a: figures 29-31
(male genitalia). Hallcck, 1940b: 207, 210, 215, 221-222, figures
116-118 (key, locality record, male genitalia). Merrill and Hutson,
1953: 678 (biology).

M (Walker). Roback, 1954: 74, figures 240-241 (synonymy,
male genitalia). German, 1954: 44 (biology). Dodge and Seago,
1954: 52-53, 56 (biology). Downes, 1955: 524, figures 13, 22
(morphology). Dodge, 1956a: 188, 189 (synmymy). Domes, 1965: 954
(locality records). Coffey, 1966: 213, 214, 217 (biology). Sanders
ad Man, 1966: 957 (biology). Homer ard Virgo, 1967: 731, 732
(parasitoid W) . m, M, 1968: 1013, 1014 (parasitoid
data). Poorbaugh, My 1968: 22, 35, figures 1-2 (biology).
'Ihanas ard Wirrgo, 1968: 148, 151-152 (parasitoid records). Blane,
1970: 1023 (biology). Runnel ard Knap, 1970 (biology). Hayes and
thinner, 1971: 444-447 (parasitoid records). 'nms arr! )brgan,

1147

1972: 169 (biology). Merritt ard Amiersm, 1977: 43, 57 (biology).
Davis and Turner, 1978: 122, table 4-7 (biology). marton and Moon,
1979: 81, 88, figure 2 (descriptim of plparium). Pharton, 1979:
182-184 (parasitoid and predator records). Peterscn, My 1981:
512 (biology). Figg, fl, 1982: 476 (biology). Figg, m,
1983a: 961-965 (parasitoid records). Figg, £1311. 1983b: 994-995
(parasitoid records).

Awm:1ength:7-12m. Genagraypruincse. Presuwral
acrostichal setae well differentiated. Four postsutural dorsocentral
setae, the setae becanirg longer ard thicker posteriorly. Usually with 2
pairs of well developed lateral scutellar setae adjacent to the long
preapical pair. 'Iergite 5 of abdalen grey pruinose in ground color.

MALE:Posteriorarnsofsternite5notexterr1ingnmmmial
lanes and sternite not cmspicuously lcnger than wide (figure 31a).
Apices of cerci large with inner mrgins siruous in appearance (figure
31b) . Corpls not conspicuously widening at juncture with hillae (figure
’ 31c). Anteromeaal plate present between the hillae.

FEMALE: Sternite7almstbareofpruimsityanddeeporangish-red
ingroundcolor; verywide, rectangularinarpearameandmitedistimt,
even in pinned specinens (figure 31d).

HIPARILM:Adescriptia10fthepparimnisgivenbyfl1artmard
Moon (1979).

1148 g

 

 

 

 

 

 

 

Figure 31: Genitalia of W. (a) Male fifth sternite; (b)
Male cerci; (c) Aedeagus: (d) Female sternites 6, 7 and 8.

1H9

DIAQDSIS: ‘melargesize,4postamn'aldorsocentralsetae,ard
graypollimsegenaandtergiteSseparatethisspeciesfrmallbut
m. ItcanbeseparatedfrcngmigineasternNorthAnericabythe
preserveonpairsofwelldevelcpedhteralsartellarsetaeadjacentto
thepreapical saxtellar setae. InwestemNorth Americathediagnostic
stapeofthemleardfatalegenitaliawillseparateitfranmmersof

the anxia carplex.

TYPES: Male holotype, ?N. America (EN). Famd tobe in good
condition when examined in 1986.

MERIAL m: 864 neles and 972 feneles (mm, mo, (ms, (50,
mm, PMNH, GAD, MCZ, LSU, (BU, SUNY, WID). KL: Nelson Co., an-loly
Cross, 19 May 1985, G.A. oahlen, EX: Malaise trap (GAD-male; used for
figures 3la-c). KL: Holy Cross, Brion Cb., 8 June 1985, G.A. mhlan
(GAD-fanale; used for figure 31d) .

DIS'IRIHJTIGNI: Canada (Alberta, British Colmbia, Manitoba, Ontario,
glebec), United States (Arizona, Arkansas, California, Colorado,
Camectialt, Georgia, Idaho, Illinois, Indiana, Iowa, Kansas, Kentucky,
Marylani, mssadmsetts, Michigan, Minnesota, Missouri, Montana,
Nebraska, Nevada, New Jersey, New Neadco, New York, North Carolina, Ohio,
Oregon, Penrsylvania, South Dakota, 'nexas, Utah, Vermmt, Virginia,
Washington, West Virginia, Wiscmsin, Wyaning).

150

BIOLOGY: Parker (1914) nerrticns that this species is often found
feeding or larvipositin; m Imman excranent. Aldrich (1916) nenticns a
sirgle specimen reared fran cow dung. Wilson (1932) reports cannonly
rearing this species fran sheep dung. mipling (1936) reared this
speciesmhanhirgermeat, cowmarure, andhorsedung inthe laboratory.
SardersandDobsm (1966) rqaortcamxlyrearingthisspeciesfrmcow
manure througtnxt the sumer in Indiana. Coffey (1966) reared m1;
francowandhorsedung. Healsoreportsthat itwascollectedmmnnan,
sheep, swimanddogdtmginsaltheasternWashingtm. Poorbaugh, El;
(1968) reports rearing this species frun cow manure in California.
‘Ihanas and Morgan (1972) used this species as an indicator organism in
cowmamrewhen studyingparasitoids ofthehorn fly, asthe
develcpnentaltinetothewpalstagewasfardtocorrespaflwiflathat
of the horn fly. Merritt and Anderson (1977) reared this species fran
cow renure at all four of their study sites in mifornia. We is
notedforcolmizingcownemredrcppeddm'ingthenightandms
cmsidered to have a definite nocturnal activity period. Fig, fl;
(1982) reared this species fran cow fews in central Missouri. Blume
(1970) collected gm; in associatim with bovine droppings in 'Dexas.
mvis and Turner collected a few speciuens of this species in association
with cow dung in northeastern Oregon.

Richardsm (1916) collected the specimen in traps baited with
anmximcarbmatebutcollectedtmspecinensinhiscmtrol traps.

151

Knipling (1936) cammly collected this species in flytraps baited with
liver. Dodge and Seago (1954) collected nany specinens in traps baited
withdaickenentrailsorl'nnnanfecss. amnelandxnap (1970) may
collectedm intrapsbaitedwith liver, banana, ormaltmixulre in
W. Peterson, et_aL. (1981) report that this species was attracted
toSWiSSpellets, abaittoxicantsystanforscrewworms.

Merrill and Hutsm (1953) report trapping and netting this species
in Michigan onion fields. (raglan (1954) reports that this species was
observedtoprefersmmitareasinlmtanaoveradjacent lowerareas.
DodgeandSeago (1954) alsonotedthisqaeciesfrequentingamitsof
mmtains in Georgia. Collecticrs by myself and W.L. Dowms, Jr. confirm
this association of the species with the highest point in a given area.
At such sites many more males are collected than females, indicating that
this my be a station site location for this species.

'meaveragerudaeroflarvaecollectedfranSgravid fanalesby
Knipling (1936) was 29.6. Khipling notes that the developnerrtal period,
franfirstinstartoadult, ofthisspecieswhenrearedmhanhugerneat
1513 days. misspeciesismtedasoverwintering (diapausing) inthe
puparia by Khipling (1936) .

Blickle (1961) reports rearing the parasitoids W
(Say) (Kym-meta: Brawfidae). MW (Say)
(I-lynerrptera: Figitidae), and mil; sp. (Hymenoptera: Cynipidae) frun
this species in New rhnpshire. In Missouri, Kaiser and Wingo (1967)
famitlutggmlawasthepreferedhostoffihpallig, ascmparedto

152

otherDiptera speciesbreedirg incavnanlre. 'mrner, M; (1968)
mmmitoidsmnfi: Weaken”
mam sppo W: Brwmidae). We: (Say)
(Hynenoptera: whipidae), W Laterille (Hymemptera:
Pteranalidae), and 51m sun. (Coleqrtera: Stanlylinidae). This
suldywasmdertakeninpasulresinVirginiaarddiartsaregivenof
percent parasitization over time for 1965 and 1966. 'Ihanas and Wingo
(1968) found A._mllip_e_s to be the most cannon parasitoid of Mg,
with 81W (Grammt) (001W: mlinidae) and
W as lesser, although significant, parasitoids. In Virginia,
Hayes and Inner (1971) colonized the parasitoids W, E_,_
Mardhtwasmnblemaccessflnlymlmizexmmm
this species, although parasitism was noted. Merton (1979) reported A;
emcee-a. Kimmie. ammussp- asparasitoids ofgusmls
inmlifomiaanialsomtesthatthelarvaearesubjecttopredatimby
W sp. (doleoptera: Staphylinidae) adults. In central Missouri,
Fig, fl; (1983a) collected the parasitoids W, W
89- W: Diapriidae). WM (Aslmead) and
Wen (Hynerwtera:Ptermalidae)frunthep1pariaofggmlg
found in bovine durg. Fig, M (1983b) report that the highest
ixcidenceofparasitismofboviredm'gbreediJgDipterabyW
ocmrsinufisspeciesbutthattheanergernesuccessoftheparasitoid
fran the puparium was low. *

mtz (1916)nentimsthisspeciesasanexperimentalorganisminhis

153

stuiy,h1tdoesmtwntimitbynmeinhisdiswssimofthe
diranosanes of several species of Saroqriagidae.

RENAME: Frun1916tol928thisspeciesardthespeciesgmgiawere
lmownmrierthenamems. Hall (1928)symlymizedmmj§mfier
fienanemlljmis. Aldrich (1930) WWwifl)
W. Itwasnotmrtilnoge(1956a)p.lblishedmd\aracters
thatseparatedggmlafrunamiathatthisspeciesmsreported
separately. Therefore, references before 1956 concerning one of the
earlier names are considered aspotentially referingtothis species,
mlessotherwiseirdicatedbytheauthor'sfiguresorcmmlts. 'Ihe
amrentsymnymyisbasedmthemyexamixatimoftheholotypesofall
syncnymcusspecies. 'meonlydifferenceinsynonymyfranthatgivenby
Dodge(1956a)isfliep1acanentofm_miyiyaWalkerasasyrmym
ofthisspecies.

Determined specimens of this species can be fund in collections
mixedwithspecinensofanxjamderthenanejmmaswellasmder

senile.

W (W)
(Figures 32a - d)
W Townsend, 1917: 195, 196. Type-locality: Huariaca,
canycn of the Rio Huallaga, Pam. 'lbwnserd, 1935: 11
(redescriptim) . Lopes, 1962: 165-169, figures 1-11 (key,

15H

redescripticn). lopes and Tibana, 1982a: 136 (lomlity records).
W ('Ibvmsend) . Hall, 1928: 334, 344-345, figure 16
(W01. key)-

W ('Ibmsend). Roback, 1954: 74 (synonymy).
W 'nownsend. Lopes, 1962: 165-169, figures 1-
11 (key, redescription). Lopes, 1969: 22 (locality records).
WW Lopes, 1962: 169-170, figures 12-18.

Type-locality: Pasto, Departamerrto Narino, Colunbia. limes, 1969:
22 (locality records).

AflJL'I': Ia'lgth: 6 - 10 um. Gena and parafacials golden pruincse.
Palpusorange. Dorsal setaeoanextendingtoberdofSc. Ventral
setulaeofcostaextendirgpastSctoRl. Oftenwithbrightorange legs.

MAIE: Sternite Sunlwiderthanlong, posteriorarmsextending
alnnst laterally (figure 32a). Apices of cerci with nearly straight
inner margins (figure 32b) . Hillae bulbous distally (figure 32c) .

FEMAIE: Sternite Gulchwiderthanlcrg. Sternite7with lateral
margins convergent in apical half. Sternite 8 with thickened anterior

margin, appearing trilobed (figure 32d) .

DIAMIS: 'Ihe wing setaticn and red palpi easily separate this
species fran all others.

TYPE: Fenale holotype. I-Iuariaca, canyon of the Rio Huallaga,

155

 

 

 

 

 

 

Figure 32: Genitalia of W. (a) Male fifth sternite: (b)
Male cerci; (c) Aedeagus; (d) Fatele sternites 6, 7 and 8.

156

km, 10,750 feet, Decenber 20, 1913, C.H.T. 'mwnseni (IBM).

m m: 34 males and 69 females (mac, m, :50, Win).
mm, Napo, 43 km. W. Baeza, 350m, 4-7.III.76, G.E. Shalell (HIC-
male: used for figures 32a-c). Napo, mm, 0°2o's, 78° 3'w, 6.III.76,

2500 m, G. & M. Wood (HiC—famle: used for figure 32d).

DIS'IRIHJI‘ICN: Chile, Colunbia, Ecuador, Peru, Venezuela.

Distribution fran material examined and lope (1962) .

BIOIDGY: Unknown.

REMARKS: minerals dissectims of classic, red-leged m
specimens and the dark-leged form, decribed as the subspecie
1m, yeilded no casistent rnorpholcgical characters to searate
these two morphs except for color variation. Both color morphs were
examined fran specimee collected in Colmrbia. As such, I do not believe
thatthedark-legged formdeervesubspecie statusanditishere
synonymized. Lope’ (1962) subspecie W was found to be
distictardwasraisedtospecie status.

W11 n-SP-
(Figure 33a - c)

157

W n.sp. Type-locality: Iovelard Pass, Colorado.

AIIJHI‘MAIE: Length: 10 m. Gena, genal groove, parafacials, and
fra'rto-orbital plate bright silver pruinose. Outer vertical setae well
differentiated. Presutural and postman-a1 acrostichal setae not
differentiated. 'Ihree postsutural dorsocentral setae. 'Iiegula black.
Genital capsiledarkgrey, withmanylaigsetaeandwithoutstout
bristle-like setae along posterior margins. Sternite 5 with sides
approximatelyequaltodistancebebaeenlateralarns, resemblingan
equilateral triangle in outline (figure 33a). Oerci nearly as wide as
long with short stout apices which are blunt at the tips (figure 33b).
Corpus thin apically and bent anteriorly (figure 33b).

DIAGNOSIS: 'me dark tegula and genital capsule easily separate this
species frun all otters.

TYPES: Male holotype. Loveland Pass, Colo., August 12, 1958, F.C.
Hamston collector. Deposited at ISNM.

DIS'IRIHJI'ICN: United States (Colorado) .

BIOLOGY: Unknown.

W6: mismrkablespeciesisnamedinl‘umorofalymll

158

 

Figure 33: Genitalia of mm. (a) Male fifth sternite: (b) Male
cerci; (c) Aedeagus '

159

of the Biosystaratic Research 0a1tre.

mm (Walker)
(Figures 34a - d)

W Walker, 1849: 817-818. 'Iype-locality: m. America
(fanale).

W Walker, 1852: 328. ‘IYpe-locality: United States
(male ard female). Aldrich, 1930: 18-19, figure 5 (redscription) .
Dodge, 1956a: 186, 188 (key, famle genitalia). Rohderdorf and
Gregor, 1973: 8 (key).

W Walker, 1852: 328 (preoc. Macquart, 1843). Type-
1ocality: United States (female).

W Parker, 1914: 63-65, figures 26, 45. 'IYpe-locality: ?
(hale). Roback, 1954: 15, 27, 42, 73, figures 225—226 (mle
genitalia). m, 1965: 954 (locality records). Payne and King,
1972: 160 (biology). Rimmel and mm, 1970: 167, 168 (biology).
Hayes and Turner (1971): 444—447 (parasitoid records). Wharton arr!
mm), 1979: 81, 87. Fig, ML, 1982: 476. Fig), in“ 1983a:
962-965 (parasitoid records) . Figg, fl, 1983b: 994-995
(parasitoid records).

W (Parker). Aldrich, 1916: 248, 299, figure 144
(key, description of male). Hall, 1928: 334, 343, figure 15 (key,
male genitalia). Hall, 1929: 87 (locality record). Hall, 1931: 131

(key). Knipling, 1936: 425 (biology). Hallock, 1940a: figures 26-28

160

(male genitalia). Hallock, 19401:: 208, 210, 221, figures 113-115
(key, locality records, male genitalia). Sanjean, 1957: 15, 16, 17,
18, 19, 20, 33, 34, 35, 37, 39, 40, 41, 54-55, figures 106-114 (key,
biology, descriptim of larva). lbed, 1958: 241 (biology).
Blickle, 1961: 802 (parasitoid records). Sanders and m, 1966:
957 (biology).

WM (Parker). Imes, 1946a: figures 27-31, 34 (male
genitalia). Dodge and Seago, 1954: 53 (biology). Dodge, 1956a:
186-187, 188, figure 17 (key, fanale genitalia). Lopes, 1975b: 163
(descriptim).

W (Knipling, nec. l‘hll). Khipling, 1936: 425, 433-434,
figures 5, 32, 50, 64, 80 (biology, description of larva).

W (Hallock, neo. Hall). Hallock, 1940b: 208, 210, 221,
figure 112 (key, locality records, mle genitalia).

W1 (RdDaCk, nec. Hall). Rdlack, 1954: 15, 27, 42, 73,
figures 220-224 (male genitalia). Downes, 1965: 954 (locality
records).

W (Dodge, nec. Hall). Dodge, 1956a: 186, 188, figure
20 (key, fanale genitalia).

W (Walker). Irpes, 1977: 44-46, figures 7-20
(redescriptim. W)-

W (Walker). Peterem, My 1981: 512.

m: Iagth:5-10nm. mite-orbitalplategreyprujlnsewifl'i

161

yellowish tinge. 'n'e geographical race, originally described as m,
with reddish orange legs.

HAIR: Usually with well developed cuter vertical seta. Fifth
sternite generally U-shaped, with vartrally produced, semlcse medial
projectims near apex of posterior lcbes (figure 34a). Derci with short
parallel apices (figure 34b) . Aedeagus with well developed vesica and
hillae with pointed tips in lateral viav (figure 34c) .

W: Sternites 6, 7, and 8 progressively snaller in size.
Sternite 7 bilobed, with rounded anterior margins. Sternite 8 reduced,
siblated in mesal indentation of sternite 7 (figure 34d).

me: me first instar is described by Khipling (1936) under the
name 1ga_kei. Sanjean (1957) provides descriptions of all instars.

DIAGmSIS: Genitalia very similar to M, but males with
pointed tips of the hillae and females usually with rcurried anterior
margins of lobes of sternite 7. lbs two species appear to be allopatric
or slightly parapatric along a southern line extending frun San Diego,
CalifomiatoI-bJstm, Tens. 'mestnlcmreoftheneleardfamle
genitalia easily separatethisqaecies franall othernarbersofthe

W.

TYPES: Fanale holotype, ?N. America (RINK). Redescribed by Iepes
(1977).

162

 

Figure 34: Genitalia of W. (a) Male fifth sternite; (b)
Male cerci; (c) Aedeagus: (d) Fenale sternites 6, 7 and 8.

183

mu. m: 920 males and 1008 fmles (82C, (as, am, GAD,
1:2, 160, NYSM, 0w, wm). Moorhead, Clay co., Minn, VII-11-79, J.R.
Powers, 0o11r., collected at nelaise trap (male-ms: used for figures
34a-c). Dborhead, Clay Co., Mixm., VI-20-197l, J .R. Raters, (131113,

collected in malaise trap (female-(ms: used for figure 34d) .

DISIRIHJI‘IQ‘: Canada (Alberta, m'itish (blunbia, whim, New
Brurswick, Nova Scotia, Ontario, Qlebec, Saskatchewan). Cmtinental
United States except Alaska, not extending to Mexico. Distribution fran
material examined.

BIOIDGY: Averyoammcqquhagousspecies. Itisbelieved that
theusualfoodsalrceforthelarvaeofthisspeciesiscarnivoremmg,
especiallydcgdung. Irearedlargenmbersofthisspeciesfrundcg
durgcollected at several sites innidiigan, mic, Kentucky, and franone
site in Alabama. Nearly every pile of dog dung collected durirg the
smerateadloftrnsesitesmsinfestedwimlarvaeofm.
Evenverydrydcgfeesswasfotmtocmtainlarvae. MWSp.
rearedfrundcgmentbydeoof,g§_gL(1954)ispreanmdtobe,in
largepart,thisspecies. Ialsorearedthisspeciesfranwolfdimg,
collected atPotterParkZooinLansing, Michigan. Itwasrutreared,
however,francotgardtmgorblad<bear¢mgoollectedatthesame
localityarddate. 'naebearmmthatwasoollectedwasveryrmmy(a3
omparedtobearscat thatwas seen, butmtoollected, inuiohigan's

16D,

Upper Peninsula) and this may offer an explanatial of why M was
mtrearedfranit. 'nleoougardrgseanedperfectlyacceptable, yet
yieldednosarcoghagids. Outofl'urlredsofrearedspecinalsfrundcg
dung, ally a couple of irriividuals of other Sar'oofilagidae species have
beenseen. 'misspecieshasalsobeenrearedmcownarure (Sandersand
Dobson, 1966: Vhartm and Lbon, 1979: Fig, My 1983a). Knipling
(1936) reared this species on decalposing hanburger neat and Sanjean
(1957) reared it on beef or pork liver.

me life cycle, from freshly deposited, first instar to adult, was
recorded by Knipling (1936) to take a period of 16 days. Sanjean (1957)
found that the length of the life cycle was partially taperamre
dependantarrirangedfranlz-n days forlarvaerearedat27°Ctol3-18
days for larvae reared at 23° c. Sanjean (1957, Table 1-3) also provides
informticmmthelergthoftinespmtineadminmatnlrestageatzf,
25°, and 27° c. 'me life cycle averages between 15-17 days on dog dung,
with plpariation ocmring 6-8 days after deposition. The larvae migrate
awayfranthepileofdungandintothesoil beforepupariationtakes
place. 'Iheunderside ofdcgdtmghasadlaracteristicappearameafter
thelarvae leave, lookirgnxhlikesmeonehadaqalodeda firecracker
under it (persmal cbservatims). mast adults alergeduringthenight or
early mrning hours.

'lhemaximlmnmberof firstinstarethathavebeendissectedfrun
the abdmen of a single fanale is 40 (mipling, 1936) , with nest gravid
fanaleshavingnxzh feder. Idissectedneny fanalesoverthe

165

distrihlticralrangeofthisspeciesardhasmltinelyfanrioverw
larvae in some fables where nultiple specinens were collected, accept in
thestateofChlifornia. ‘Ihenexinmnmberof firstinstarsdissected
frun California specinens was twelve. Extinatims of fecmdity based on
nmbereof larvaecontainedinpin'ledspecinensofthisspeciesmaybe
difficult, due to the atpilsim of larvae frun gravid fanales as they die
inakillingvial. 'Ihisspeciesdcesnot, Maven dothisascauncnly
or cmpletely as females of Wig; (personal observatims) .
Trapping records for adults include Knipling (1936) who collected a.
m in flytraps baited with liver in the vicinity of Ames, Iowa.
Dodge and Seago (1954) collected nmerous specimens at several sites in
Georgia inflytrapsbaitedwithdlickenentrailsormmanfeess. 'Ihey
collectedalnnsteighttinesasnenyfaalesasmalesinthesetraps.
Sanjean (1957) collected this species at the following baits: horse
chm, hlmlan feces, sheep liver, hog liver, shoat intestines, deer heart,
woodchuckcarcass, chickencarcess, andrabbitcarmss. Mostspecimens
were collected at htman feces. He also notes collecting W
ispecimensonastcnewall, cntreetnmks, ardseveralotherresting
sites. Reed (1958) collected this species fran dog carcasses. Rmmel
and Knapp (1970) collected this species in Kentucky fran traps baited
with liver. Payne and King (1972) list this species as a smvenger m
babypigcamassesplacedoncpengrun'ld, intrees, andinwater.
Felelesofthisspeciescanbeeasilycollectedastheycmetodogfeces
inthesmduringthefirst 10-20m1'rutesafterdepositim (peremal

166

observations). Peterson, ML (1981) reportcnespecinencollecbed
frunlsuldyareabaitedwithSWASSpellets,abaittacicantfor
screamrns.
Malestatim-takimbehaviorismtwellmderstcodforthis
species, altrnlghIhavespentmdltinetryirgtofirdsitesardcbserve
thisbehavior. Haleshavebeenobservedadlibitingthisbehaviorinthe
aftermmthetnnflcsofisolatedtreesinfieldsarflmlowhangihg
branches of isolated trees (personal daservaticns and observations by Dr.
W.L. Downes, Jr.). Such observations, however, are not cm and a low
nrrberofmaleshavebeaminvolvedatsrdmsites. 'Iheydonotserveto
acplainwhereardvmenmatingistakingplaceinthisverycamm
species. Ectalsivesear'diinginardar'otn'd(evenmtopof)myperscnal
residencehasprovidednocluestothismystery,eventtnlghrearing
shadiesbasedmdmgdepositedbymydcginiicatescatirualfly
meantimfrunlatespringtolatefall. Bytheprocessof
elimination, it is hypothesized that mating takes place sanewhere in the
moflargertreesinthealburbanandurbanawiraments.
MatirgbetaviorofLstigglanswasobservedinalZWlZ")! 12"
collapsible field cage durirq the sunlner of 1986 in the suburban
cammity of Okenns, Ingham Co., Michigan. ‘Ihe following ample
providestheproceduresusedandtypical observaticxsofthisspeciee'
cmrtshipardcopdlatim. Mydcg, Pippin, is allele mixed-breed of
collie and German shepard origin. He sugalied all the feces used as
larvalmediaforthisstuiy. Pippinwasfedadietconsistirqcfmrina

167

Dogdnvardccmsiaaaltablescraps. anallflagswereplacedbypiles
ofbgfewsdepositedintheearlymrrflmardlateafterrnmmfi?
June. m9Jme,thefcurpilesoffeceswerecollectedandplacedm
aleirdlofslightlydanpsardinalargeplasticshoebat. 'meshcebcx
mscovered,inordertoprotectthedevelopinglarvaefrunparasitoids
orpredatorsardrain,aniwasleftartsideinashadedarea. Paparia
weresiftedoutofthesardmummearriwereplacedmalflirdl
layerofsandinirdividual6oz.paperclps. 'Ihepapercupswere
coveredwithplasticwrap,heldinplacebyarubberband. Eachalpwas
outfitted with a plastic centrifuge vial, held in place with mdeling
clay,cmtainirqmgarwaterardcappedwithhalfofacottmball. A
total of seventy-three puparia were collected.
EnergemebegananOJlmewiththreespecinens,ardall73adults
hadalergedbyzzmme. mergalcewasmtdlservedhltusuallycccurred
atnightorintheearlymmingrnlrs. Emergencenistocalrfairly
rapidly,assevaalfreshlyanergedspecinemmredaserveddurirgwing
acparsimmthesidesofthealplealcbservatimslSmirutesearlier
indicated non-energetics. Significant mortality was observed over the
first 48 bars after eclcsicn, with 18 mles and 5 females dying during
thistime. Survivingnmberscmsistedof 20mlesand30 fanales. m
zsmthemaleswereplacedinalecollapsible fieldmgeardthe
fanaleswerecadaihedintoamther. Water,drysrgar,anddcgfeess (as
aproteinsource)wereprovided.
m29Jmlethemleswereplacedintothecegewith‘females. One

168

nelewaskilledinthetransferprocess. 'metransfera'lcewasperformed
fran 12:00-12:15 P.M., E.D.'I'. The cage was placed in direct sunlight
outside. 'metalperamreranairedbeWeal80°-85°F.thraglnrtthetine
spartcbservingtheflies. mr'ingthefirst half-hair, amroximately?
wellnessfulnatihgattaptswerecbserved. Pairsof flies felltothe
bottanofthecagehltthefanalesweremtreceptivetothemales. 'Ihe
fanalesvmldrntstaymtheirfeetanipairestnmledupside-dam
\mtilthenalewasplshedoff. Saneweedswereacfledtothebcttanof
thecageatthistine,incaseam1gherfootholdwasneededthanflle
metalbottanofthecage. llinropairssuccessfully1’)oinedbet:ween1:00and
1:15 PM. me pair stayed m until 4:35 P.M., the other pair
until4:50P.M. Afavothermlcesssfulattalptsveremtedduringthe
firstcwpleoftnlrs,hxtthefrequencyofattaptsdremedoffover
time. Between6:00-9:00 P.M.thecagewasobservedforshcrtpericdsat
half-Inmintervalsandmmatirgattaptsweredaserved. 'mecagewas
mtwservedagaintmtilllflonmardtwopairsweremtedmiat
thistime.
'nlelandihgofthenalemthedorsmnofthefanaleisthefirst
bodycmtactinthisspeciescan'tshipa'dvirmallyallstrikesbythe
maleocanedasafanaleflew,ratherthanasshesatorcrawledmthe
insidecfthecage. 'mepairwmldinlnediatelyfalltothebottanofthe
cagewherethefamlemlldstnrggletodislcdgethemlefcrseveral
securisbyflimingoveranikickingthemalewithherlegs (thisoccdred
evalinsuccessful ccpulatims). Inslccessfulnetirqattapts,the

169

fanale would an rej ectim behavior ard the following matirq position was
noted: male or back of female: faale's wings mead slightly: mle's
fore tarsi at farale's wing base, cccasimally roving up to lumeral area;
nele’s midlegs ranging over fanale’s costa of wing, with tarsi suspended
abitabovethegrami:nale'shirdlegsrestingmthegrundbehirdthe
female. wring cqaulatim, the fanale would often clean her frmt legs
ard rubbed her hird legs together occasimally. 'Ihe farale would crawl
arunldthecagemlringthefirsthmr, thensettleddownincmeplace for
theranainderofthecopdlaticn. Anoddnalalentcalsistirgofthepair
reddng frcn side to side was noticed sporadically throughout the
copllaticn. 'Ihis may be carparable to the rocking notim noted for
W Meigen by Saniean (1957: 18) -

Although a few apparently successful copulatims were noticed in
nearly all rearing attarpts during the sunrner of 1986, mst mating
attaptswererejectedbythefenales. Iwasnotabletocolonizethis
speciesarrimleofthefanaleswereeverfanritocmtaindeveloped
larvae.

Potential parasitoid records (records form or outplaces of
Millie Species which imllfle W) include: W Say
(F199. steel... 1982. 1983a) and W (Say) W:
Bracmidae) (Blickle, 1961: Hayes and Turner, 1971: F199. M1. 1982,
1983a, 1983b): Emile sp. (Blickle, 1961: Fig, M, 1982) and
W (say) W: wuipidae) (mites and Tamar.
1971): m 89- (F199. as]... 1982) and WW

170

(Say) (Hynerrptera: Figitidae) (Blickle, 1961: Hayes and Turner, 1971):
m sp. (I-Iymenoprtera: Diapriidae) (Figg, M“ 1982, 1983a):
W (W) (F193: My 1983a), W SP-
(Figq. 93.31.. 1982). 1112mm 89- (F199. stall. 1983a). W
Girault and Sanders (rigg, egg“ 1982), W Asrmead
(Figs. Me, 1982, 1983a), m Iatr. (Figg, My, 1982, 1983a)
andfihnim Curtis (Fig, in“ 1982, 1983a) (Hymenoptera:
Pterunalidae): ard film sp. (Goleqrtera: Staphylinidae) (Hayes and
thinner, 1971: Fig, m, 1982, 1983a).

REMARIG: mstdeterminedspecimensofthisspecieswillbefamd
underthename ELM. Aldrich (1930) incorrectly synonymizedthe
nanefiimlamwithm. 'nlisledtomchconfusicnard
specinersdetermiredasstjnglargalmstalwaysrepresatthespecies
W- Dodge(19563)medflienmmflmmas
it was not possible to separate several species of closely related
fanales. Iopes(l977) rearaminedWalker'stypesardprovidedthe
syrulymyadoptedhere. 'nlefanalegalitalstemitesardthehillaeof
themales,aswellasotherpartsofthenelegenitalia,aremre
variablethanseeninotherspeciesofgmma. 'Ihis,cmbined
with an insufficient original descriptim of the species m1, has led
nenyauthoretousethenanelaakgiasadmpinggrumiforvarious
genitalmrflrsofflrespeciesminaflitimtom
specinem. meverywellbeacarplarofcloselyrelated

171

synpatric species, but I have beentmable to find cmsistent
mrprologicaldlaracterstoseparatethesespecies, iftheyadst. In
all cases, reared specimens fruncne lomlitywere ranarkably similar in
garitalstrwbnevfimcmparedtospecinerscollectedfrmctheneven
nearby, areas.
Mechmmnanethatmystnwupincollectimsisagma.
'Ihis none was applied to a red-legged geographial race which replaces
tremrreldark-leggedmrwinareasofthescutheastemrangeofthis
species (Florida, adjacent states). Unfortunately, the name was
fimrrectly applied to W (Wulp) by Aldrich (1916).
Althcngh Aldrich (1930) corrected this mistake and provided a
redescription of assim, many determinatims in collections were based
onhisoriginalwork. Asmdnanyspecinensthathavebeendetermined
asMstmldberedleckedtoseeifflienanewasircorrectlyapplied
mmmm- Wmmwctegulaemfi
lacksdorsalsetaeoan. Altl'nlghspecimenshavemtbeenseen,itis
believedthattheBLJgidiaoffiayesardTlmeruWnardWhartmard

‘ Hour (1979) are actually W.

Mega (van der Wulp)
(Figures. 35a - d)
W van der Wulp, 1896: 281. Type-locality: Mexico, Amla
in Guerrero. Aldrich, 1930: 34 (synonymy).
W (van der Wulp). Lopes, 1946a: 137-139, figures 41-43

172

(redescriptim, locality records). Lopes, 1948: 567 (locality
records). Iqaes, 1969: 22 (locality records).

m: Iagth: 10 - 14 m. Gan, parafacials, ard fronto-orbital
plate golden pruinose. Gama abruptly charging to grey pruinose posterior
tolineextariingfrunposteriormrginofeye. Garalgrooveless
dalsely golden pruirnse. Presutural acrostichals differentiated,
especially near anterior margin.

MALE: SterniteSmldllcngerthanwideandwithcormveanterior
margin (figure 35a). Apices of cerci begin to diverge very high, with
inner margins siruous (figure 35b) . Aedeagus with anteriorly projecting
mesal plate between hillae (figure 35c) .

FEMALE: Sternite 6 and 7 golden pollinose, very wide and rolnlded,
resarbling a broad oval when viewed together (figure 35d).

DIAMIS: 'Ihe genitalia of this species will squarate it fran all
others.

TYPES: Hale lectotype. Mexico, Annla in Guerrero, 6000 feet, H.H.
Snith (mum . 'Iwo male paratypes frun same locality.

MATERIAL EXAMINED: 14 males ard 28 fanales (HE, as, GAD, m1).
'Iepic, Enrico, 7-5-56, R.& K. Dreisbach We: used for figures 35a-
c). 'Iepoztlan, Dbrelos, Mexico, 8-20-56, R.& K. meisbach (DELI-fanale:

173

 

 

' Figure 35:'Genitaiia of EVE ia sueta. (a) Male fifth sternite; (b) Male

cerci; (c) Aedeagus; (d) Farale sternites 6, 7 and 8.

171+

used for figure 35d) .

DISIRIEH‘IQI: United States (Texas), Manico (Chiapas, Chihuahua,
Dirango, Mexico, Macico D.F., Morelos, San Luis Potosi, Tabasco).
Distributim frun material aamined ard Iqaes (1948) .

BIOLOGY: Unknown.

REMARKS: Aldrich (1930) mistakenly synonymized this species with
hem. Ibstinformatimplblishedmmderthenaneggg,mltil
1opes(1946a) separated these two species, actually applies to
1mm. 'Ihe Argentine locality data provided by alarrriard (1939)
mflertheramemmdoesmtarplytoeitterofulesespecies
asArgartinaiswellsarthoftheirrargesardthenameiscasideredan
error.

1opes(1946a)notedthattheapices ofthe cerci ofgm are
divergartardgivesthisasadistimtivedlaracterfortheseparatimof
mmM(=-W). Itslmldbenotedthattheceroido
notappeartobedivergartinpimuedspecimasorincerefullydissected
specimasbuttheceroiareratherlooselyjoinedardabitofpressme
mdissectedcercicausestheapicestodiverge. 'Iheshapeofthe
farele'sgamitalsternaarpeartobeverydistirctive,butitslnndbe
mtedthatnnchofthelateralareaofstemites6and7arehiddalin
pimredspecimerstn'dertheabdaninalterga.

175

Detemiredspecinarsofthisspeciesmaybefanrimixedwith
lhemimrimflerfllenmessetaardm.

Winn Rdoack
(Figures 36a - d)
W Raback, 1952: 48, figures 6a-c. Type-locality: Mexico,
Michoacan, W minim. W, 1954: 74, figlm 254-255

(syncnymy, male genitalia). Lopes, 1969: 22 (locality data).

mm: length: 6-9nln. Gaiaandparafacialsgreypruinose, often
withacqperytinge. Onepairofpresutural acrostichalsetaennre
differaltiated than others, erect. 'Ihree postaltural dorsocartral setae.

MAIB: Sternite 5 not mach wider posteriorly than long (figure 36a).
Apices of cerci with nearly parallel irmer nergins and cmvergent outer
margins (figme 36b). Hillaeverybroadardwithdistinctvartralnotdl
near apac, giving the aedeagus an ax-like appearance (figure 36c) .

FEMALE: Sternite7squaretoshortrectangularinappearance, with
apiml margin slightly carver: (figure 36d) . Sternite 8 nearly
sanicircular.

DIAQJOSIS: 'nlethreepcstsumraldorsocaltralsarrirelatively small
sizeservetoseparatethisspeciesfrunallothersatcartgggthlaover
itsrange. Itiseasilyseparatedfranthisspeciesmthebasisof
galitalc‘haractere.

176

- _-~--“
7 ‘~
9

[[1 i
(a) Male fifth sternite; (b)

’

~

I

1

. .. soccer . a a -
... w...~..-m.¢.....v...¢..sw\u.. e .3 ... .
.3...- O I O I ~

  

Figure 36: Genitalia of W.

(o) Aedeagus; (d) Female sternites 6, 7 and 8.

o
I

i

Malecero

177

TYPES: Holotype nele. Dunn: ‘I‘arcituro, sweeping in neadow, 780
feet, Midnacan, Mexico, Farrth Hoogstraal Mexican Biological Expedition,
1941, H. Hocgstraal (Chicago Natural History mean).

mm: 43 males and 21 fanales (sac, as, GAD, wm).
Barfoot Lookout, Chiricahua Mts., 8800’, ARIZ., 8 Sept. 1966, Wm.L.
Downes, Jr. (WLD-male: used for figures 36a-c). m., Gallatin Oo.,
Fairy Lake, 9500’, 18 All]. 1983, W.L. mares, Jr. (WLD-fanale: used for
figure 36d) .

DISTRIBUTOR United States (Arizona, Colorado, Martana) , Mexico
almanac, Mexico, Midnacan, Morellos). Distrihrtim frun neterial
aramined.

BIOLOGY: Unknown.

ms: misspeciescamuyocalr‘swithmsiglginmixedseries.

mlybyspreadingthegalitaliaordissectimcznthesetwospeciesbe
differentiated.

W (van der Mp)
(Figures 37a - d)

178

W Wulp, 1895: 270, pl.7, figure 4. Type-locality:
Mexico (male ard fanale) .

W Hall, 1931: 181-182, figure 1b. M Type-
locality: United States, Tans, Menard (male).

WW (Wulp). ropes, 1946a: 132-135, figures 23—26, 33
(descriptim, male gaiitalia, locality record). Lopes, 1948: 566
(locality records). ropes, 1978a: 221-222, figures 13-15

(redescription, lectotype designation) .

awnr: Length: 7 -9nm. Pronto-orbital plategreypruinosewith
yellowish tinge.

MALE: Head as in figures 15a-b. Usually with well develcpted outer
vertical seta. Fifth sternite generally U-shaped, with vaxtrally
produced, setulcse medial projections near apaf of posterior lobes
(figure 15c). Cerci with short parallel apices (figure 15d). Aedeagus
with well developed vesica and hillae with romded tips in lateral view
(figme 15c).

FEMALE: Sternites 6, 7, and 8 progressively smaller in size.
Sternite 7 bilcbed, with pointed anterior margins. Sternite 8 reduced,
situated in mesal irdartatim of sternite 7 (figure 15f) .

DIME: Galitalia very similar to Mans, but males with
rumded tips of the hillae and fanales usually with pointed anterior
margins of ldaes of sternite 7. lbs two species appear to be allopatric

179

 

 

Figure 37: Genitalia of W. (a) Male fifth sternite: (13)
Male cerci: (c) Aedeagus: (d) Fanale sternites 6, 7 and 8.

180

orsligntlyparapatricalalgaswthermlireatta'dirgfrunSanDiego,
alifomiatornlstmfllexas. 'nlestruzmreofthemaleandfanale

genitalia easily separatethis specie franall othermabereofthe
gamma.

TYPES: Male lectotype, Mexico City, May 88, H.H.S. (EN-I). 8 nele
paralectotype. Anula, Guerrero, 6000 ft., Aug. 8.1!. Smith (5 male-
BdNH). Mexico City, May 88, I-I.H.S. (2 male-BN1). Xualmanatlan, 7000

ft., July 8.1-1. anith (male—BM). Redecriptim of type by Lope (1978) .

mm. ammo: 73 male and 39 fanale (me, as, ($0, GAD,
m2, 160, Win). 11 mi. w. mrango, Dgo. m., 7000', June 20, 1964, J.F.
McAlpine (male—32C: used for figure 15a-e). Almolcngo, 6000', Gro.,
m., VII-29-62, Guy L. ash collector (fanale-iBU: used for figure 15f) .

DISIRIHJI'IQJ: Sartheastern U.S. (Ariza'ia, California, New Mexico,
Texas), mxico (Baja california Norte, Baja Chlifornia Sur, mrango,
' Guerrero, Jalisco, Mexico, Mexico D.F., Michoaen, Morelos, Puebla,
Sabra, Veracruz, Zacatecas) . Distribution fran material acamined and

che (1946a, 1948) .

BIOLOGY: Uhknom. 'IwO male collected at flowers of m

m (MEXICD: Dirango, l7 road mile W. DJrango, 10-IX-1966, 6600',
D.E. Breedlove, (ES) . (he nele collected at mnnan accr'anent (msa

181

Grands, Ariz., III-25-34, A.J. Basinger coll. GS). Entirely collected
atelevationsabove6000'. Maxinnnnmberof firstirstarsdissected
fran are fanale was 43.

REMARKS: Wulp’s name M was not applied to this specie
Initil Lope (1946a) redescribed this specie and illustrated its
genitalia. Prior to this time, Fall's name mi was cannonly used,
both for this specie and for abberrant forms of the specie m.
Aldrioh (1930) erroniously placed W as a synonym of M and
m. Unforttmately, Sarooghagidae specialists in the U.S. seem
to have ignored Lope’ intonation, evai after his 1978 redecription.
'Iherefore, specimens of m in North Anerien collections will
usuallybefomrimderthenamelaakii, epeciallythosedeterminedby
H.R. Dodge.

APPENDICES

182

 

mommaamocor‘mm

 

183

AWOF‘IHEMLA

m Robineau-Devoidy, 1863: 434. ‘Iype specie: W
W Meigen (orig. de.).

M Mead, 1915: 408-409. 'Iype glacie: We
W W (orig. des.).

Minis W, 1917: 191, 194, 195. Type species: Ravinia cannmis
Parker (orig. de.) = giggle (Walker).

mugs ‘Iownsad, 1917: 191, 194, 195. Type species: m
m Aldrich (orig. des.).

W ‘Iownsad, 1917: 191, 194, 195. Type specie: m
miners Townsad (orig. des.).

W W, 1917: 190, 193, 195, 196. 'IYpe specie:
W Townsend (orig- 695-)-

W W, 1917: 190, 193, 195. Type specie: Mia
mm Coquillett (orig- des-) =areliste Walker).

W W, 1927: 220, 295. Type specie: W
212mm (orig- 636-) =m miter)-

angling Erderlein, 1928: 16. Type specie: W
Bderlein, nec. van der Wulp (orig. de.) = misidentification of
m (Tomsad).

W Enderlein, 1928: 42-43. 'Iype specie: mm
Enderlein, nec. Fabricius (orig. de.) - misidaltificatim of

1814

W (?)-

m (Walker) 1849: 824-825 (M) - Nova Scotia; Canada
(Alberta, British Colunbia, New mick, Northwet 'nerritory, Nova

Scotia, Ontario, Qiebec), United State (Alaska, California,
Cblorado, mine, Massadmsetts, Michigan, Mirmeota, New Hanpshire,
New Jersey, New York, Ohio, Oregm, Permsylvania, Vermmt,
Was-1W)-

M Parker, 1914: 58-59, fig. 45 - ?Massachusetts.

m (Walker) 1852: 324 (W) - Brazil; Argentina, Bolivia,
Brazil onto Grosso, Minas Gerais, Sao Paulo, Santa Catarina),

WY- m
we: Walker. 1852: 327 (W) - Brazil-
3321311222 Townsend, 1927: 220, 295 (W) - Brazil, Sao

Paulo.
M Engel, 1931: 149, fig. 18 (Mega) - Bolivia.

M (Lope) 1946):: 227-230, figs. 1-5 W) - Brazil, Mato
Grosso: Argetim, Brazil (Ceara, Minas Gerais, Parana, Sao Paulo).

gum (Dodge) 1956a: 187. fig. 15 W) - Georgia: (made
(Ontario), United State (Arkansas, Delaware, Florida, Georgia,
Hawaii, Illinois, Indiana, Kansas, Ka'rtuzky, Imisiana, Narylani,

185

Michigan, Mississippi, Missouri, Nebraska, New Jersey, New York,
North Carolina, (1110, Peru'sylvania, South Chrolina, Tennessee,
'Dexas, Virginia, West Virginia, Wisomsin).

m (Walker) 1849: 818 (W) - ?N. America: Canada (Alberta,
British Colunbia, Ontario, Quebec, Saskatchewan) , United State
(Arizona, California, Colorado, Hawaii, Idaho, Illinois, Indiana,
Iowa, Kansas, Kentudq, Maryland, mssachusetts, Midfigan,
Mimeota, Missouri, Montana, Nebraska, Nevada, Newbie-doc, New
York, North erolina, North mkota, (hio, Oklahala, Oregon, Rhode
Island, South thkota, 'beas, Utah, Virginia, Washingtm, Wisomsin,
Wyanirg), Mexico (Baja California Norte, mrargo, Hidalgo, Jalisco,
Mexico, Mexico D.F., Puebla, Sabra, Zacatecas).

Walker, 1852: 323 W) - U.S.A. .
Wis ‘Ihansm, 1869: 535 (W) - Hawaii, Kaiolulu

W (Hall) 1928: 339-340, fig. 10 W) - Chile, Argol:
Argentina, Brazil, Chile.

m (Townsend) 1912: 357-358 M) - Peru, Piura: Chile,
Cbsta Rica, Ecuador, Panama, Peru.

W (WI!) 1915: 409-410 (M) - Fem. Croya:
Ecuador, Pam.

186

m (Dodge) 1956b: 97-99. fiQS- 1-5- W) - Chiba. Habana:
lea.

M (Prado and Foneeca) 1932: 39, fig. 7 W) - Brazil, Sao
Paulo: Argentina, Brazil, Cblunbia, Paraguay, Trinidad.

giglli - Kethcky, Marion Co., Rman’s Knob: Kentucky, New York, (hio.
m

mes (Hall) 1931: 182. fig. lc (Sewage) - Galifornia.
Coadiella.

man; (Iqae) 1962: 170, fig. 19-22 (AW) - Colunbia:
Venezuela. m

m (Lope) 1946a: 135-137, figs. 36-40 W) - mioo,
Chapfltepec: Mexico (Chiapas, wrango, Hidalgo, Mexico D.F.,
lbrelos).

m (Walker) 1852: 322 W) - U.S.: eastern North America,
MassadmssettsswthtoVeracruz, wettoManitobaand'Dexas.
W Coquillett, 1901: 17-18 W) - U.S., District of
Columbia.

187

M Aldrich. nec- Walker. 1930: 15. 17 W)-

m - Ms: Arizona. m

m (Walker) 1861: 309-310 (M) - Mexico; Florida to
Califonfia, south to Colmbia, imluding West Indie.
W van der Whlp, 1895: 272 (W) - Mexico.
W van der Wulp, 1895: 272 (W) - Mexico.
mil Hall, 1928: 345-346, fig. 17 (W) - Jamaica.

W (van der Wolp) 1896: 278 (W) - Mexico, Tabasco:
sartheastern U.S. (Arizona, Chlifornia, Iansas, Louisiana, Nevada,
New Mexico, Texas), maxim (Baja mlifomia Norte, Chiapas, wrango,
Hidalgo, Jalisco, Mexico, Mexico D.F., Lbrelos, Oaxaca, Paela, San
Luis Potosi, Sabra).
mm mm, 1928: 346—347, fig. 20 (m) - Texas.

W (Aldrich) 1916: 249, fig. 117 (W) - Florida.

M (Aldrich) 1916: 249, 299-301, fig. 145 (W) - Cuba:
Hispaniola, met-to Rioo.

W (Meiqm) 1826: 29 (W) - ?Germny. Aadxen-
m authors, nec. Harris 1780: 84 (1933)

188

m Fabricius, 1794: 315 (um) (Preooc. by Gnelin, 1790) -
Da'lnark.

Wis Fallen, 1817: 237 M) (Preooc. by Villers, 1789)
- Sweden.

W Robineau-Desvo , 1830: 346 mg) - France.
1mm Robineau-Devo , 1830: 346 (More) - France.
M Robineau-Devoidy, 1830: 346 W) - France.
m Robineau-Devoidy, 1830: 346 W) - France.
m Rnbineau-Devoidy, 1863: 427 (21mg) - Rance.

m Robineau-Devoidy, 1863: 436 W) - France.

m Robineau-Devoidy, 1863: 437 (Millie) - France.

m Siebke, 1877: 94 (gm) - Norway.

mi; Seguy, 1935: 5 W) - Chad.

m Seguy, 1938: 43 W) - lbroooo.

m1 Lope, 1975a: 485-486, figs. 1-4 - Chsta Rica.

W1 (Robineau-Devoidy) 1830: 339 W) - U.S.A., Chrolina.

m Aldrich, 1916: 255-256, fig. 121 as m variant

W) - Mississippi-

].mi [2111311, 1989: - Chile.

M (Blanchard) 1939: 835, fig. 12 W) - Argentina.

189

Wis (Hall) 1928: 341. fig- 13 (8mm) - Peru.
Ollantaytanbo.

mg (Aldrich) 1916: 251-253, fig. 119 W) - Indiana.
M Hall, 1928: 337-338, fig. 6 W) - Colorado (as
Colunbia, error).

21m (Aldrid'x) 1916: 249-251, fig. 118 (W) - NewMexioo.
M Hall, 1928: 338, fig. 7 W) - NewMexioo.

mm; (Dodge) 1968: 423. 426-428. figs- 4-5 W) - Maniac.
San Louis Potei: Mexico (Guerro, Sonora, Veracruz), Guatanala,
Honduras, Costa Rica, Panama, Colunbia, Fandor, British Guiana,

Trinidad. M

2519],; (van der Wulp) 1895: 268, 1896: 278 W) - Mexico, Mexico
City.
W Hall, 1928: 335, fig. 1 (W) (preoc. Rabineau-
Devoidy, 1830) - Meadoo, meme.
W (Hall) 1928: 335-336, fig. 2 W) - Mexico,
Sierra Madre.
mg (fhll) 1929: 71 (W) - Moo, Puebla.

190

M; (Walker) 1849: 821-822 M) - ?N. America.
m Walker, 1849: 822-823 (M) - Nova Scotia.

m Walker, 1849: 823 (W) - Canada, Ontario.

ragga Walker, 1849: 823-824 (m) - Nova Scotia.

gm Walker, 1849: 825 (W) - ?N. Marie.

mi; Parker, 1914: 55-57, pls. 1-5 m) - ?Massachusetts.

M (W) 1917: 196 (W) - Pam: Columbia, Ecuador,
Chile.
W Lope, 1962: 169, fig. 12-18 (Marnie) -
Oolunbia. u, SYN.

my. Dahlem, 1989: - Canada, Yukm.

More (Walker) 1849: 817-818 (M) - ‘31:. America: canada
(Alberta, British Colunbia, Manitoba, Nev answick, Nova Scotia,
mtario, Qiebec, Saskatchewan), omtinental United State, except
Alaska, ard not extending to Mexico.

M Walker, 1852: 328 W) - U.S.

m Walker, 1852: 328 (W) (preoc. Maoquart, 1843) -
U.S.

M Parker, 1914: 63-65, figs. 26, 45. - ?Massad'msetts.

m (van der mlp) 1895: 268, 1896: 281 (W) - Mexico,

191

m Roback, 1952: 48, fig. 6a-c. - Mexico, Michoacan.

W (van der Wulp) 1895: 270, pl. 7, fig. 4 W) - Mexico;
soutlmetem U.S. (Arizona, California, Ne Mexico, Texas), Mexico
(Baja California Norte, Baja California Sur, wrango, Guerrero,
Jalisco, Mexico, Mexico D.F., Miohoacan, Morelos, Riebla, Sonora,
Veracruz, Zacateoas).

laakei Hall, 1931: 181-182, fig. 1b (W) - Texas. N. SYN.

192

APPENDIXB: FIGJRESOF'DIEHEAIBOFMESOFSEVERALMSPECIES

193

 

Figure 38: Head of W male. Front and lateral view.

19L}

 

Figure 39: Head of W male. Front and lateral view.

 

 

195

 

Figure 40: Head of mm; male. Front arri lateral vie.

196

 

Figure 41: Head of minia mfi male. Front and lateral view.

197

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M "u Hm

      

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Figure'42: Head of W male.

 

Front and lateral view.

 

198

 

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l

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Figure 43: Head of mm; male. Emit and lateral view.

199

 

 

Front and lateral view.

frenata male.

a

44: Head of m

Fi

     
 
 
   

 

.' /
\ . an: “Vii:

\3

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«i 3
31

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Figure 45: Head of mm mle. Front and lateral view.

201

Figure 46: Head of Balm 'a globulus male.

 

Front view.

,“n‘fiflm

 

202

 

Figure 47: Head of W male. Front view.

203

  

     
 

v' 'a stinulans male. Front and lateral View.

Figure 48: Head of

201+

 

Figure 49: Head of Balinia vagaAJLda male. Front view.

 

205

mm

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206

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