Studia Entomologica,
vol. l2,fasc. l4,novembro 1969

Notes on the Tropical American Speciesof Tipulidae (Diptera)'VIII.
Subfumily Tipulinae

Charles P. Alexander':

Edit6ra Vozes Ltda., Petr6polis,Rio de Janeiro, Brasil

I 3 7

Studia Ent., vol. 12, fasc. l--1, novembro1969

I6l

Notes on the Tropical American Speciesof Tipulidae (Diptera)
VI|I. Subfamily Tipulinae
CharlesP. Alexander
Emeritus, University of Massachusetts,Department of Entomology,
Amherst, Massachusetts(With 70 figures)

In the precedingnumber of the present series of reports(Studia Entomologica,10: 277-352, 78 f igs.; 1967) a list of
the hitherto publishedparts was provided, all having appearedin the Revistctde Entomologiabetween 1945 and 1951. In thisnumber it was indicated that a concluding report would be
prepared to include the subfamily Tipulinae and so complete
the record of speciesin the family for the Neotropicalregion.
At approximatelythe date of the present report a contribution
by CharlesP. and Mabel M. Alexandershould be publishedas
Fascicle4 of <<ACatalogueof the Diptera of the Americassotrth
sponsoredby the Departamentode Zoolo'of the United States>>,g i a , s e c r e t a r i ac l a A g r i c u l t u r a ,S i o P a u l t t ,B r a s i l . T h i s v o l u n t eproviclesbibliographicreferences to the lllore tltan 3,000 Neotropical speciesof Tipulidaepresentlyknown and it is believccl
withincluded,usedin conjunctionthat the bibliographicreferences
the presentseriesof Notes,will prove of serviceto future workers
on this extensiveand involvedf amily of Diptera.
The present paper includes a discussionof the subfamily
Tipulinae, an extensivearray of species,including the larger
membersof the f amily. Recent studies on the more primitive
Tipuline genera,especiallythose made in Australia by Dr. N'
Dobrotworsky,in England by Mr. Richard I. Vane-wright, andsome significantchanges
by the presentwriter, have necessitatedin ttre hitherto acceptednomenclatureand distributionof [{enerain the subfamily.
At about 1930 it was realized that in the great genus
Limonis Meigen,in the subtamilyLimoniinae,a markedreductionin the numberof supposeddistinct related generawas necessary
and many of these were relegated to subgenericrank under
Limonis or were placed in the synonymy,as was discussedin
PartsVI and VII of the presentseriesof Notesand in a companion
ll

r62

A I e x a n d e r , T r o p i c a l A m e r i c a nT i p u l i d a e ( V I I I )reportby the writer ( PhilippineJour.Sci.,g3: 427-433; I 96a).
Morerecently
it hasbecome
apparent
thata comparable
consolidation of nanles was advisablein the subfamily Tipulinae and a
prelirninary discussion,as it cclncernedthe genus Ctenophora
Meigen,was provided by the writer ( PhitippineJour. Sci., 83 :264-265; l95 ). As indicated above, recent studies by Dobrotworsky and Vane-Wright are of primary importance in thisconsideration,as discussedlater in this paper.
In this reporta considerable
proportionof the includedgenera
pertain to the so-calledprimitive Tipulinae,the remainderbeing
what apparentlyare more specialized,
such as CtenophoraMeigen,
Dolichopeza Curtis, Nephrotomu Meigen, and Tipula Linnaeus.
The generalizedtype of wing venation is shown irr Megistocero
Wiedemann ( Fig. 2) where vein Sc, is strongly preservedandthe free tip of Sc, lies far proximad, at the wing nrargin being
confluentor virtually so with the thickenedapex of Scr to close
the narrowedcell Scr. The continuingvein Rl longitudinallybisects the stigma as a pale element,joining the very short Rz
at the distal end of the stignra. Brochypremna Osten Sacken( Fig. I ) is similar but has the f ree tip of Sc, more distal, acondition that is even more accentuatedin the remaininggeneraof the subfamily, as shown in the various f igures.
The generic, subgenericand species names arc arrangedalphabeticallyfor ease of reference.
Brochypremna.
Osten Sacken
Ctenophora Meigen
Dolichopeza Curtis
SubgenusMegistoma.sfixAlexancler
Elnoretta Alexander
Holorusia Loew
IschnotornaSkuse
Icriomastax Enderlein
Ischnotoma Skuse
Neotipula Alexander
Leptotarsus Cuerin-Mdneville
Aldrovandia Enderlein
Araucomyia Alexander
CeoneuraAlexander
Leptotarsus Gu€rin-Menevi
lle
Limoniodes AlexanderLongurio Loew
Pehlkea Enderlein
Tanypremna Osten Sacken'fanypr emnella Alexander'fanypremnodes Alexander
Xenotipula AlexanderStudia Ent., vol. 12, fasc. l-4, novembro1969

r63

MegistoceraWiedemann
NephrotornaMeigen
OzodiceraMacquart
DihexaclorutsEnderlein
OzodiceraMacquart
PhucclodoceroEnderlein'l'ipula
Linnaeus
Bellardina Edwards
EumicrotipulaAlexancler
Lunatipula Edwarcls
Microtipula Alexander
PectinotipulaAlexancler
Pt erelochisusRonclani'l'richotipulo Alexancler'l'ri plicitipula Alexander
YamatotipulaMatsumura
ValdivianaAlexander
Z elandot
i pula Alexancler

BrachypremnaOstcn Sacken
B r - a c - h y , p r e m nOas t e n S a c k e n; B e r l i n . E n t . Z e i t s c h r
l6l; 1886; (tvpe dispetlens^ w a r k e r , d e s i g n a t e t l b y c o q u i n e t t , ryg. r 0 . a s a i " p r i h n l f'. t
IJrachl'premna Alexander; Jour. N.
Ent. Soi:., zo zzj-zza, t f ig.. pl. (with 6
figs.); 1912.

BrochyTtre
mna is an essentiallyNeotropicalgenus with asinslewiclespreacl
species(dispellen.s)that extendsinto the eastern
UniteclStates.Two further speciesare Australasian,thesebeing
R. tigrivenlrisAlexancler,of north QueenslandandB. waigeuensii
Alexander,of the Papuan subregion.Tlrree f ossil speciesf rom
the lower Tertiary have been clescribedthat should be discussed
as to their possible affinities with Brachypremnoand with
MegistocerrWiedemann,as consideredlater. MegistocerogurnelensisCockerell(Ann. Mag. Nat. Hist.,ser. g, 7: 4s6,fig., wing;l92l) appearsto be a nrenrberof the presentgenus but may
pertain to Megistocera.It was clescribedfronr the Curnet Bay
beds in the Isle of Wight, consideredas being Oligocene.The
speciesclescribedas Macromastixbornhardti Meunier,from Baltic
Amber ( Upper Eocene) very probably belongs to the genus
Megistoce
re, as nrentionedunder that genus.The greaily lengthened male antennaeand general structure of the hypopygiunr
support this reference,since the conditionsare not unlike thosefound in the recent Old World Megistoceraf ilipes ( Fabricius).
All known speciesof the presentgenus have the antennaevery
short in both sexes.The type was re-examinedby the writer andwas further discussedand figured under the name Macromastix
bornhardti(Bernstein-Forschunsen,
AnrberStudies,2: lg-21, fig.
l l t

r64

A I e x a n d e r, Tropical AmericanTipulidae(VIII)

4 [wing], fig. 5 [male hypopygium]; l93l). The third fossil
species,describedby Meunier as Brochypremno eocenica,I 906,f rom Baltic Amber,evidentlydoes not belong to this genusand
was furtherdescribedand figuredby the writer as Tipula eocenica( 1 .c . , l 9 3 l : 3 4 ) .
The generalmorphologyof the adult flies of Brachypremna
has been discussedin the Alexander l9l2 paper cited above.
The generalfeaturesare mentionedbriefly at this time. Nasus
Antennae( Figs. 3, 4) shorter than the maxillary
conspicuous.
Eyes
in caseswith l0 segments.
palpi, contmonly1I -segmented,
with ommatidiasmall and very numerous,as in Megistoceroandvirtuallyall other generain the primitiveTipulinae.Wings (Fig.
I , Brachypremnoilludens) with vein Sc2*R1 longer than thefree tip of Scr, the latter a short distancebeyondSct and not
closingthe cell; vein Rn-s elongatein all speciesexceptingB unicolorwhere it is in longitudinalalignmentwith Rs and with
r-m distinct;ceil 2nd A very narrow.VeinsMr, Ms and M 1 at
or closeto the wing marginnarrowlywhitened.Male hypopygium( Figs. 6, 7 ) with the clavateouter dististyleinsertedon lower
margin of the larger inner style,with a small tubercleor spine
on outer margin of style at this point. The whiteningat the tipsof the outer medialveins,as described,should be emphasized.
It occursin all Neotropicalspeciesand alsoin the two Australasianf orms abovenrentioned.It similarly occursin speciesof Megis'
the relationshipbetweenthesetwo genera.
toceraand emphasizes
The immaturestagesare poorly known. The pupa has beenfound in rotting moist logs, a very difterent habitat from thatof Megistocero(Alexander,Crane- fties of New York, Part II,
pp. 984-985,f igs. 47 4-477 [detailsof PuPa| ; I 920)
List of Species
Ecuador.
abitaguaeAlexander.
(albimana Wiedemann,see candida')
Brazil.
angusta Alexander.
Ecuador, Peru.
appendigeraAlexander.
Ecuador.
arajuno Alexander.
arcuaria arcuaria Alexander.- Ecuador.
Venezuela.
a. triangularis Alexander.
Argentina.
australis Alexander.
Peru.
basilica Alexander.
Brazil.
hrevigerutaAlexander.
Brazil.
breviterebraAlexander.
Bolivia, Brazil, British Guiana,
breviventris (Wiedemann,as Tipula)
Peru, Surinam,Venezuela.

165

Studia Ent., vol. 12, fasc. 14, novembro 1969

I
6

7

Fig. I. Brachypremna illudens Alexander; venation. - Fig. 2. Megistocera longipennis( M a c q u a r t ) ; v e n a t i o n . - F i g . 3 . B r a c h y p r e m n aa r c u a r i a A l e x a n d e r ; m a l e a n t e n n a .
Fig. 4. Brachypremna dispellens (Walker); male antenna, two. - Fig. 5. Megistoceral o n g i p e n n r s( M a c q u a r t ) ; m a l e a n t e n n a .
Fig. 6. Brachypremna candidella sp. n.;m a l e h y p o p y g i u m .F i g . 7 . B r a c h y p r e m n ac a n d i d a A l e x a n d e r ; m a l e h y p o p y g i u m .
S y m b o l s : b , b a s i s t y l e ; d , d i s t i s t y l e ; p , p h a l l o s o m e ;t , t e r g i t e .

candida Alexander.- re-namingof albimana (Wiedemann),nec albimana(Fabricius).
Brazil, Ecuador, Panama, Peru, Surinam.
Peru.
candidella sp. n.
clymene Alexander.
Brazil.
dispellens (Walker, as Tipula). - Brazil, Costa Rica, Guatemala,Mexico,
Panama, Surinam, Venezuela;Trinidad; Eastern United States.
Peru.
diversipes Alexander.
geijskesi Alexander.
Surinam.
Alexander.
Peru.
illudens
Brazil.
integristigma Alexander.
itatiayana Alexander. - Brazil.
karma Alexander.
Ecuador.
laetiventris Alexander.
Venezuela.
Brazil.
nigrolemorata Alexander.

166

A I e x a n d e r, TropicalAmericanTipulidae(VIII)

phrixus Alexander. Ecuador.
pictipes Osten Sacken. Brazil.
pictiventris Alexander.
Brazil.
quasimodoAlexander. Ecuador.
sappho Alexander. Brazil.
Costa Rica, Mexico,Nicaragua,Panama,Venezuela.
sini/is Williston
Alexancler. Bolivia.
subevanescens
subsimilisAlexander. Argentina,Brazil
subuniformisAlexander. Venezuela.
Peru.
thyestesAlexander.
Cuba, DominicanRepublic,Haiti, Grenada,
unicolor Osten Sacken.
Jamaica,Puerto Rico.
uniformisAlexander. Brazil, Peru.
variitibia Alexander. Brazil.
williamsoniAlexander. Brazil, British Guiana,Peru, Surinam.

Brachypremnacandidellasp. n.
Closelyallied to candida,differing especiallyin the male hypopygium,including the conformationof the inner dististyle and the very slenderaedeagus.

L e n g t h ,a b o u t l l - 1 2 n l n l . ; w i n g , l 3 - 1 5 m m .
Male.
Frontal prolongationof head light yellow above,includingthe
long nasus,lower half dark brown; proximalsegmentsof palpi
brown, the incisurespale, third segnlent more yellowed, theterminalone dark brown. Antennaewith scapeand pedicellightyellow; f lagellum brownish black. Head light grayish brown;
anterior vertex f ronr about equal to nearly twice the diameterof the scape.
Pronotum obscure yellow. Mesonotal praescutumobscureyellow with four light cinnamonbrown stripesthat are narrowly
borderedby slightly darker brown; posteriorscleritesof notunr
light brown. Pleura yellowed,without pattern. Halteres withstem whitened,the small knobs brown. Legs with coxae andtrochanters
yellow; femorayellowishbrown, tips narrowlywhitened; tibiae darkerbrown, basewhitened,slightly narrowerthan
the f emoral apex, tips paling to yellowishwhite, narrowly so
on f ore and middle legs, extensivelyon posterior pair where
nearly half the segmentis included;tarsi whitened.Wings pale
brown,stigmadark brown,solidydarkened;morewhitenedareas
before and beyond stigma and more vaguely in centersof allouter medialcells and in I st Mzi no whiteneddropletsin outer
endsof cellsRs and Mr, as in sonlealliedspecies;veinsbrown,
M and lsf A paler and more delicate.Venation: Basal sectionof R+-s long, in punctiformcontactwith Mr-2.

Studia Ent., vol. 12, fasc. 1-4, novembro1969

t67

Abdominaltergitesbrown, basesof segmentsnarrowlyyellow€d, the color becomingmore restrictedon the outer segments,
sternitesmore uniformly yellow; eighth segme
nt brownish black
to f ornr a narrow ring; ninth segment light brown, yelloweci

outwardly,includingthe dististylesand outer halt of basistyle.
Male hypopygium(Fig. 6) with apexof innerdististylerelatively
narrow,decurved,tip obtuse.Aedeagusunusuallyslender.
Habitat: Ecuador,Peru.
Holotype,6 , Jatun Yacu, Napo-Pastaza,
Ecuadrlr,altitude
700 meters,March 1937(Wm. Clarke-Macintyre).
Paratopotypes,
3 nrales.Paratype,6 , Cerro Azul, Loreto,Peru, April 27, 1947
(Jos6Schunke).
I have shown the male hypopygium of Brachypremno candida
Alexanclerfor comparisonwith the presentfly (Fig. 7).

CtenophoraMeigen
C t e . n o p h o r.aM e i g c -;n I l l i g e r ' s M a g . . 2 : 2 6 J ; | 8 0 3 ; ( t y p e p e c t i n i c o r n i s L i n n a e u r s :_ - d e s i g . n a t e d _ Fwre s t w o o d , ! r y O ; p s e u d o t y p ei t a v e o t a i a F i b r i c i ' u i , u v n o n o a n i , i s t 6 . 'F l a b e l l i f e r aM e i g e n ; N o u v . C l a s s . ' . M o u c h .p, '. ' _ 1 3 ; 1 8 0 0 ; i n v a l i d ' n i i n e , s u p p r e s s e db y
l n t e r n a t i o n a l C o - m m i s s i o nZ o o l o g i c a l N o m i n c l a t u r e . O i i n i o n O Z g ; 1 9 6 3 .
S u b g e n . uPs s e l l i o p h o r aO s t e n S a c k t n ; B e r l i n . E n t . Z e i t s ' c h r . ,3 O : i O S - t O O :t g g 6 ; ( s u b g e n o t y p e^ -l g - e . t aF a b r i c i u s ; d e s i g n a t e db y E n d e r l e i n , F e b r u a r y 1 9 1 2 ; B r u n e t t i , ' N o vemberl9t2).

The occurrenceof three speciesof the subgenus Pselliophorain Tropical America providesa problem in distributionsince thegreat majority of speciesare f ound in the Oriental and eastern
Palaearcticregions.In the Nearctic region three other subgeneraof Ctenophora occur, including typical Ctenophora, Tanyptera
Latreille,and PhorocteniaCoquillett(Malpighia Enderlein).Thesevarious groups had been maintainedas valid genera until 1954
when they were placedas subgenera(for discussion,see Alexander, Philippine Jour. Sci., 83 : 264-265).
Antennaeof ntale (Fig. 22) l2-segmented,
with nine branchedflagellar segments;pedicelvery small, disciform; first f lagellar
segmentstout,simple,roughly suboval;secondto ninth segments
each with f our long branches,the basal pair longer than theouters; tenth f lagellar segmentwith a single pair of branches
that are basal in position,subequalin length to the axis of theseement; all branchesand the axis of terminal one with very
abundanterect black setae that slightly exceedthe diameter ofsegnrent; apex of terminal segment narrowly yellowed andglabrous; longest brancheswith sparse scatteredblack verticils

I _riA I e x a n d e r , T r o p i c a l A m e r i c a n T i p u l i c l a e( V I I I )near outer ends,outer brancheswithout such verticils;branchof first segmentwith a seriesof similarverticilsfor virtually
the entirelengthof lowersurface,totallingabouttwelve.
The three Nearcticsubgeneraabovelisted have the flagellar
branchesmuch shorter,the longestbeing subequalto the segtnent,
all branchesvirtually glabrousexceptfor very sparsescatteredverticils.In Tanypteraeach such segmenthas three branches,theunpairedouter one short; Ctenophoraand Phorocteniaeach havef our such brancheson each segment,these being subequalin
length in the latter subgenus,the outer pair slightly shorter inCtenophora.
The Old World speciesof Pselliophorahave the antennae
andwith the elongateterminalelementunbranched,l3-segnrented,
with the basal f lagellarsegmentmore or less producedinto alobe or point. All specieshave the f lagellar branchesprovided
rvith conspicuoussetae as in the Neotropicalmembersof thesubgenus.
The Neotropicalspeciesall are rare and little known. It
is very probable that the immature stages occur in decaying
In Asia, Ctenophoro
rvoodas is the casein the Nearcticsubgenera.
(Pselliophora)chrysophiloWalker was observedwhile laying
eggs in tree hollows, ?s recordedby Brunetti, 1912, while C.
(P.) dendrobio( Edwards), in Penang,was rearedfrom larvae
found damagingthe roots of Dendrobiumformosum (Orchidaceae).
List of Species
Mexico.
margarita Alexander.
Cuatemala.
mesamericanaAlexander.
Venezuela.
Alexander.
venezuelen.si.s

DotichopezaCurtis
D o l i c h o p e z aC u r t i s ; B r i t i s h E n t . , 2 , p l a t e 6 2 ; t 8 2 5 ; ( t y p e a l h i p e s S t r d m , a s s y l v i c o l a
Curtis; monotypic).
Lcptina Meigen; Syst. Beschr., 6. plate 65, fig. l0; 1830; (type albipes Strdm, as
sl,lvicola Curtis; monotypic).
A p e i l e s i sM a c q u a r t ; D i p t . e x o t . , s u p p l . | : 8 ; 1 8 4 6 ; t y p e c i n e r e a M a c q u a r t .

R e f e r e n c e sA:l e x a n c l e r , C . P . , P h i l i p p i n eJ o u r . S c i . , 9 4 : 5 6 4 - 5 8 2 ,f i g s . 4 0 - 6 1 ; 1 9 6 5 ( s u m m a r yo f s u b g e n e r a ) .
B y e r s, C. W. The crane fly genus Dolichopezain North America.
U n i v . K a n s a sS c i . B u l l . , 4 2 : 6 6 5 - 9 2 4 , 2 4 4 f i g s . , 1 3 m a p s ; 1 9 6 l ( r e v i e w
o f N e a r c t i c s p e c i e s ,i n c l u d i n g b i o l o g y ) .

Studia Ent., vol. 12, fasc. l-4, novembro1969

r69

The genusDolichopezopresentlyincludes1I subgenera,asconsideredin the Alexanderreference above cited. There arescoresof specles,being f ound in all biotic regions,including
Madagascarand New Zealand.The only one of these groups
found in TropicalAmericais MegistomastixAlexander,discussed
hereinafter.In the Nearctic region two other subgeneraoccur,
DolichopezoCurtis and OropezaNeedham,treated in detail byByers,l96l .
The immaturestages and their habitats of Megistomastix
rcmain unknownbut may be assumedto be generallysimilar to
thoseof Dolichopezaand Oropeza.The early stagesof the latter
two occur in cushionsor patchesof bryophytes,includingbothmossesand liverworts,preferably in moist areas but in certain
casesunder much dryer conditions,as discussedby Byers.
Subg. MegistomastixAlexander
I'IegistomastixAlexander; Psyche, l9: 63-64, 3A l e x a n d e r , m o n o t y p i c a n d o r i g i n a l d e s i g n a t i o )n.

figs.;

l9l2;(type portoricensis

presentlyknown are Neotropical
All speciesof Megistomastix
and are restrictedto the Creater Antilles, including Cuba,
Hispaniolaand Puerto Rico, but not occurringin Jamaicaas
presentlyknown.It is probablethat severalfurtherspeciesremain
to be discoverecl.
It may be noted that all eight speciesknown
f rom PuertoRico are f ound in the LuquilloNationalForestandthat six of thesehave beentaken at light at a singletime. The
made in 1945by Dr. Harry D. Pratt and Mr. Jenaro
collections
were f ronr an altitude of about 2,600 f eet
Maldonado-Capriles
in an environmentof Sierra Palm, Euterpe globosa Gaertner.
all taken at light, consisted
The extensiveseriesof specimens,
only of males, a condition quite different from that found in
most other craneflies that are attractedto light, where the great
are females.The ecologicalconditionsin
majority of specimens
the Luquillo Mountainshave been discussedby H. A. Gleason
and M. T. Cook (Sci6ntificSurveyof Porto Rico and the Virgin
Isfands,vol. 7, pt. 2: 126-129,plates27-35; 1927,on the Plant
Ecology of Porto Rico).
Charactersthat separatethe subgenusf rom others include
especiallythe very long male antennae(Figs. I l, l2), in somevestiture;the presenceof abundantspecieswith long conspicuous
macrotrichiain the outer wing cells ( Figs. 8- 10), and in hypopygial structure,especiallythe lobed dististylesin the majority
iedof the known species,being more generalizedand unmodif

t70

A I e x a n d e r , T r o p i c a l A m e r i c a nT i p u t i d a e ( V I I I )in domingensis
and someothers.In nrostspeciesthe maleantennac
are longer than the wing, being shortestin darlingtoniand domingensis.
The vestitureof the antennalf lagellumis noteworthy
and provides strong charactersfor the separationof species.
This includesunilaterallyarrangedverticils,with about two or
three on each segntent,and a more abundantnormal vestiture,
consistingof erect white or pale very delicatesetae.As utilized
in the accompanying
k.y, certainspecieshave such normal setaevery long and abundant,much exceedingthe verticils,?S in
portoricensrs
and multifila, f or example.Other specieshave these
normalsetaemuchshorterthan the blackened
verticils,ds foundin prattianaand others.
The wings (Figs.8-10) commonlyhavevein R,.-, atrophied,beingpresentand completein darlingtoniand dontingensis,
which
specieslikewisehavethe mostabundantand generallydistributed
macrotrichiain the cells, includingall cells with the exceptionof Sc and extendingbasadalmostto the arculus.Most Puerto
Rican specieshave the trichia restrictedto the cells beyondthe
cord or virtually So, being more numerousin polytricho andsome others. The hypopygialstructure has been discussedin
detail under the variousspecies( Figs. l3-20).
List of Species
acutiloba Alexander.
Puerto Rico.
borinqueniasp. n.
Puerto Rico.
cubensisAlexander.
Cuba.
darlingtoni Alexancler.
Cuba.
devexaAlexander. Cuba.
domingensisAlexancler. Hispaniola:DominicanRepublic.
jenaro sp. n.
Puerto Rico.
multifila sp. n.
Puerto Rico.
obtusilobaAlexander.
Puerto Rico.
polytricha sp. n.
Puerto Rico.
portoricen.sis(Alexander).
Puerto Rico.
prattiana sp. n.
Puerto Rico.
vittiner,is Alexander.
Cuba.
Key to the Puerto Rican species

t . Antennae of male with normal setae very long and delicate (Fig.

'
I I ), subequal to (polytricha) or exceeding the sparse unilaterally
arranged more darkened verticils, the longest setae more than onethird the length of segment
2
Antennae of male with normal setae shorter (Fig. l2); less than
one-half the verticils and much shorter than the segments
5
2. Generalcolorationof thorax, yellow, including the pleura; male hypopygium (Fig. l4) with dististylesvirtually unbranched;lateral tergal
arms narrow, glabrous
borinqueniasp. n.

Studia Ent., vol. 12, fasc. l-4, novembro1969

t7l-

General coloration of thorax dark brown, including the pleura; male3
hypopygiumwith both dististylesbearing lobes or projections
3. Trichia of wing cells very abundant,including outer end of cell R;
male hypopygiumwith apicesof lateral tergal arms narrow, glabrous
polytricha sp. n.
(Fig. 18)
- Trichia of wing cells less extensive,lacking in outer end of cellR; male hypopygiumwith lateral tergal arms obtuse,with setae to
4
the tips
4. Setae of flagellar segments(Fig. I I ) very long, approximatelyonehalf the segmentor more; male hypopygium (Fig. 16) with central
lobe of ninth tergite broad, its apex rounded or subtruncate;margin
multifila sp. n.
of outer lobe of outer dististyle untoothed- Setae of flagellar segmentsshorter, less than one-half the longest
segments; male hypopygium (Fig. l9) with central lobe of ninth
tergite relativelysmall, subtriangularin outline; inner margin of outer
lobe of outer clististylewith two separatedteeth or roughenings
portoricensis(Alexander)
5. Apices of lateral lobes of tergite of hypopygiumacute or subacttte,......6t h e i r m a r g i n sg l a b r o u s- Apices of lateral lobes of tergite obtuse,with setae throughout
7
hypopetiole
subequalto m;
6. Wings with cell Mr relativelyshort, its
in a single
lobes
(Fig.
terminating
pygiuml3) with lateral tergal
slenderspine, central produced area of tergal border small, with few
acutilobaAlexander
setae-- Wings (Fig. 9) with cell M, deep,sessileor virtually so; hypopygium(Fig. l5) with lateral tergal lobes stouter, unequallybilobed, central
proclucedarea of tergal border extensive,with abundant.setae
Ienaro sp. n.
7. Wings with anterior cord scarcelydarkened;male hypopygium (Fig.
17) with lateraltergal lobeslong, outer end more dilated,with numeroustlarkenedspinoid setae, central producedarea of tergal border subtriangular; inner lobe of outer dististyleenlarged,with abundantlong
obtttsilobaAlexander
setae,beak of inner style very stout
spincricl- Wings (Fig. l0) with a narro\r' brown seam over the anterior cord;male hypopygium (Fig. 20) 'with lateral tergal lobes relatively short,
not expandedoutwartlly; central producedarea of tergal border low
and obtuse; inner lobe of outer clististylesmall, beak of inner styleslender
Prattiana sP. n.

Dolichopeza(Megistomastix)acutiloba Alexander
D o t i c h o p e z o( M e g i s t o m a s t i x )a c u t i l o b a A l e x a n d e r ;J o u r . A g r . U n i v .
l ? 9 - 1 8 0 ,f i g . 6 ( d h y p o p y g i u m ) ; 1 9 3 7 .

Puerto Rico, 2l:

Typesfront the LuquilloMountains,PuertoRico,June7-ll,
Male hypopygium(Fig. l3).
1935,takenby JulfoGarcfa-Diaz.

Dolichopeza(Megistomastix)borinqueniasp. n.
Size relatively small (wing about 7.5 mm.); general coloration ofthorax yellowed,the darkenedpattern restricted,pleura without markings;
male antennaewith normal flagellar vestiturelong and delicate,exceeding
the verticils;wings brownishyellow, stigma and a seam over the anterior
corclbrown, petioleof cell M' and m subequal;male hypopygiumunusuallysimple and unmodified, especially the dististyles, lateral tergal arms
appearingas narrow glabrous blades.

172

A I e x a n d e r, TropicalAmericanTipulidae(Vl{l)M a I e.
Length, about 2.8-8 mm.; wing, 7.3-7.S mm.;
antenna,about 8-9 mm. Rostrumand palpi brown. Antennaeof
nlale elongate,exceedingthe wing or body, brown throughout;flagellarsegmentsvery long; normal vestiturelong and delicate,
the longestabout one-fourth the segment,verticilssnraller.Headbrown.
Prothoraxyellow. Mesonotumyellowed,the brown patternvery restricted,including the scutal lobes and vague areas on
praescutum;pleura yellow, virtually unpatterned.Haltereswith
stemyellow,knob slightlydarker.Legswith coxaeand trochanters
yellowl remainderof legsbrown,femoralbasesrestrictedly
yellowed. Wings brownishyellow, prearcularand costal f ields slightly
brighter yellow; stigma and a conspicuousconfluent cloud on
anteriorcord brown,posteriorcord morenarrowlydarkened;veinsbrownish yellow, slightly darker in the patternedareas. Cells
beyondcord with abundanttrichia, includingalso the outer ends
of cells Cu and /s/ A; trichia of stigma stronger and darker.
Venation: Petioleof cell M t variablein length,in type subequal
to or shorter than m, in one paratype with the cell virtuallysessile.
Abdomenchiefly yellow, the incisuresrestrictedlyinfuscated,the basaldarkeningsbroader;hypopygiumbrownishyellow.Malehypopygium( Fig. | 4) of unusuallygeneralized conformationf ora memberof this subgenus,
as shownparticularlyby the dististyles.
Ninth tergite, t, with lateral tergal arms or lobes appearingas
slender glabrous blades, inner edge near base slightly moreproduced;central lobe low, the nearly truncateborder with relatively few long black setae, the lateral ones longest.Outerdististyle,d, a flattenedbtadethat is gently narrowedoutwardly,
apex very obtuse,on inner rnarginnear base with a very small
lobe that is provided with few stout black setae; inner stylesmaller,boomerang-shaped,
outer end glabrous,obtuse,on f ace
with a low lobe providedwith a brush of long setae,nearerthebase with a very low glabrousblade.
Habitat: PuertoRico.
Holotype,d , EI Yunque,Luquillo National Forest,altitude2,600feet, May 25, 1945 (H. D. Pratt). Paratopotypes,
3 6 6,
May 30-31, 1945.
This species is the most generalizedmember of the subgenus so
far discovered in Puerto Rico, especially in hypopygial structure, withboth dististyles being virtually unmoclified.

Studia Ent., vol. 12, fasc. 14, novembro1969

t73

9

'b
73

14

FiS. 9.
Fig. 8. Dolichopeza (Megistomastix) domingensis Atexander; venation.
Fig. 10. Dolichopeza
Dolichopeza (Megtstomostix) jenaro sp. n.; venation.
(Megtstomastix) prattiana sp. n.; venation.
Fig. I l. Dolichopeza (Megistomastix)
Fig. 12. Dolichopeza (Megistomastix) prattiana
multitila sp. n.; male antenna.
Fig. 13. Dolichopeza (Megistomastlx) acutiloba Alexander;
sp. n. ; male antenna.
Fig. 14. Dolichopeza (Megistomastix) borinquenia sp. n.; male
male hypopygium.
trypopygium. - Fig. 15. Dolichopeza (Megistomastix) ienaro sp. n.; male hypopygium.
Fig. 16. Dolichopez.a- (Mysistomastix) -multi.fita- rp.. n.; .male hypopygium.
Symbols: b, basistyle; d, dististyle; f, tergite.

t7.L

A l e x a n c le r , T r ' p i c a l A m e r i c a nT i l l , l i r l a e ( v l t t )Dolichopeza(Megistomastix)jenaro sp. n.
Antennaetlf male with nclrmal vestiture short, to about one-halft h e l e n g t h o f t h e v e r t i c i l s ;w i n g s f a i n t l y d a r k e n e c lv, i r t u a l l y u n p a t t e r n e r l
e x c e p t f o r t h e . b r o w n s t i g m a , c e l l M L l o n g , s e s s i l eo r v i r t u a l l y s o ; n r a l ehypopygiumwith lateral tergal arms unequally bilobecl,both lobes w'itlr
tips glabrous; central lobe very broacl; beak of clististylenarro\4,;guterd i s t i s t y l e u n e q u a l l yb i l o b e d ,t h e o u t e r a r m a n e l o n g a t ec l u b .

M a leL e n g t ha
, b o u t8 - g m m . ; w i n g ,g - g m m . ; a n t e n n a ,about 9- 10 rnm. Frontal prolongationof head and palpi lightbrown, distal half of terminalsegmentof latter darker brown.
Antennaewitlr scapeand pediceldark brown, f lagellunrblack;flagellarsegmentseklngate,the normalvestiturerelativelyshrlrt,'verticils.
up to aboutone-halfthe blackened
Head brownishgray.
Pronotalscututttbrownishyellow, scutellunrcleareryellow.
Meson<-ltal
praescutumbrownish gray with three darker brown
stripes,the centralone narrow; scutal lobessimilarlydarkenecl,
posteriorscleritesIight gray.Pleuraobscureyellow,conspicuously
patternedwith dark brown on anepisternum,
ventralsternopleurite
and meron.Haltereswith stem yellow, knob brown. Legs withcoxaeand trochanters
yellowed;femorabrownishyellow,brighter
basally,tips darker; tibiae and tarsi darker brown. Wings ( F ig.
9 ) f aintly darkened,virtually unpatternedexceptf or the brownstigma; veins brown. Macrotrichiain cells beyondgeneralleveltrf outer end of cell I st Mr, includingalso the stigma and outerends of cells M4 to /sf A. Venation:Cell Mr long, sessile,incasesbroadly So, in others more punctiform;cell /s/ Mz longand narrow, Ms-q short, usually less than one-half the basal
sectionol M1-,, in casesto abouttwo-thirdsthis length,in othercasesvirtually lacking,in still otherswith m-cu on M4 beyond
the base.
Abdonrinal tergites chiefly dark brown, posterior bordersof outer segmentsnarrowly yellowed; sternitesinconspicuously
bicolored,the yellow apicesbroad; hypopygiumbrownishyellow.
Male hypopygiunr( Fig. l5) with lateral tergal arnrs, t, broad,the outer end unequallybilobed,both bladesglabrous,the apical
one narrower,subacute
I central tergal region very broad, with
abundantlong setae.Outer dististyle,d, unequallybilobed,outerarnl an elongateclub with long coarsesetae,on inner face witha dense concentrationof setae of various lengths, inner lobe
sntaller,dilated at apex, both lobes with abundantlong setae;

stuclia Ent., vol. 12, fasc. l-4, novembrolt)6g

l7s

inner style with beak narrow, dorsal crest erect, with blackened
setae,posteriorcrest a flattened glabrous blade.
Habitat: Puerto Rico.
Holotype,d, El Yunque,Luquillo National Forest,altitude
alrtrut2,000feet, November27-28,rg43 (H. D. pratt andJenaroMafdonado-Capriles).
Paratopotypes,
numerousd d , 2,600 feet,
May 25-30, l94b (H. D. Pratt).
I am pleasedttl name this speciesfrlr the collector of the type,'toJenaro MaldonadoCapriles,of Cayey RegionalCollege,Puerto Ricol
whr-rmI am indebteclfor severalinterestingTipulittaefrom Puerto Ricoand northernSouth America.As is inclicatecl
by the key to the regional
speciesthe most similar form is Dolichopeza(Megistomastix)oriiitotro
Alexancler,separateclby the characterslistecl.

Dolichopeza(Megistornastix)multifila sp. n.
Antennaeof male elongate,about rine-fourthlonger than
the wing;
normal vestiture rlf flagellar segmentsvery long un,rabunclant,auoi,tone-halfthe longestsegmentanclmuch e*.e.ding Tn.
verticits;mesonotum
br'wnish yellow, patternecrwith crarker glow-n; wrngs
weakry tinged
with brown, sJi.smaclarker, petiole of cell Mr ,r,u.irur
hypopygiumwith lateral tergal arms simple, tips obtuse; to m; maleouter arm of,uter clististylebrraclly expanclecl,
margin smroth.

M a l e . - L e n g t h ,a b o u tT - T . sm m . ; w i n g , 7 - g m m . ; a n t e n n a ,about 9-9.5 nlm. Rostrunrand palpi dark brown. Antennaeofnrale very long, about one-half longer than the wing;
scape,
pediceland proximal end of first flagellar segnrentligfit brown,
the renrainderdarker; flageltar segmentslong cylindrical,
thenorntal vestiture very long and abundant, about one-half thelongestsegnlents,ntuch exceedingthe only slightly differentiatedunilaterallydistributedverticils.Head brown.
Pronotunt yellow, darker medially. Mesonotal praescutum
brownishyellow, vaguelypatternedwith darker brown, including
the centralline and lateralborders,posteriorscleriteslight brown.
Pleura brownish yellow, dorsopleuralmembrane light yellow.
Halteresbrown, base of stent narrowly dull orange.-Legs withcoxae and trochantersyellowed; remainderbrown, femoral bases
yellowed.wings weakly tinged with brown, stignradarker; veinsdark brown. venation: cell M 1 with petiole subequalto or alittle shorter than m.
Abdominalsegmentsbrownishyellow, patternedbasallywith
clark brown, more broadly so on tergites; hypopygiumbrownish
yellow. Male hypopygium ( Fig. l6) with lateral tergal arms,
t, stout, tips obtuse,apical setae relativelyshort; central tergal

176

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

region low-convex, obtuse, with relatively f ew setae. Outerdististyle,d, unequallybilobed,outer arm very long, apex broadly
expanded,ntargins smooth, inner lobe slender,bent on outer
half, margin with short black setae;inner style with beak very
slender,dorsal crest relativelystout, posteriorcrest a flattened
glabrousblade.
Habitat: Puerto Rico.
Holotype,d , El Yunque,Luquillo National Forest,altitude
3 6 6,
2,600feet,May 31, 1945 (H. D. Pratt). Paratopotypes,
May 25-30, 1945 (H. D. Pratt).
In the nature of vestitureof the male antennae,the presentfly ismost as in Dolichopeza(Megistomastix)portoricensis(Alexander),differing in the charactersprovidedin the key.
Dolichopeza(Megistomastix)obtusiloba Alexander
Dotichopeza (Megistomastix) obtusiloba Alexander; Jour. Agr. Univ. Puerto Rico, 2l:
1 8 0 - 1 8 1 ,f i g . 7 ( d h y p o p y g i u m;) 1 9 3 7 .

Types f rom the El Yunque trail, Luquillo National Forest,
P u e r t oR i c o ,a t K m . 1 . 3 ,J u l y 2 7 , 1 9 3 5 ,c o l l e c t ebdy J u l i oG a r c f a Diaz. Male hypopygiunr( Fig. 17).

Dolichopeza(Megistomastix)polytrichasp. n.
S i z e l a r g e ( w i n g o f m a l e t o 8 . 5 m m . ) ; m a l e a n t e n n a es l i g h t l y e x c e e c l ing the body or wings, normal vestiture of flagellar segmentsclarkened,e l o n g a t e ,e x c e e d i n gt h e v e r t i c i l s ; m a c r o t r i c h i ao f w i n g c e l l s u n u s u a l l yabundant,including all cells beyond cord, as well as the outer ends ofcells R, M, Cu, and /sf A; male hypopygium with apices of lateral,
tergal arms narrowed and glabrous.

M a I e.
Length, about 8-8.5 mm.; wing, 8-8.5 nlnt.;
antenna,about 9- l0 mm. Frontal prolongationof head obscureyellow; palpi beyond the basal segmentdark brown. Antennac
of male slightly exceedingthe wing, black; f lagellar segments
very long-cylindrical,
normalvestituredark brown, exceedingthe
verticils,the longestabout one-thirdthe segrnent.Head brown,
restrictedlyorange immediatelybehind the antennalbases.
Pronotumobscureyellow, patternedwith brown. Mesonotal
praescutumbrownishyellow, with brown stripesthat includean
abbreviatedcentral'areaand a U-shapedlateralmark that involves
the darker outer margin; scutal lobes reddish brown to darker
brown; posterior scleritesof notum brownish yellow to lightbrowir. Pleura dark brown with an obscure yellow ventral

Studia Ent., vol. 12, fasc. l-4, novembro1969

t77

Fig. 17. Dotichopeza (Megistomastix) obtusiloba Alexander;- male .hypopygilm. -=FiE. 18. Dotichopbza(Megiitomastix) polytricha sp. l.; ma-le-hypopvg.imu.
I!g. 19.
Dotichopeza (Migistomastix) portoricensis (Alexander); male hypopygiury:l. Fig'.2U.
Dotichopeza(ne&stomastix) firattiana sp. n.; male hypopy_giuq:- lig. 2l-._Ct-enophora
22. CtenoOlora (Pselllophor-a)Fig.
(Psetliophora') riargarita Alexander; venation. -basistyle;d, dististyle; p, phallob,
Symbols:
male antenna.
margarita Al-exande-r;some; t, tergite.
t2

178

A I e x a n d e r, TropicalAmericanTipulidae(VIII)

longitudinalstripethat widensbehind,dorsopleuralregionyellowed. Haltereswith stemyellow,knob weakly darkened.Legs withcoxae brown, trochantersyellow; femora brownish yellow, tips
darker, tibiae and tarsi brown. Wings weakly darkened,stigma
and veins dark brown. Macrotrichiaof cells unusuallyabundant,
includingall cells beyondcord, as well as outer ends of cells
R, M, Cu and Ist A. Venation:Petioleof cell ML slightlylonger
than m.
Abdomen bicolored,incisures,including the bases of segments,dark brown, apicesyellowed.Margin of eighth sternite
trilobed,with long setae.Male hypopygium(Fig. l8) with lateral
tergal arms,/, stout,narrowedoutwardly,terminatingin a narrowglabrousblade, its tip obtuse;central tergal region low convex,
the marginwith severalstout black setae,more ventralextensive,ly at base of lateral arms with a dusky setiferouscushion as
is commonin the subgenus.Outer dististyle,d, with outer arm
long, dilated outwardly into a clavate head, on its inner facewith a low darkenedf lange that is vaguely crenulatebut notdistinctly toothed,oS in portoricensis,inner lobe compact,innerstyle as f igured
Habitat: Puerto Rico.
Holotype,6 , El Yunque,Luquillo NationalForest,altitude
3 6 6,
2,600feet,May 31, 1945 (H. D. Pratt). Paratopotypes,
Alay 25-30, lg45 (H. D. Pratt).
The most similar species is Dolichopeza (Megistomastix) porto'
ricensis(Alexander),as indicatedin the accompanyingkey. Other regional
members of the subgenusthat have macrotrichia in outer end of cell
R of the wings are D. (M.) acutiloba Alexander and D. (M.) obtttsiloba
Alexander, readily told by the antennal vestiture and the hypopygial
structure, 3s shown in the key.

Dolichopeza(Megistomastix)portoricensis(Alexander)
i l l e g i s t o m a s t i xp o r t o r i c e n s i s A l e x a n d e r ; P s y c h e , l 9 : 6 3 - 6 6 , p l . 5 , w i t h 3 f i g s .; 1 9 1 2 .

The uniquetype ntalewas capturedon the El Yunquetrail,
Puerto Rico, at 2,800 feet on February 20, 1900, by C. W.
Richmond,preservedin the United States National Museum(AccessionNo. 14,592).Nunrerousfurther specimenslater weretaken on El Yunque,chiefly at about 2,600 feet, in 1943 and1945by Dr. Harry D. Pratt and Mr. JenaroMaldonadoCapriles.
In the figures abovecited, the conspicuous
vestitureof thenrale antennaeand arrangementof the nracrotrichiain the wingcells were shown satisfactorily.The drawing of the nralehypo-StudiaEnt., vol. 12, fasc. l-4, novembro1969

179

pygiumwas madefrorn the dried type and is highly diagrammatic.
A more satisfactory f igure is providedherewith ( Fig. l9).
now availablecertainindividuals
In the seriesof specimens
have the true verticils greatly reducedor lacking and with thenormal vestitureslightly longer than in other specimenswherethe verticils are present,though reduced.I believe that thisconditionof the verticils is the normal one, as shown by thetype. In other regards,ds in the venation,wing trichiation,and
structureof the hypopygiunr,I can see no essentialdifferences
betweenthe two sets of specimenspresentlyreferred to thisspecies.

Dolichopeza(Megistomastix)prattianasp. n.
Antennaeof male with normal vestitureof flagellar segmentsshort,about one-half the verticils; wings faintly darkened,with a brown seamover the cord; petiole of cell Mt subequalto or longer than m; malehypopygiumwith lateral tergal arms stout, their apices obtuse, medianregion low-convex.

M a I e.
L e n g t h , a b o u t 8 - 8 . 5 n l n r . ; w i n g, 6 . 5 - 7 m m . ;antenna, about 6.5-7 mm. Rostrum dark brown, palpi black.
Antennae of male ( Fig. 12) elongate,subequalto wing; scapebrownish black, remainderof organ dark brown; f lagellar seglnents long-cylindrical,the whitened normal vestitureshort andclense,only about one-half as long as the blackenedverticils.
Head light brown.
Pronotum light yellow, sides slightly darker. Mesonotal
praescutumwith f our light brown stripes that are very narrowly
bordered exteriorly by slightly darker brown, the central interspace similarly colored, humeral region yellowed; scutal lobes
light brown, central region and remainderof notum pale brownish yellow, sparsely pruinose.Pleura dark brown with a vague
more yellowedlongitudinalstripe extendingfrom behind the fore
coxaebackwardto the metapleuralregion; dorsopleuralmembranelight yellow. Haltereswith stem f aintly darkened,base yellow,
knob more darkened.Legs with fore coxae dark brown, remaining
coxae more yellowed;trochantersyellow; remainderof legs lightbrown. Wings ( Fig. l0) f aintly darkened,prearcularand costal
regionssliglttly nlore yellowecl;stigrnaand a confluent seamoveranterior cord darker brolvn, narrowed behind; veins brown.
Macrotrichiain cells beyond cord, including also the outer ends
of cells Cu and I st A. Venation: Rs in transversealignmentwith
rernainderof cord, subequal to basal section of Rn-u; petioleof cell M 1 subequalto m.
t2.

180

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

Abdominal tergites bicolored,bases narrowly brown, outerparts paler; hypopygiumbrown. Male hypopygium (Fig. 20)

with lateraltergal arms,t, stout,with a vaguelow flange nearouter end, apex obtuse;centraltergal regionlow convex.Outerdististyle,d, unequallybilobed,outer arm longer,clavate,withlong delicatesetae,apex broadlyobtuse;outerend of inner lobe
dilated,the setaeshorterand stouter;innerstyle with beak pale,triangular in outline, dorsal crest darkened,narrow, with aof black setae,posteriorcrest a flattenedglabrousconcentration
dark colored blade.
Habitat: Puerto Rico.
Holotype,6 , El Yunque,Luquillo NationalForest,altitude2,000 feet, November27, 1943 ( H. D. Pratt and J. Maldonado
numerous6 6 , 2,600 feet, May 30 Capriles).Paratopotypes,
June 6, 1945 (H. D. Pratt).
I have named this interestingfly for my former student and outstandingentomologist,Dr. Harry D. Pratt, who collectedmany speciesof Tipulidae in Puerto Rico. As is shown by the key the most similar
speciesis Dotichopeza(Megistomastix)obtusilobaAlexander,readily toldby the hypopygialstructure,3s described.

'

ElnorettaAlexander

E l n o r e t t a A l e x a n d e r ; D i p t e r a P a t a g o n i a .- a n d- S o u t h. C h i l e , l : l 5 - 1 6 , f i g . 3 l
f i g . 1I I ( a n t e n n a) ; ( t y p e a c r a c a n t l t o l d e sA l e x a n d e r ; r n o n o t y p i c ) .

(wing),

There is a single known species,apparentlyallied to theAustralian.genus AcrocanthaSkuse. The antennaeof the male( Fig. 25) are I 2-segmented;f lagellar segmentsI to I eachwith a single long terminal branch, all being longer than thesegmentwith the exceptionof the last which is slightly shorter;f lagellar segmentswith scatteredstrong setae on outer f ace,branchesstrongly wrinkled, without setae but with numerous
nearly circular areas that lack setae. In the female ( Fig. 26)
antennae l3-segmentedor l4-segmented,the lower f ace ofproximal three f lagellar segmentsslightly produced to appear
serrate; f ourth segmentstouter than the succeedingones, thelatter cylindrical,providedwith very long setae,the longestmuch
exceedingthe segments.Wings (Fig. 23) with cell ML sessile.
Other structuralfeaturesof the genusare discussedin the original
definition.
List of Species
acracanthoidesAlexander.

Chile.

Studia Ent., vol. 12, fasc. 1-4, novembro 1969

l8l

Holorusia Loew
Ho_lorusiaLoew; Berl. Ent. Zs. 7: 276, 1863; type, grandls Bergroth :_ L o e w , p r e o c c u p i e dn a m e ; N e a r c t i c ; m o n o t y p i c .
Ctenacroscelis Enderlein; Zool. Jahrb., Syst., 32
l-2; l9l2; (typeEnderlein; original designation).

rubiginosa
dohrnianus

The recent paper by Vane-Wright is of great importancein an understandingof this genus and its synonymy.* Holorusia
was based on a single speciesfrom western North America but
now is united with the extensivegenus Ctenacroscelis
Enderlein,with approximately90 f .orms describedf ronr the Old World,with a marked concentrationof speciesin easternAsia. During
the past half century a large number of chiefly South American
specieshave been assigned to H olorusia by the present writer
but it has been shown by Vane-Wright that thesemore properly
should be placed in two other genericgroups,IschnotomaSkuse
and ZelandotipulaAlexander,under which names they may beconsultedin the presentpaper.As it now stands,Holorusicin the.
New World includes only the genotype, grandis Bergroth,(rubiginosaLoew), describedf rom California,with a wide range
in westernNorth Anrerica.Vane-Wright suggeststhat this isolated speciesmay have reachedits present range via the former
Bering land connectionand this belief is furtheredby the recent
discoveryof the speciesin central Alaska, where two females
have been taken in McKinley National Park (by F. Morand;in the United States National Museum, reported to the writerby Dr. Alan Stone).

lschnotomaSkuse
I s c h n o t o m aS k u s e ; - - P r o cL. i n n . S o c . N e w S o u t h W a l e s , ( z .\ 5 : I l . l - l t 5 ; l 8 g 0 ; ( t y p e
Tlpyta _eburnea Walker, as Tipula serricornis Macquart, designated by Ale'xairdir,
1920:53).

In a recent paper, discussedunder the preceding genus,
Vane-Wright has made a critical survey of IschnotomaSkuseand some related groups. In Tropical America he recognized
two generic entities f ormcrly included in our conceptionof thegenus H olorusia, these bcing I schnotoma and Zelandotipula
Alexander.In the f ornrer he recognizedtwo subgenerain thisfaunal area,thesebeing Ischnotomas. s. and IcriomastaxEnder* ) Vane-Wright, R. I., A re-assessmentof the genera Holorusia
Loew (- Ctenacrosceli.s
Enderlein),IschnotomaSkuse and Zelandotipula
Atexander(Diptera: Tipulidae),with notes on their phylogenyand zoogeography.
Jour. nat. Hist., I : 5ll-547, 46 f i g s . ; 1 9 6 7 .

t82

A I e x a n d e r, TropicalAmericanTipulidae(VIII)

lein, to which shouldbe addedthe more recentlyproposedNeotipula Alexander. Vane-Wright placed virtually all American
speciesin I criomastoxbut it now appearsthat these species
would be placed more satisfactorilyin typical Ischnotomoand
it seems questionableto me as to whether I criomastax,and
probably also N eotipula, can be maintainedas distinct f rom
the male hypopygiaof virtually
Ischnotomo.I have re-exarnined
all speciesknown in the genus and these show a noteworthy
rangein structurein both the Australianand Neotropicalspecies.
In the presentreport I am assigningthe majority of the regional
restrictingIcriomastaxto thespeciesto the subgenus/scftnotoma,
so-calledjuiuyensisgroup, includingten species,all f rom Argentina and Brazil. N eotipulahas f our species,all being moreof the other subgenera.Twonorthernthan the representatives
aberrant species,problematicaAlexanderand penai Alexander,
for the presentare placed in typical I schnotoma.
in detailby Vanehavebeendiscussed
The genericcharacters
Wright. The normal type of antennais illustrated(Fig. 27,
decorata)but certain other specieshave a greater enlargement
Wingsof the flagellarsegmentsto producea weak pectination.
with veins unusuallyglabrous,the trichia more numerouson
veins C, Sc and R, , more rarely on the posteriorouter radialveins. Squamanaked or weakly setose.Male hypopygiumwith
the ninth tergite emarginatein I schnotomo,entire and produced
nrediallyin I criomqstox.It should be noted that in the latter
subgenus,including the type, ocellataEnderlein,that spinoid
setaeare presenton the inner dististyle,somewhatas f ound in
LeptotarsAsand Zelondotipulaand apparentlyindicating a relationshipto thesegroups.In I criomastaxthesespinoidelements
are relativelyfew in number,being reducedto a single spine
in calliopewhile being lacking in nitra.
I schnotomavasifera (Cockerell
Of interestis a f ossilspecies,
and Haines, l92l), describedfront the Bembridgebeds of theIsle of Wight in the EnglishChannel,assignedto the Oligocene
and originally placed in H olorusia.Vane-Wright has re-studiecl
the type specinlenof this and emphasizesthe close generalof this fossil to the recentIschnotomo(lschnotoma)
resemblancepor (Walker) of easternAustralia.
Varre-Wright suggests that iuiuyensis may prove to be
knownnudicornis(Macquart).
identicalwith the still insufficiently
The latter was describedf rom the vicinity of Buenos Aires
whereasjujuyensisis fronr the westernnrore mountainouspro-

1,93

StucliaEnt., vol. 12, fasc. l'4, novembro196927-_ Fig.. 2{. Ischnotoma
Fig. 23. Etnoretta acracanthoides Atexander; vertation_._(Philippi);
venation.
(I"schnotoma)decorata''nraleFig. 26. Elnoretta acracanthoides.. Alexa-nder;
antenna. -'hlexander;. f emale(Philippi);
male
antenna.
(lschnotoma)decorata
Frg. 27. Ischnotoma
antenna.

vinces of Argentina. There are certain Brazilian speciesthat
are as closeto BuenosAires as is iuiuyensis,andgeographically
nudicornismay be found to pertain to one of these or perhaps
may representa further memberof the ocellatagroup.
The immaturestages of iuiuyensisoccur in wet mats ofplants growing along stream margins. These have been welldescribedand figured by Carlos Bruch (Contribucional conocimientode los TipulidosArgentinos( Diptera). Physis,17: l-28,
pp. 12-16,pl. l, fig. 2 (habitatof immaturel l plates-references
stages),pl. 3, f ig. 7, pl. 7, f igs. 26-28 (larva and pupa), pl. 8,f igs. I - 16 (detailsof egg, larva and pupa).

184

Alexander,T r o p i c a lA m e r i c a nT i p u l i d a e( V I I I )List of Species

SubgenusI schnotoma Skuse
concinna (Philippi), as Tipula
Chile.
decoratadecorata (Philippi), as Tipula
Chile.
d. araucana (Alexander)
Chile.
delpontei (Alexander), ?s Holorusia
Argentina, Chile.
as Holorusia - Argentina.
fagetorumfagetorum (Alexander),'Holorusia
Argentina.
f. trunculata (Alexander), 4s
Chile.
f rauenfeldi (Schiner), as Tipula(may be decorata (Philippi).
Chile.
fttscostigmosa(Alexander), ?S Holorusia
homochroa(Alexander),?S Holorusia
Chile.
larotypa (Alexander), ?s Holorusia
Chile.
pefiai (Alexander),?s Holorusia (lschnotoma)
Chile.
guzmani (Alexander), dS Holorusia (lschnotoma)Syn.
lapsus for pefiai.
porteri (Alexander),?s Holorusia
Chile.
postnotalis (Alexander), ds Holorusia
Argentina.
problematica(Alexander), ?S Holoru.sia Chile.
ruf ostigmosa (Macquart), as Tipula
Chile.
schineriana(Alexander),ds Holorusia
Chile.
shannttniana(Alexander),3S Holorusia
Chile.
silvai (Alexancler),3SHolorusia
Chile.
vittigera (Philippi), as Tipula
Chile.
SubgenusIcriomastax Enderlein
I c r i o m a s t a x E n d e rl e ijn ? o 9 1 ..Ja h1b.
, Syst., 32: 9;m o n o t y p i ca n d o r i g i n a l d e s i g n a t i o) n.

l9l2;( type octilata Ender l ei r r ,antinymtrtha(Alexander), ?s Holorusia
Brazil.
calliope (Alexander), ?s Holorusia
Brazil.
euterpe (Alexander), ds Holorusia
Brazil.
helios (Alexander), 3s Holorusia
Brazil.
juiuyensis (Alexander),flS Holorusia
Argentina.
(may be nudicornis (Macquart), auct. Vane-\Mright)
nitra (Alexander), 4S Holorusia
Brazil.
nudicornis (Macquart), as Tipula
Argentina.
ocellata (Enderlein), as Icriomastax
Brazil.
phaeton (Alexander), &s Holorusia
Brazil.
zikani (Alexander), 8S Holorusia
Brazil.
SubgenusN eotipula Alexander
Tl p u l a . ( N e oti p u l a ). A l e xa .n d e rjJogr . r !. Y. Ent. Soc., 48: 105- 106; 1940; ( ty pe
p e c t i n e l l aA l e x a n d e r ,o r i g i n a l d e s i g n a t i o n ) .

maya (Alexander),ns Tipula
Guatenrala.
paprzyckii (Alexander), ds Tipula (Neotipula)
pectinella (Alexander), ?S Tipula (Neotipula)pefi.ata(Alexander),4s Holorusia (Holorusia)

Peru.
Panama.
Ecuador, Peru.

Studia Ent., vol. 12, fasc. l-1, novembro1969

185

LeptotarsusGudrin-MdnevilleL e p t o t a r s u sG u d r i n - M d n e v i l l e V; oy. corvette La Coquille, Zool., 2. pt. 2: 286, pl. 20,
f i g . l ; 1 8 3 0 ; ( t y p e L e p t o t a r s u sM a c q u a r t i O u d r i n - M e n e v i l l e A; ustralian; monotypic).

As presentlyrecognizedin the Neotropicalregion,the genus
Leptotarsus includes eleven subgenericgroups. As had been
indicatedby the writer in various recent papers ( as in South
African Animal Life, l0: 244-245; 1964) certain groups ofTipulinc crane f lies that had been maintainedas valid genera
now appear to be of subgenericranl< while some others inpossiblywill f all invariousparts of the SouthernHemispherethe syynonynry..
It had been believedthat the oldest genus in the complexof the problem
was Longurio Loew, 1869,but a consideration
being ntade in Australiaby Dr. N. Dobrotworsky,and by theAustraliangenusLeptotarsus
writer, indicatesthat the essentiallybelongshere and apparentlyis the valid name for the genus.
In the Tipulinaethereare five still oldernamesbut theseevidently
clo not apply to the present group. These nafiles are Tipula

Linnaeus,1758; NephrotomoMeigen, 1803; CtenophoraMeigen'ro Wiedenlann,I 828 (MaekistoceraWiedemann,
I 803; Me gistoce
I 82I ) , and DolichopezaCurtis, 1825.
The chief charactersused in referring these groups to asingle genus are found in the nrale hypopygium,especiallythe
clististyles and phallosonre. Virtually all species in this complex

have modified setae on the inner dististyle that have been
to true spines.
designatedas spinoid setcrt:f rom their resemblance
There seentsto be no question but that this characterindicatesaffinities among the various groups despite their differencesin
seneral appearance.It may be indicatedthat comparablespinoid
setae on the dististyle occur in other groups of the primitive
Tipulinae which on other charactersare consideredas being
rlistinct f ronr Leptotarsus, such including Clytocosmus Skuse,PtatyphttsiaSkuse,ZelondotipulaAlexander,and others.Additional
to the dististylesthe very generalizedphallosorne,with unusuallyalso appearsto indicateclose
and aecleagus,
simple gonapophyses
but is f ound in various other genera in theinterrelationships
Tipulinae.
Itshould be

noted

that

two

furthergroupsthatareMacromastixOsten Sacken,1886,and
nraintainedas subgenera,
HehromastixSkuse, 1890, are Australasianin distribution.The

A I e x a n d e r , T r o p i c a l A m e r i c a nT i p u l i d a e ( V I I I )186variousrecordsf or thesegroups f rom Tropical Anrericapertain
to other subgenera,
chiefly to Longurio.
To the writer there appearsto be an analogy between
and Limonia Meigen,the typical
Leptotarsus,
as here discussed,
genusof the subfamilyLimoniinae.In most subgeneraof Limonia
there similarly are found spinoid setae on the dististylethat
provideimportantcharactersin definingthe variousgroups.Also
range
in Limonia,as in the presentgenus,thereis a considerable
in body sizeand structure,in venation,and in the relativelengtlt
and slendernessof the legs. In someLimonia s. s. and in MetalimnobiaMatsumurathe body is large and bulky, a condition
to that found in Leptotarsusand Semnotes
somewhatcomparable
Skuse
in the Tipulinae,while stitl othergroups,as Thrypticomyio
or EuglochinaAlexander,have a delicateetherealstructure,witlt
to the condition
slenderbodiesand unusuallylong legs,analagous
f ound in Tonypremna and sonle allied subgenerain the present

genus.
In most subgeneraof Leptotarsusthe antennaeof both sexes

are very small but speciesoccurwherethe maleshave the organ
excessively
elongated,as in Araucomyia(as paulseni),Macromastix(as coslalisSwederus),
Hobromastix,
and in certainspecies
that presentlyare assignedto Longurio.
The intmaturestages of a nunrberof specieshave been
described,especiallyin the subgenusLongurio in South Africa

lies of the South-WestCape (Diptera,[H. G. Wood, The crane-f
Tipuloidea). Ann. South African Mus., 39: l-327, 105 f igs.;
19521.Most of these speciesoccur in soil among clumps ofplantsof the monocotyledonous
family Restionaceae,
often underrelativelydry conditions.In America,the type of the subgenus
Longurio (testaceusLoew) has been f ound in wet sandy soilat the margirtsof springs, while the related L. rivertonensis(Johnson)is nlore nearly aquatic,the larvae occurringin cold
springs [referencesin Alexander,The crane-flies of New York.
Part 2: 990-993, f igs. 488-491 ( testaceus), f ig. 492 (rivertonensis);19201.In New Zealand,variousspeciesthat are referred to the subgeneraLongurio and Macromastix,have been
f ound living in soil beneathleaf mold, as describedby J. S.
Rogers(Trans.New ZealandInst.,58: 301-309,14 figs.; 1927).
The immaturesof speciesin the subgenusChlorotipulaAlexander
havebeentound living in decayingwood in New Zealand,including L. (C.) olbistigma (Edwards) and L. (C.) holochlora( Nowicki), the latter having been describedand f igured b-v G.

Stuclia Ent., vol. 12, fasc. l-4, novembro 1969

187

V. Hudson(ManualNew ZealandInsects,pp. 47-48,pl. 5, figs.
flarva,pupal 1892),in casesbeingfound in the decayingwoodof old buildings and of some economicimportance.
List of Species
SubgenusAldrovandia EnderleinA l d r o v a n d i a E n d e r l e i n ; Z o o l . J a h r b . , S y s t ,. 3 2 : 5 2 , f i g .
g e s n e r i E n d e r l e i n; m o n o t y p i c a n d o r i g i n a l d e s i g n a t i o)n.

gesneri (Enderlein), as Aldrovandia(venation); l9l2:(type

Brazil.

SubgenusArancomyiaAlexander
Araucomyia Alexander; Diptera Patagonia and South Chile, l: t4-15;. 1929; asM a c r o m a s t i x - A r a u c o m y i a(; t y p e p a u l s e n i ( P h i l i p p i ) ; o r i g i n a l d e s i g n a t i o n ) .

brevihirsutus (Alexander), as Macromastix-Araucomyia Chile.
Chile.
paulseni (Philippi), as Tipula(Alexander),as Macromastix-Araucomyia Chile.
penittts
SubgenusCeoneura Alexander
1 9 2 4 ; a s T a n y p r e m n a - C e o n e u r a( t; y p eCeoneuraAtexander;Ent. News,35:289;
idloneura (Alexander); monotypic and original designation).

(Fig. 28, venation) idioneurus (Alexander), as Tanypremna-Ceoneura
British Guiana.
Subgenus Leptotarsus Guirin-Mineville
Reference as under the genus; type Leptotarsus Macquarti.

annulipes (Philippi), as Tipulain this subgenus.

Chile. May not be placed correctly

SubgenusLimoniodes Alexander
Limoniodes Alexander; Ann. Mag. Nat. Hist., (t t ) 1: 337; t938; as Tanyp.remnaL i m o n i o d e s ,t.y p e s u l p h u r e l l a ( A t e x a n d e r ) ; m o n o t y p i c a n d o r i g i n a l d e s i g n a t i o n ) .

(Fig. 29, venation)
sttlphurella(Alexancler),as Tanypremna-Limoniodes
Ecuador.

SubgenusLongurioLoew
L o n g u r l o L o e w ; D i p t e r a A m e r i c a e s e p t e n t r i o n a l i si n d i g e n a , C e n t u r i a o c t a v a . B e r l i n .
E n t . Z e i t s c h r . , l 3 : 3 ; 1 8 6 9 ; ( t y p e t e s t a c e u sL o e w ; N e a r c t i c , o r i g i n a l d e s c r i p t i o n
and monotypic).

atrirostris (Alexander), as Macromastix
Chile.
horgmeieranussp. n. - (Fig. 38, 6 hypopygium)
Brazil.
brasiliae (Alexander), as Macromastix (Macromastix)
Brazil.
hrowni (Alexander), as Macromastix
Ecuador.
(Alexander),(Araucomyia)
Argentina, Chile.
bttllocki
as Macromastix(Alexander),(Macromastix)
as Macromastix
caparaona.
Brazil.
Brazil.
carreranus (Alexander), as Macromastix

188

Alexander,T r o p i c a lA m e r i c a nT i p u l i d a e( V I I I )chrysostigma(Alexander), as Macromastix (Macromasfix)
Brazil.
decoloratus(Alexander),as Habromastix Paraguay.
espinozai (Alexander), 4S Macromastix
Chile.
eucrypta (Alexander),as Longurio
Chile.
exemptus (Alexander), as Habromastix (Fig. 39, 6 hypopygium)
Brazil.
goyazana (Alexander), as Macromastix (Macromastix)
Brazil.
guimaraesi (Alexander), as Macromastix
Brazil.
gymnocera(Alexander),as Macromastix
Brazil.
helota sp. n. ( Fig. 40, 6 hypopygium)
Brazil.
huanucensis(Alexander),as Longurio (Fig. 36, venation) - Peru.
insidiostts(Alexander),as Macromastix(Macromasfix) Brazil.
ixion (Alexander),as Macromastix(Macromastix)
Brazil.
Iemniscatus(Alexander),as Habromastix
Brazil.
lustralis (Alexander), as Tipula
Brazil.
luteistigma(Alexander),as Macromastix(Araucomyia)
Brazil.
nahuelbutae(Alexander),as Macromastix Chile.
nigroabdominalis(Alexander),as Habromastix- Brazil.
nocivus (Alexander),as Longurio
Chile.
paraguayanus(Riedel), 4s Tipulodina
Paraguay.
(Alexander),as Tipula
pdroguayen.sls
Paraguay.
perglabratus(Alexander),as Longurio (Longurio)
Brazil.
phaedra (Alexander),as Macromastix
Brazil.
pygmaeus(Alexander),as MacromastixPeru.
rabelloi (Alexander),as Longurio (Longurio)
Brazil.
(Alexander),serotinella
as trlacroma.sfi-r Chile.
sessoris (Alexander), as Longurio (Longurio) (Fig. 37, venation) Peru.
stenostylus(Alexancler),
as Longurio (Longurio)
Ecuador.
.sfyx (Alexander),as Longurio (Longurio)
Brazil.
tijucanus (Alexander),as Macromastix(Macromastix)
Brazil.
(Alexander),as Macromastix(Macromastix) Brazil.
travassosanus
zikanella (Alexander), as Longurio (Longurio)
Brazil.
SubgenusPehlkea Enderlein
Pehlkea Enderlein; Z o o l . J a h r b . , .S
. y s t ,. 3 2 : l 5 ;m o n o t y p i ca n d o r i g i n a l d e s i g n a t i o n ) .

l9l2;( t y p e c o t u m h i a n u sE n d e r l e i ;ncolumbianus (Enderlein), as Pehlkea
Colombia.
pallitarsis (Alexander), as Tanypremna (pehlkea) - Ecuaclor.
reginus (Alexander), as Tanypremna
Colombia.
(Alexander),regultts
as Tanypremna (pehtkea) Ecuador.
Subgenus Tanypremno Osten Sacken
T a - n ^ 1 ' p r e m on sat e n . s a ^ c k e n ;- B i o l . c e n t .
' . lg, pl. l, fig. 2 (wing);.Amer., .Diptera, p^ . 1 8 8 6 ; ( t y p e o p i t i o O s t e n S a c k e n , o r i g i n a t c l e s i g n a t i o' nll).
s t e g a s m o n o t u s E n d - e r l e i n ; z o o l . J a h r 5 . , s y s t ., - 3 2 :; lgl2; (type longissimus
E n d e r l e i n ,o r i g i n a l d e s i g n a t i o n ) .

albobasalissp. n. (Fig. 30, venation) - Venezuela.
aurantiothorax(Alexander),as Longurio-Tanypremna(Fig. 31, venation)Peru.
bezzianus(Alexander),as TipulaPeru.
borgmeieri (Alexander),as Tanypremna
Brazil.
(Alexander),calliope
as Tanypremna- Brazil.

Studia Ent., Vol. 12, fasc. 14, novembro1969

189

carbonipes(Alexander),as Tanypremna Ecuador.
cerritus (Alexander),as Tanypremna Brazil
clotho (Alexander),as Tanypremna Brazil.
elegantior (Alexander),as Tanypremna
Colombia.
fieldianus(Alexander),as Longurio-Tanypremna Panama.
fuscitarsis(Alexander),as Tanypremna Colombia.
hodgei (Alexander),as Tanypremna- LesserAntilles: Dominica.
horridus (Alexander),as Tanypremna Peru.
incompletus(Alexander),as Tanypremna Venezuela.
Brazil.
invaripes (Alexander), as Tanypremna
kadeni (Alexander),as Tanypremna- Venezuela.
<<SouthAmerica>.
longipes.(Fabricius),as Tipula
longissimus(Enderlein),as Stegasmonotus Brazll.
manicatus(Osten Sacken),?s Tanypremna Brazil.
mirandus(Alexander),as Tanypremna- Venezuela.
opilio (Osten Sacken), as Tanypremna-

Guatemala,Panama,Venezuela.

Brazil.
ornatipes (Alexander),as Tanypremna
perornatus(Alexander),as Tanypremna Ecuador,Peru, Venezuela.
phylax (Alexander), as Longurio-Tanypremna(Fig. 32, venation)Peru.
picturellus(Alexander),as Tanypremna- Ecuador.
porterianusporterianus(Alexander),as Tanypremna Brazil.
p. limai (Alexander),as Tonypremna Brazil.
proavitus (Alexander),as Tanypremna Brazil.
salome (Alexander),as Tanypremna Brazil
saltatrix (Alexander),as Tanypremna Costa Rica.
(Alexander),as Tanypremna Brazil.
sonctaecatharinae(Fig. 33, venation)
uniguttatus(Alexander),as Longurio-Tanypremna
Peru.
Subgenus TanypremnellaAlexander
T a n y p r e m n e l l a A l e x a n d e r ; A n r ! . S " s . N a t . - H i s t . , _- ( J l ) l : 3 3 9 - 3 4 0 , f i g . 2 ; l g 3 g ; a s
Tanypremna (Tanypremnella); type cryssot|no (Alexander); original"designatiori.

antenniferus(Alexancler),as Tanypremna (Tanypremnella)- peru.
cry'st,""inus(Alexander), as Tanypremno (Tanypremnella)
Ecuador.
gentilis (Alexander), as Tanypremna.(Tanypremnella)
Venezuela.
maldonadoi(Alexander), as Tanypremna (Tanypremnella)- Venezuela.
mediocornis(Alexander), as Tanypremna (Tanypremnella)
Ecuador.
megacerus(Alexander), as Tanypremna (Tanypremnella)
Ecuador.
(Alexander),microcerus
as Tanypremna (Tanypremnella)
Ecuador.
perdistinr:/us(Alexander), as Tanypremna (Tanypremnella)Peru.
segnipes(Alexander), as Tanypremna(Tanypremnella) (Fig. 34, venation)
Bolivia.
transfasciatus(Alexander), as Tanypremna (Tanypremnella)
Ecuador.
SubgenusTanyprentnodesAlexander
Tanypremnod.es Ale.xander; Ent. News, 35: 290-291; 1921; as Tanypremna (Tanyp r e m n o d e s ) ; t y p e l e u c o p l a c a ( A l e x a n d e r ); m o n o t y p i c a n d o r i g i n a l d 6 s i g n a t i o n .

Ieucoplacus(Alexander),as Tanypremno(Tanypremnodes)- Brazil.
subapicalis(Alexander),as Tanypremna(Tanypremnodes)- Brazil.

r90

A I e x a n d e r, TropicalAmericanTipulidae(VIII)
SubgenusXenotipulaAlexander

Xenotipula Alexander; Ann. Mag.Nat.
Hist., (9) 8: l7tt92l;( A l e x a n d e r ) ,E t h i o p i a n , m o n o t y p i c a n d o r i g i n a l d e s i g n a t i o n ) .

(type munroi

cisatlanticus(Alexander),as Xenotipula (FiS. 35, venation)-

Brazil.

Leptotarsus(Longurio) borgmeieranus
sp. n.
Allied to exemptus; mesonotumpolished dark brown, pleura andpleurotergite light yello'w with a transversebrown stripe on mesepisternum; femora yellow, tips brown; wings pale brown, stigma and costal
border dark brown, prearcular field and base of cell R not darkened;
abdominaltergites conspicuouslypatternedwith black and yellow; malehypopygiumwith inner dististyle slightly decurvedat tip, spinoid setae
concentratedon a low lobe.

Male.
Length,about 8-8.5 mm.; wing, 9.5-l0 mm.;
antenna,about ll.5-12 mm. Frontalprolongationof headyellow,
without nasus;palpi light brown, terminalsegmentdarker.Antennaeof male very long, exceedingthe wing; proximal threeyellow,outersegments
segments
brownishblackto black; f lagellar
with long erect white setaeover
segmentsvery long-cylindrical,
the whole length.Head orangeyellow.
Pronotumtestaceous
yellow.Mesonotum
polisheddark brown,
scutellumpaler brown, parascutellapale yellow. pleura andpleurotergite
light yellow, includingthe dorsopleural
membrane,rnesepisternum
with a transverse
darkenedband. Halteresyellow,
knob dark brown. Legs with coxaeand trochanterslight yellow;
fentoraand tibiae yellow, tips brown, darker and more extensive
on femora;tarsi brown.Wings pale brown,cellsC and Sc, withthe stigma,dark brown; prearcularf ield and baseof cell R not
darkened,as in exemptus;two conspicuouspale yellow areas,
one near outer end of cell M, the secondon basal half of cells
Rs and Rr,; veins brown, darker in costalregion.
Abdominaltergitesconspicuously
patternedwith black andyellow, the latter on sides of basal half of segments,leaving
black -shapedareas; sterniteschiefly yellow, posteriorbordersof interntediate
segntentsfaintly darkened;a narrow subterminal
ring, hypopygiumyellow. Male hypopygium(Fig. 38) with thetergite,t, slightly narrowedoutwardly,posteriorborder shallowly
emarginate,with morethan the outerhalf of platewith abundantsetae.Dististyles,d, terminal, outer style broad, apex obtuse;inner style with apex slightly decurved,obtuse;spinoid setae
chiefly in a compactgroup on a low lobe, with a few others

Studia Ent., vol. 12, fasc. 14, novembro1969l9lidioneura ( Alerander ) ; venation. - _fig. 29. . L.eptoFig. 28. Leptotarsus (Ceoneura)'Fig. 3O.._Leptofcrsus(Lihoniotles)
sulphurelia (Alexander);taisus'- venation. :(Tany.premna)
Leptotarsus(Tanyprimna)
3t.
Fig.
n.;
venation.
atbo'basalii rp.
'venation.
Fig. 32. Leptotarsus (TanypremryQ ph.ylax
aur:aiintnorax (Alexander);'Fig.
(Alexander); venation. 3:1. Leptotarsis (Tanypiemna) uniguttatus (Alexander);r)enation.
F i g . S 5 . L t ' p t o t a r s u s ( X e n o t i p u t a ) c i s a t l a n t i c u s -( A l e x a n d e r ) ; v e n a t i o n .

on disk of style. Leptotarsus (Longurio) exemptus ( Fig. 39)
differs chiefly in the conformation of the inner dististyle,asshown.
Habitat: Brazil.
Holotype,6 , Petropolis,Rio de Janeiro,altitude2,7OOfeet,
April 13, 1940 (Thomas Borgmeier).Paratopotype,6 .

r92

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

The speciesis dedicatedto Father Thomas Borgmeier,distinguishedentomologist
and world authorityon variousdifficult familiesof insects.
All scientistswho study the vast insect fauna of the Neotropicsareindebtedto Father Borgmeieras being the founderand directoi of the
Revistade Entomologiaand the Studia Entomologica.The most similar
speciesis Leptotarsus(Longurio)exemptus(Alexander),differingespecially in the wing pattern and in the structure of the hypopigium,asdiscussedabove.

Leptotarsus(Longurio) helota sp. n.
Size small (wing of male I mm.); general coloration of mesonotum
mediumbrown; head velvety black, the lo'w vertical tuberclelight orange;
antennaeshort; wings strongly infuscated,vein Rr-, preserved;abdominal-ro;segments bicolored. the sternites more strongly
male hypopygiumwith tergal emargination shallow; inner clististylewith spinoid setae
elongate, naillike.

M a I eLength,about 12 mm.; wing, g mm.; antenna,
about l.l n1nr.Frontal prolongationof head light yellow; palpi
black.Antennaeshort;scapebrownishyellow,remainderof organ
black; f irst f lagellar segmentoval, secondsmaller,remaining
segmentssubcylindrical,
shorterthan their verticils.Head velvety
black, the low vertical tubercleabrupily light orange.
Pronotuntorange,centerof scutelluminfuscated.
Mesonotum
mediumbrown,scutellunr
and mediotergite
very sparselypruinose.
Pleura light brown, anepisternum
slightly clarker.Halteresinfuscated.
Legswith coxaetight brown; trochantcrs
yellow;femora
brownishyellow, cleareryellow basally,tips blackened;tibiae
brown, tips narrowly black, tarsi brownishblack; claws toothed.
Wings strongly infuscated,costal border and tfte snlall oval
stigma slightly darker brown; veins light brown. Veins posterior
to R glabrous.venation: Rt-z preserved,less stronglyupcurved
than in styx; petiole of cell ML slighfly longer than m; m-cu
shortlybefore f ork of Ms-1; cell 2nd A narrowerthan in styx.
Abdomen elongate;tergites vaguely bicolored,black withnarrow light brown areas on posterior half of basal rings ofintermediatesegments,basal tergite brownish yellow; sternites
yellow, conspicuouslyringed with black on basal rings; segments seven through nine black. Male hypopygiunr( Fig. 40)
with posteriorborder of tergite,t, shallowlyemarginate,vestiture
relativelyshort. Outer dististyle,d, narrowedat outer end; innerstyle with body oval, provided outwardly with erect blackenecl
spinoid setae,beak of style decurved,its margin with a row
of strong blackenedspines.

StucliaEnt., vol. 12, fasc. l-4, novembro 1969

r33

39

(Longurio) huanucensis(Alexander); venation- -- F!g' 37.
Fig. 36. Leptotarsus'(ton,gurto)'
Le-ptotars.us.
38.
venatiort.
(Alexander);.
ses"soris
t-ipton|'su{^; _p'i8:(,Longurio.)
Leptotarsus
njg.
t
mhle
n.;
sp.
OoremZt.erarius-Iig..
vpopvsirlm.
.99.
7i:oie;;{;)'(niexanOer);
helotaLe.oto,tapys,-(Longurio).
hyboprgiud,'mal'e ---'Sfmbols:bii;'iiu;---10:b, b-sistyle;d, dististyles;f, tergite.
rtr- n)j mate hybbpygium.

Habitat: Brazil.
Holotype, 6 , Borac6a, S5o Paulo, altitude 850 meters'
January30, 1949 (Lauro Travassos).
The only other regional speciesof the subgenusLongurio having
the spinoid ietae of tha inner dististyle of the hypopygiumelongate as
in th; present fly is Leptotarsus(l,ongurio) styx (Alexander), likewise
from southeastern Brazi\. This differs- in the coloration of the head,thorax, abdomenand legs, in the trichiation of the wing veins, and in
hypopygialdetails,particuiarly the tergite and inner dististyle
l3

194

A l e x a n d e r , T r r p i c a l A m e r i c a nT i p u l i d a e( v l l l )Leptotarsus(Tanypremna)albobasalissp. n.
S i z e r e l a t i v e l yl a r g e ( w i n g o f f e m a l e l 8 m m . ) ; m e s o n g t u mp o t i s h e dyelkrw, conspicuouslypatterneclwith dark brown, pleura with transverse
dark brown.stripes; palpi brown, terminal segment whitened; femora
light brown, tips blacliened;tibiae brownishblack, base narrowly whitened,tarsi white; wing.sweakiy brownish yellow, patterneclwith darker brown,
including especiallythe stigma anrl a confluent cloucl over the anterior
ctlrcl; R.s more than one-half Rr-,r, base of cell R^ narrowed, cell /sf'iletioleM, large,
rtblique,Ionger tha-nof cell ill.,";abtlgminaltergites
ry
ctlnspicuously
patterneclwith brown and yellow, sternitesalmost unifoimly
yellow.

F c n r a I e . - L e n g t h ,a b o u t2 g m r n . ;w i n g , I t t m m . ;antenna,about I .5 nrnr. Frontal proloneationof head brown;
palpi much
I0nger than the antennae,proxinrar three segments
brown, theelongate terntinal segrnentwhitened. Antennaewith
scape andpedicel yellowed, flagellurn brown, the elongate
terminal segtttent paler. Head with front attd anterior vertex paler
than thebrown posterior parts.
Pronotulttbrownish black, the color cxtendeclventrad
overthe propleuraas a narrow bancl that includesthe bases ()fthefore coxae.Mesonotalpraescutunrpolishedvellow, conspicuously
patternedwith dark brown, includingthe anteriorend andlateralbrlrderbehindthe hunteri,the latter continuedventradas anarrowtransversestripe that includesthe posteriorhalf of the mesepisternuln' more intenselybrownish black on the dorsopleuralmembrane, slightly suffusing the anterior face of the nriddle coxa;
posterior sclerites of notunr chiefly light brown, parascutella
yellow, pleurotergitewith a conspicuousdark brown area.pleura
yellow, transverselypatternedwith brown, as describedab've.
Haltereselongate,ste.n.tgreenishwhite, knob clarkenecl.
Legs withcoxae yellowed, darkened as clescribecl;trochanters greenish
yellow; femora tight brown, nlore yelloweclbasally,tips -onspicutlusly black; tibiae brownish black, bases narrowly whitened,slightly less than the darkenedfenroralapex; tarsi w5ite, ternrinal
segntenta little darker. Wings ( Fig. 30) weakly Srownishyellow, the broad cell C clearer yellow; stierna brown, anterior
cord with a slightly nlore extensivedarker hrown area; outerends of cells Rz and R,r pale brown; veins beyoncl cord very
narrowly and inconspicuously
seamedwith brgwn, veins Cu andnt-ctt similarly darkened;veins brown, costa tingeclwith green,especiallyoutwardly.Macrotrichiaof outer veins relativelysparse,
lacking on veins behinclM z. Venation: veins sc,r, f ree tip of

Studra Ent., vol. 12, fasc. l-4, novembro 1969r95

Scr and R,-, about equallyspacedon costa; Rs nlore than onehalf Rr-r; base of cell prr narrowed,r-m very short; cell Ist M2
large, m oblique,longer tlran the petioleof cell M t.
Abdominal tergites medium brown, each with a transverse
yellow ring beyondbase,followed by a darker area that narrowly
includesthe lateraland posteriorbordersof the segrnents;sternites
almost uniformly yellow, proximal segmentsweakly more darkened before apices. Ovipositor with both cerci and hypovalvae
long and straight.
H abitat: Venezuela.
Holotype, g, Choroni, Aragua, R. Maracay, altitude 980
nretersD, e c e m b e r2 2 , 1 9 3 8 ( A n d u z e ) .
The most similar speciesis the larger Leptotarsus ('l'anypremna)ntiranda (Alexancler),likewise f rom Venezuela,the two f lies differing
evidently in the rletails of colrlrationof the bocly, legs ancl wings, andin the venation.L. (f.) kadeni (Alexander) also is related but is more
distinct in coloration ancl venation.

MegistoceraWiedemanni l l a t k i s t o t t r u W i e d e m a n n; D i p t . e x o t i c i r , p a r t | : . l l ; I t t 2 0 .
,lltgistocero Wiedemann; Aussererrr. zs'eill. Ins.. I: 515; ltt2t{: (t1'pe Tipula litipts_ Fabricius; designated by Ilacquart. lftiJfi: ti:J); Ethiopian.
T h e p r i o r n a m e I l l a e k i s t o c e r a\ \ ' a s : rr l l i t r : r r i l v r r i c c t e d h v i t s ; rr r l h o r : r n t l t l t c l a t e r o n e
proposed and is norn'generalll' ;rcccptcd.

Two recent ntentberstlf tlre genus art: knorvn, incluclingttre
Old World Megistocera filipes, rcpresented by two geographicalraces or clines, the typical filipes being wide-spreadin Africa
whife the racef uscanaWiedemannhas a vast range in Australasia
and South Asia, occurring as f ar west as southern India. In
this species the antennaeof the male are greatly lengthened
whereasin the New World species,here discussed,the antennae
are very short in both sexes,with only eight segments( Fig. 5)
As was indicatedunder the genus Brachyprenlne, it appearsthat
the Baltic Amber (Upper Eocene)MacromostixbornhurdliMeunieris more properlyplaceclin the presentgenusand enrphasizes
theatfinities betweenthe two genera.
In Megistoceraand to a lesscrdegreein Brachylrremno,thewings ( Fig. 2) have veins Sc, ancl Sc, closelyapproximatedat
costa,closingcell Scr or virtually so. A peculiarityof the venationis the strongly angulatedvein Rr-,r which in some cases has ashort spur at the angle, lying along the proxinral edge of thestigma. A second venational f eature is the course of vein M 4
at its base, where it is directeclstrongly backward and thence
extendedto the nrargin at a slightly acute angle, with m-cu at
l3r

t96

A I e x a n d e r, TropicalAmericanTipulidae(VIII)

the point of angulation.Both speciesof Megistocerohave thecuter ends of veins Mr, Mz and M 4 pale to subatrophied,acondition found also in Brachypremno,and emphasizingthe
relationshipsbetweenthe two groups. Cell 2nd A of the wings
is very narrow in the local species,much broaderin f ilipes.The
legs in both sexesare excessivelylong and slender,especiallythe middle and hind pairs where the basitarsialone are nearly
twice the remainderof the tarsi and equal to or longer than
the wing.
The habits of the adult f lies and the immaturestageshave
in an outstandingpaper by J. S. Rogers(Univ.
been discussed
Michigan,Occas.Pap. Mus. Zoology,No. 521: l-14, 2 pls. with
7 figs. 1949).The remarkablef light of the adults,as describedin this report, has been observedin a somewhatsimilar mannerin Megistocerafilipes fuscanain New Guineaby Dr. JeanLaffoon( in litt.) . The immature stages of M. longipennisare aquatic,
occurringclose to the surfacefilnr in quiet water bodies,thelarvaeliving in detachedsectionsof- hollow plant stemsfloating
the larval casesof certain
at the surface,in this habit suggestinglarger stems
Trichoptera.As larval growth proceeds,successively
are utilized and pupationoccursin one end of the f inal tube.
List of Species
Iongipennis(Macquart),as Tipula (Fig. 2, venation;,Fig.5, 6 antenna).
Syn. tenuis (van der Wulp), as Tipula - Surinam.
United States,south to Argentinaand Bolivia; Creater
Southeastern
Antilles.

NephrotomaAlcigen
N e p h r o t o m a M e i g e n ; M a g . f i i r I n s e k t e n k u n d e 2, : 2 6 2 ; 1 8 0 3 ; ( t l ' p e d o r s a l i s ( F a b r i c i u s ) ;P a l a e a r c t i c ,m o n o t y p i c ) .
P a l e s M e i g e n ; N o u v . C l a s s . M o u c h e sd d e u x a i l e s . p .
1 8 0 0 ( n a m e s u p p r e s s e db y
I n t e r n a t . C o m m . Z o o l . N o m e n c l . ,t 9 6 3 : 3 3 9 ) .
Pachyrhina Macquart, Hist. nat. des Insectes, Dipttres, p. 88; 1834: (type crocata
L i n n a e u s , d e s i g n a t e db y W e s t w o o d , 1 8 4 0 : 1 2 8 ) ; e m e n d e ds p e l l i n g s i n c l u d e P a c h y r i n a ,Pachyrhyna, Pachyrrhina.

Nephrotomais a major genuswith nunlerousspeciesoccurring chieflythroughoutthe Holarcticand Ethiopianregions,withrelativelyfew recordedf rom the Neotropicsto the presenttime,
It seemscertain that , many additional specieswill be found;including several presently being studied by Dr. George W.
Byers.

Studia Ent., vol. 12, fasc. l-4, novembro1969

197

Importantrecent papers on the taxonomyand biology ofthe genus include the f ollowing.
A l e x a n d e r , C . P . , S o u t hA f r i c a nA n i m a lL i f e , l 0 : 265-281,2 figs.;
1964 (Lund)
SouthernEthiopian.
- RuwenzoriExpedition,1934-35,l, no. 7: Tipulidae, 1 3 5 - 1 6 4 , 3 1
figs.;(London)1956
Tropical Ethiopian.
B r i n d I e, Allan, Trans. Soc. British Ent., 14: 63-114,179 figs. European,
Immature stages.
D i e t z, W. G., Trans. Amer. Ent. Soc., 44: 105-140,4 pls. (with 34
f i g s . ) ; 1 9 1 8 -. N e a r c t i c .
Nearctic.
D o a n e, R. W., Ent. News, 19: 173-179;1908.
M a n n h e i m s , B e r n h a r d ,I n E r w i n L i n d n e r ,D i e F l i e g e nd e r P a l a e arktischen Region. Lief. 167: 32-59, figs. 10-26, pl. VI; 1951.
Western Palaearctic.
T h e o w a I d, Br., Die Entwicklungs-stadiender Tipuliden (Diptera,
Arten, pp. vii-xi,
der West-Palaearktischen
Nematocera)insbesondere
Immature stages.
195-308,332 figs.; 1957.
- Bestimnungsbticher
zur BodenfaunaEuropas,Lief. 7, Tipulidae, 1-100,
Immaturestagesof Europeanspecies.
344 figs.; 1967.

The immaturestages are f ound in soil, commonlybeneatha layer of leaf mold. Certain speciesare of some economic
irnportance,particularly to agriculture ( Brindle, Theowald referencesabove).
List of Species'tipulaMexico (identiy uncertain).
alfinis (Bellardi), as
Colombia.
alleni (Alexander),as Pachyrhinu
Ecuaclor.
aurocomatasp. n.
Bolivia.
Alexander
boliviana
cacuminisAlexander- Ecuaclor.
circumscripta(Loew), as Pachyrrhina- Cuba, Jamaica.
consularisconsularis(Osten Sacken), as Pachyrrhina.- Costa Rica, Nicaragua;Argentina,Brazil, Paraguay.
c. eminensAlexander- Venezuela.
dampfi Alexancler- Mexico.
Alexander(Fig. 44, 3 hypopygium)- Mexico.
durangensf.s(identity uncertain).
America>>elegans(Fabricius),as Tiputa - <<Southelegantula(williston), as Pachl,rrhina- LesserAntilles: st. Vincent.
North America, south
ferrugineaferruginea (Fabricius), as'tipula
to Mexico.
Syn.proxima (Bellardi),as Tipula,'auct.OstenSacken.- Mexico.
quadrilineata(Macquart), as Pachyrhina,auct. Osten Sacken.- Mexico.
glabricristataAlexander. Cuba, Haiti; Dominica.
globosa sp. n.
Colombia.
Alexande
Iateropolita
mediotigulaAlexander- Colombia,Venezuela.
Alexander(Fig. 41, venation). Bolivia.
melanoxantha
mexicano(Macquart),as Pachyrhina- Mexico (identity uncertain).
Costa Rica, Mexico.
nigrolutea (Bellardi), as Tipula

198

A l e x a n c le r , T r o p i c a lA m e r i c a nT i p u l i d a e( V I I I )nigropilosa sp. n.
Peru.
ordinaria (Osten Sacken), as Pachyrrhina - Mexico.
pulchella (Rondani), as Pachyrhina
Brazil.
punctifrons (Macquart), as Pachyrhina
Argentina, Brazil.
sparsicomasp. n.
Ecuador.
triobtusa sp. n.
Brazil.
usta (Osten Sacken), as Pachyrrhina
Costa Rica.
Doubtful species: elongata (Macquart), as Pachyrhina.
unknown, possibly Neotropical.

Country

Nephrotornaaurocomatasp. 11.
Generalcolorationyellow, conspicuously
patterneclwith black, inclutling the head ancl thorax; antennaeentirely black; wings fulvous brown,
cells C and Sc more yello'wecl;male hypopygiumwith the outer basal
lobe of the inner dististyleextendedinto a long slenclerspine; posteriorborder of eighth sternite conspicuously
emarginate,with a broad obtuse
central plate, the lateral lobes with abundant incurveclgolden yelklw
setae.

M a I e.
L e n g t h ,a b o u t 1 2 - 1 3 n u n . ; w i n g , l 0 . s - 1 2 m n r . ;antenna,about 3.8-4 mnl.
F e m a I e.
L e n g t h ,a b o u t 1 4 n t n l . ; w i n g , 1 4 m n l .
Frontal prolongation of head brownish yellow, narrowly
black above,includingthe nasus; palpi brownishblack, terminal
segment black. Antennae black throughout; flagellar segments
moderatelyincised,slightly exceedingthe longestverticils.Headorange, the elevated vertical tubercle more yellowed, incisedm e d i a l l y ;o c c i p i t a lb r a n d l a r g e a n d b r o a d , s h i n y n a c r e o u sa, n t e r i o r l y w i t h a d u l l b l a c k c e n t r a l c x t e n s i o nt h a t i s c o n t i g u o u swith large black lateralareas;hcaclu'ith corrspicuous
black setae,
those of the genac rrery long.
Pronotuntblack,scuturnrnecliallybroadly light yellow. Mesonotal praescutuntyellow with three polished black stripes, thernedianone reachingthe suture'behincl,latc'ralpair outcurvedto
margin,not opaque;scutumyellun',each lobe with two confluent
black areasthat are extendedacrossthe suture;scutellunrbrownish black, parascutellalight yellow, clarkencdlaterally; central
part of mediotergitebroadly blackened,expandedand slightly
paler behind, pleurotergiteblackened,the nlore elevatedkatapleurotergiteyellowed. Pleura btack, conspicuouslyvariegatedwith yellow, including the posterior anepisternumand pteropleurite,dorsal sternopleurite
and meron,and much of the metapleura. Halteresyellow. Legs with f ore coxae brownish black,
midcoxaeyellowedanteriorly,dark brown behind,posteriorcoxae
more uniformly yellow; trochantersdull orange; femora obscureSttrtlia Ent., vol. 12, fasc. l-4, novembro ltl69l t)9

yellow to brownish yellow, tibiae passing into brown; tarsi
black, claws of male toothed.Wings fulvous brown, cells C andSc, with vein Cu, more yellowed, base of cell 2nd A slightly
more infuscated; stigma light brown, with macrotrichia.Very
sparsetrichia in outer end of cell Rs. Venation: Cell M 1 sessile
to short-petiolate;m-clt at or close to fork of M4.
Abdomenyellowed,posteriorborders of tergites pale brown,
the amount increasingon outer segments,in male including theouter f our segnrents;genital seglnentof female yellowed.Maleouter margin
hypopygium(Fig. 42) with the tergite,t, transverse,rounded, posterior border with a circular emargination, theconspicuouslobes with blackenedspicules,including two or threeof snrallerspinoid
larger marginalpoints and with a concentration
setae at mesal end of lobe. Outer dististyle,d, broadest before

rnidlength,
narrowedto the obtusetip; inner stylewith beak pale,narrowerin the holotype,tip obtuse,lower beak blackened;outerbasal lobe yellow basally,extendedinto a long slenderblackspine. Eighth sternite,S s, large, sheathing,posterior borderwith a broad obtusecentralplate, theentarginate,conspicuously
larse lateral lobes with abundantlong incurvedgolden yellow
brushes.
setaeto fornt conspicuous
Habitat: Ecuaclor.
Holotype,6,, Banos,Tungurahua,altitude 2,20Ometers,
October15, 1938 (Macintyre).Allotopotype,9 , rvith type.Paraaltitude1,500
types,| 6, | 9, Upper Rio Topo, Tungurahua,
meters,Februaryt 949 (Velastegui); 2 6 6, Yunguilla,,NapoPastaza,altitude 1,700 meters,May 1944 (Velastegui).
especiallythe wing coloration,the present
In its generalappearance,
fly is most similar to Nephrotoma usta (Osten Sacken), of Central
A m e r i c a ,d i f f e r i n g i n m a n y d e t a i l s o f c o l o r a t i o n .T h e m a l e h y p o p y g i u mof usto is nclt available to me for comparisonand was not describedby Osten Sacketr.

Nephrotomacacuminis Alexander
Nephrotoma cacuntinis Alexander; Ann. tYtag. Nat. Hist., (l I )12: 753-755; 1945.

Ceneral coloration of thorax yellow, the praescutunrwith
three black stripes, the polished.lateral pair extended to themargin in f ront and again at the suture to enclose a small
lateralyellow ground area,scutellumblack, postnotunland pleura20c

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

yellow, patternedwith brownishblack. Wings whitened,stigma
brown, with about 20 trichia. Abdomenyellow, basal segments
restrictedlypatternedwith brown, outer segmentsextensively
brownishblack to form a broad ring. Male hypopygium( Fig.
43) with outer basal lobe of inner dististyle,d, very short,terminatingin an equally short blackenedspine, dorsal crest
very low. Eighth sternite,8s, with a medianligula that is surroundedby long yellow setae.
The types were f rom Morro Morro, El Oro, Ecuador,colBolectedby David Laddey.A f urther male f ronr Balzapamba,
livar, altitude700 meters,takenJune4, 1938,by Macintyre.
Nephrotomaglobosa sp. n.
Antennaeblack,proximal segmentsslightly paler; frontal prolongationof head light yellow, broaclly black above; posterior part of head
conspicuouslypatternedwith black; mesonotalpraescutumlight yellow
with three polished black stripes, outer pair outcurved,remainder ofpatternedwith black; wings brownnotum and the pleura conspicuously
ish yellow, prearcularand costal fields clearer orange yellow; abdomenyellow, conspicuouslyringecl rvith black, including the broad apices of
the tergites ancl narrower margins of sternites,eighth tergite yellow;
male hypopygiumwith beak of inner clististylevery short and ohtuse,outer basal lobe a small roundeclblack knob; eighth sternitewith posteriorborder shallowly emarginate,with abundant short yellow setae.

M a I e.
Length, about 13.5 mm.; wing, 10.5 mm.; antenna, about 3 mm. Frontal prolongationof head light yellow,
broadly black above,includingthe nasus; palpi brownishblack,terminalsegmentblack. Antennaewith proximal three segrnents
llrownish black, remainderblack; segrnentsshort, the basal enIargementssubequalto the renrainderof segrnent.Head yellow,
the globularverticaltuberclelight yellow; occipitalbrand polishednacreous,borderedby velvety lllack, sending a point cephalad
onto the tuberclel antcrior vertex with large dull black areasadjoining the eye, alnrostconfluentwith the vertical darkening,
postcriorparts of head blackened.
Pronotum medially light yellow, sides broadly blackened.
Atesonotalpraescutum light yellorv with three polished black
stripcs, the lateral pair with very narrow blackenedmargins, atouter end with a large opaque black spot to appear outcurved;
further blackeningsalong suture and at lateral border completely
enclosea triangularground areabeforethe suture;scutumyellow,
lobes extensivelyblackencd;scutellumblack, parascutellayellow,

Studia Ent., vol. 12, fasc. 14, novembro1969

201

t+3

{rT

F i g . 1 2 .. N e p h r o t o . m a; venation.
F i g . 4 1 . N e p h r o t o m u m e l a n o x a n t h a A l e x a n d e r-13.
cacuminis Alexander;
licphrotoma.
lti,q.
hypopygiu_nr.
mate
n.;
sp.
au-r'ocomata'.
n t g . a A . ' N i p n r o t o m u , i u r a n g e n s i s. A l e x a n d e r ; m a l g h - y R o p v S i u m .
m;it-nypopygium.
Fig' 46. Nephrotomaboopsa--'Fig."'ii.''Neinrotoma- g- ht oyp
y gl pi i r mn. . ; m j l l e h y p o p y g i t r . m .
s . p a r s i c o m a. s p . n . ;
N
e
p
h
r
o
t
o
m
a
F
i
g
.
.
m
a
t
e
n
.
;
s
p
.
.
i
4
.
'
.
V
e
p
l
t
r
o
t
o
m
a
nisropilosa
.47.
n
.
;
s
p
.
t
r
i
o
b
.
t
u
s
a
fale lypopygium.
; ; i ; ' ' ' r ; "; ;J;rp- fr' [ "i us m
. ' 'u- o t s : n ] e .
"imd , d - i s t i s t y l e sg; , g o n a p o p t r y s i ss; , s t e r n i t e ; f , t e r g i t e .

202

A I e x a n d e r , T r o p i c a lA m e r i c a nT i p u l i c l a e( V I I I )narrowly blackenedposteriorly;mediotergiteblack, with a large
yellow area at each anteriorpart, pleurotergiteblack, variegatedwith yellow on anterior katapleurotergite.
Pleura light yellow,
conspicuously
variegatedwith black on propleura,anepisternunl,
ventral sternopleurite,dorsal and ventral pteropleurite,meronand on part of nletapleura.
Haltereswith stenrdark brown, knob
light yellow. Legs with coxaeyellow, narrowlyinfuscatedbasally,
more extensivelyso on f ore legs; trochantersorangeyellow,withblackenedpattern;fenroramediumbrown, tibiae slightly darker,
tarsi black, claws of nraletoothed.Wings with ground brownish
yellow,clearerorangeyellow in prearcularand costalfields,cells
beyondcord slightly more darkened,especiallyat tips; stigma
brown, with about I 5 trichia; veins light brown on basal halfof wing, beyond cord brownish black. Venation: nt-clt at forkof M; cell Mt very short-petiolate.
ringed with black, including
Abdomenyellow,conspicuously
the broad apices of the tergites and narrower margins of thesternites;eighth tergite yellow, eighth sternite brownish black,remainderof hypopygiumvariegatedblack and yellow. Male
hypopygium(Fig. 45) with posteriorborder of tergite,t, deeply
emarginateto f ornr two rounded lobes, each with relatively
nunlerousblackenedspiculesarrangedchiefly in two transverse
rows. Outer dististyle,d, narrowedon outer f ourth; inner styleshort and compact,beak very short, obtuse; dorsal crest high,glabrous;outer basal lobe a small roundedblack knob. Gonapoouter half dilated. Eighth sternite,
physis boonrerang-shaped,
8 s , n a r r o w e do u t w a r d l y , p o s t e r i obr o r d e ' rs h a l l o w l ye m a r g i n a t e ,with abundantshort yellow setae.
Habitat: Colonrbia.
23,
altitude3,500nleters,Decentber
Holotype,6, Manizales,

1958 (JuanFoerster).
Nephrotoma globosa is generally similar t o N. consularis (Osten
Sacken), differing in details of coloration of t h e antennae.thclrax andabdomen,ancl in hypopygial structure.

StucliaEnt., vol. 12, fasc. 14, novembro1969

203

Nephrotomanigropilosasp. n.
General coloration of thorax yellow, praescutumwith three polishedbrownish black stripes, pleura with more reddish brown areas; antennae
black, scape light yellow; wings yellowed basally, less distinctly sobeyond cord; abdomen yellow, posterior borders of segmentsbrownish
black, with abundantrelativelyshort black setae;eighth sternitedarkened,remainderof hypopygiumyellow; beak of inner dististyleobtuse,outerbasal lobe subquadrate,blackened;eighth sternite with posteriorborderemarginate,with abundantgolden yellow setae.

M a I e . - L e n g t h ,a b o u t 1 3 m m . ; w i n g , 1 0 . 8m m . ; a n t e n n a ,about 2.7 mm.
L e n g t h ,a b o t t t 1 5 t n l n . ; w i n g , 1 2 n l n l .
F e m a I e.
Frontal prolongationof head light yelkrw, including thenasus, the latter tufted with long black setae; palpi yellow,
terminalsegmentblack. Antennaeof male sltort; scapelight yelonly moderatelyincised.
low, remainderblack; tlagellarsegrnents
Head with anterior vertex light yellow, vertical tubercleentire;a small black spot adjoiningeye; occipitalbrand broad, polished
black; posteriorvertex with abundantblack setae.
Pronotumlight yellow, sidcs narrowly blackened.Mesonotal
praescutunr
yellow, with threc'polishe'dbrownish black stripes,
lateral pair outcurvedto nrargin.as a nlore intenselyblackened
spot; lateralend of sutureand posteriorlateralborder narrowly
black; scutunrlight yellow,each lobe with two confluent brorvnish black areas; scutellumbrownish black, parascutellalightyellow with a centralbrownishblack stripe
yellow; mediotergitethat is widenedposteriorly;pleurotergitebrown, the katapleurotergite yellow. Pleura yellow with reddish brown areas, mostevident on ventral sternopleuriteand meron; propleura yellow,
Halteresdark brown, base of stem obscureorange, knob clear
orange.Legs with coxae and trochantersyellow; femora yellow,
more darkenedapically;tibiae brownishblack,tarsi black; claws
of maletoothed.Wings yellowedbasally,lessdistinctlyso beyond
cord, veins black, thosebefore cord yellow; stignrapale brown,with about l5 trichia. Venation: Cell M t narrowly sessile.
Abdomen yellow, posterior borders of tcrgites brownish
black, the cotor in part produced by very numerousrelatively
short black setae;eighth sternitemore uniformly darkened,hypopygium yellow. In the female the blackenedsetae restrictedto
segmentstwo to f ive, beyond this with normal longer yellow
setae.Ovipositorwith tips of cerci narrowly obtuse.Male hypopygium ( Fig. 46) with blackenedspinoid setae of tergal lobes,

204

A I e x a n d e r, Tropical AmericanTipulidae (Vlll)t, numerous.
Outer dististyle,d, with outer half narrowed;innerstyle with beak broadly obtuse,narrowly blackenedon lower
margin,lowerbeakbroadlyrounded,black;dorsalcrestmoderately
developed;
outerbasallobesubquadrate,
blackened.
Gonapophysis
slender,outer half slightly broader,tip obtuse.Eighth sternite,
8s, with posteriorborderemarginate,with abundantgoldenyellow
setae,median part of each lobe with very abundantshorteryellow setae;emargination
with membranedenselyprovidedwith
small circularareasthat bear microscopic
setae.
Habitat: Peru.
Holotype, 6 , Tingo Maria, April 5-8, 1963 (L. E . P e n a ) .
Allotopotype,9 , pinnedwith type. Paratopotype,one, I , pinned
beneathallotype.
Nephrotoma nigropilosa is generally similar to some other regional
species,being best distinguishedby the vestiture of the abdomen andin hypopygial structure, particularly the inner clististyle and eighthsternite.

Nephrotomasparsicomasp. n.
General coloration of thorax yellow, conspicuouslypatternedwith
black; praescutalstripes polished,the lateralsoutcurvedto margin; femora
yellow, tips narro'wly darkened; abdomen orange, posterior borders oftergites narrowly darkened;male hypopygiumwith eighth sternite withrelatively few short yellow setae.

M a I e.
L e n g t h ,a b o u t 1 3 m m . ; r v i n g , I I m m . ; a n t e n n a ,a b o u t3 . 2 m m .
F e m a I e.
L e n s t h ,a b o u t 1 3 . 5m n l . ; w i n g , 1 2 m m .
Frontal prolongation of head light yellol, above more
fulvous,includingthe conspicuous
nasusthat is fringed with long
black setae;proximalthreesegrnentsof palpi yellow,the terminal
one brown, passinginto black.Antennaewith scapeyellow, pedicel
brown, flagellumblack; flagellarsegmentsonly moderatelyincised,slightl-yexceedingthe longerst
verticils.Head light yellow; occipitalbrand and a small spot adjoining eye blackened.
Pronotum yellow, sides blackened.Mesonotal praescutum
yellow, with three polished black stripes, lateral pair straight
but with a more intenselyblackened area at its anterior end,rcaching the suture and contiguouswith a smaller similar spot
on dorsopleuralmembrane;scutumyellow, each lobe with a single
polished black area, anterior and lateral parts, including thesuture,more intenselyblack; scutellumblack, parascutellayellow;
nrediotergite
black, the broad anterolateralpart light yellow; kata-

StucliaEnt., vol. 72, fasc. l-1, novembro1969

205

black. Pleura yellow, vapleurotergiteyellow, anapleurotergite
riegatedwith black, includingthe propleura,ventralanepisternum
and sternopleurite,and more restrictedareas on anterior pteropleurite and ventral meron. Halteresbrownish yellow, base ofknob slightly more darkened.Legs with all coxaeand trochantersferruginousyellow; femorayellow, tips narrowly darkened,tibiae
brownishyellow,'darkeroutwardly; tarsi black, claws of male
toothed.Wings brownishyellow,prearcularand costalfields moresaturatedyellow,especiallycell Sc,'stigma mediumbrown, with
about 20 trichia; veinsbrown, Sc, R, and Cu, with the prearculartoveins, more yellowed.Venation: Cell M I very short-petiolatebarely sessile;m-cu just beforefork of Mn.
Abdomen orange, posterior borders of segmentsnarrowly
darkened,the actual marginsblack; secondtergitewith a, central
black spot on anterior half, sternitesmore uniformly yellow,
brownishblack,the bordersand central
eighthsterniteextensively
area yellowed, remainder of hypopygium yellow. Male hypopygium (Fig. 47) with lobes of tergite, t, low, rounded, thesmall blackenedspinulesarrangedin two grouPS,thoseof lateral
Inner dististyle,d, with beak relatively
area more concentrated.
slender,unblackencdexcept along lower ntargin adjoining theobtuselower beak; outer basal lobe small,obtuse,with a blackenedpoint, basewith f our or f ive long erect setae;gonapophysisa twistedyellowblade,near its baseu'ith a slenderspine.Eighthsternite,8s, large and sheathing,posteriorborder with a small
yellow setae.
incisionthat is f ringed with f ew inconspicuous
Habitat: Ecuador.
Holotype, 6 , Puyo, Napo-Pastaza, January 1949 (Velastegui).Allotopotype,I , pinned with type.
of the present fly is much as in Nephrotomatri'
The hypopygium.southeasternin the structure
Brazil, clifferingespecially.
obtusasp. n., ofof the inner clististyle,including the much more slender beak'

Nephrotomatriobtusa sp. n.
Ceneral coloration of praescutumyellow with three chestnut brown
stripes, the lateral pair outcurved; head dull orange, occipital brandlarge, margined with paler brown; legs obscure yellow; wings weakly
darkenecl, prearcular and costal fields brownish yellow, stigma pale
brtrwn; basal abdominaltergites bicolorecl,yello'w,apices brownish black;male hypopygium with beak of inner dististyle very obtuse, clorsal crest
low, glabrous, outer basal lobe small, obtuse; eighth sternite large,
narrowed outwardly, posterior border with a very small central pocketthat is provided with small yellow setae.

206

Alexancler,T r o p i c a lA m e r i c a nT i p u l i c l a (ev l l l )Length, about I I nlm.; wing, 8 nlm. Frontal
M a I e.
palpi
prolongation of head yellowed above, darker on sides;
pedicelobscure yellow, setae dark. Antennae with scape andyellowish brown; f lagellunrbroken. Head dull orange; occipital
paler
6rand large, dark brown, anterior end pointed, margins
brr.rwn,the color extendeclanteriorly to the vertical tubercle.
Pronotum yellow, darkened laterally. Mesonotal praescutum
broad
yetlow, with three chestnutbrown stripes,the median one
end morein front, lateral pair outcurvedto margin, the anterioralmostexpanded,darkei, polishecl;scutumyellow medially,lobes
yellowed;
uniformly brown; scutellumdark brown, parascutella
yellow,with a V-shapedpalebrown area;pleurotergite
nrediotergite
extensivelyyellowed' Pleura yellowed'
brown, t<atapleurotergite
meminconspicuouslypatterned with pale brown, dorsopleLlral
knob clearer
brane yellow. Haltereswith stem brownish yellow,
coxae
yellow. Legs with coxae and trochantersyellow, posterior
outer tarsal
more infuscated;remaindertlf legs obscureyellow,
and costal
segmentsdarker. wings weakly darkened,prearcularveins palef ields more brownish yellow, stigma pale brown;
Venation:brown. Stignrawith about seventrichia, all in cell Rr.
; m-ctt shortly before fork of M,'
Cell Mr narrowly sessilebrownAbdominal tergites bicolorecl,yellow basally, apices
on tlttter segntents
ish black, the anrounttlf the latter increasingsternitesand
where the bases likewise are narrowly darkened;
brown' Male
hypopygium more yellowed, eighth sternite light
posterior
hyp,rpygium ( Fig. 48) with the tergite, t, transverse,
the broad lateralborder with a very narrow nredianemargination,setae at either
lobes transverse,with small blackenedspinoid
short' the apex
cncl of the plate. Outer clististyle,d, relatively
be'ak'lower beaksucldenlynarroweclttl a ptlint; inner style with
tlf heak pale, theancl outer basal lobe all <lbtuseapically, apex
Eighth sternite'
rentainderclarkened;dorsal crest low, glabrous'
virtually entire'
8.S,large, narrowed outwardly, posterior borderrelativelyabundantwith a very small centralpocketprovidedwith
andsetae,'theother setaeof sternitelong
yellow, inconspicuousof plate'
black, very numerouson outer three-fourths
Habitat: BrazilHolotype, 6, Rio Negrtl, ParanA,Noventbc'r21, 1944 (M'Witte).
hypoThe speciesis most reaclily tolcl from regional allies by the.
especiallyth; inner clististyleand eighth sternite'
pygial charac.ters,

Stuclia Ent., vol. 12, fasc. l-4, nrtvembrol969

207

OzodiceraMacquartO z o _ d i c e r a/ f l a c q u a r t : H i s t . n a t . I n s e c t e s D i p t e r e s , I : 9 2 ; l t l 3 4 ; ( t y p e p e c t i n a t u(Wiedemann)a, s o c h r a c e a , \ { a c q u a r tm
; onotypic).
H e m i c t e i n oW e s t w o o d ; Z o o l . J o u r . , 5 : 4 5 0 ; t t t 3 5 ; ( t y p e g r u c i l i s W e s t w o o d ; m o n o t y p i c ) .

Ozodicera is one of the major Tipuline genera in Tropical
Anrericaand appearsto be unusuallydistinct from its nearest
relativesin Australia.The nrost distinctivecharactersare foundin the antennaewhich are branchedin both sexesbut to a lesser
dcgreein the female ( Figs. 5l , 52). In the male of the typicalsubgenusthere commonly is a single short branch on each off lagellarsegmentstwo through seven,whereasin the subgenus
Dihexaclonus
thereare two suchbranches.
Thereis somedeviationin this numberof branchedsegments,in schwarzmaierano
there
being only five segmentsso involvedwhile in mocrecanthaandneivai there are seven. Certain speciesthat are assigned toDihexaclonusshow considerablemodification in the shape andarrangementof the two branchesor spurs.Norntallyboth branches
arise closetogetheron the segment,with their basescontiguousor barely united. In some, as spilophoectand umbrifera, thebranchesare unequalwith the shorter projectionarising fromthe larger branch lleyondits base,in extensuwith the spur atnear one-thirdthe length, in strohmi beyond rnidlength,while
only by a smallertubercle
the branchis represented
in noctivagans
shortly before the apex. ln thc. extreme case it is ditficult toassign the speciesto either of the recognizedsubgenera.
The male hypopygiumof certain speciesare conspicuously
nrodified, providingstrong specific characters( Figs. 53-55) . The
but in somespeciesare
wings conrnronly
are almostunicolorous
and attractivelypatterned.Thr' venationis shown
conspicuously(Fig. 4e).

List of Species
Subgenus Ozodirera Macquartattenuata AlexanderBrazil.
himaculata Enderlein
Brazil.
bispinilera AlexanderBrazil.
correrella AlexanderBrazil.
-Alexander
caudifera
Brazil.
t:inereipennisAlexander (Fig. 52, antenna 6 )corrientesanaAtexander
Argentina.
cygniformis Alexander -_ Ecuador.
duidensis Alexander
Venezuela.
eliana AlexanderBrazil.

tsrazll.

208

A I e x a n d e r, TropicalAmericanTipulidae(VIII)

Brazil.
epicosmaAlexander
eurystyla Alexander(Fig. 49, venation) - Peru.
Brazil.
extensa AlexanderBrazilgracilis (Westwood), as H emicteina
Brazil.
griseipennisLoew<<SouthAmerica>>longimana (Fabricius), as Tipula
Ecuador.
multiermisAlexanderBrazil.
nigromarginataAlexander
British Guiana.
noctivagansAlexander(ochraceaMacquart- see pectinataWiedemann).
British Guiana.
pectinata(Wiedemann),as Tipula<<SouthAmerica>>.
ochraceaMacquartSyn.
phallacanthaAlexander- Peru.
Ecuador.
piatrix Alexander
Peru.
Alexander
placata-schwarzmaierana
Btazil.
Alexander
septemtrionisAlexander Mexico.
<SouthAmerica>.
simplex (Walker), as Ptilogyna
Venezuela.
striatipennisAlexanderBrazil.
strohmi AlexanderBrazilsubvittataAlexander
Peru.
thaumastaAlexander(Fig. 55, 6 hypopygium)
Ecuador.
trispiniferaAlexander
Btazil.
witteana AlexanderBrazil.
zikaniana Alexander

SubgenusDihexaclonusEnderlein
Dihexaclonus Enderlein; Zool. Jahrb., Syst., 32: 28; l9l2;
original designation).

(type apicalis jllacquart;

apicalis Macquart - Brazil.
biaculeataAlexander(Fig. 53, 6 hypopygium) Peru.
Brazil.
effecta AlexanderBrazil.
fumipennisLoew
graciliramaAlexander
Bolivia.
Alexander
Britishguianensis
Guiana.
jesseanaAlexanderBrazil.
Alexander
Brazil.
lanei
Venezuela.
longisectorAlexanderBrazil.
Alexander(Fig. 51, 3 antenna)
macracantha
neivai Alexander Brazil.
Panama.
panamensisAlexander
perfuga AlexanderBrazil.
Brazil.
pumila AlexanderPeru.
spilophaeaAlexanderBrazil.
superarmataAlexander
Colombia.
telestylasp. n.
Brazil.
terrifica AlexanderBrazil.
triguttata AlexanderBrazil.
tripallens AlexanderBrazil.
umbrifera AlexanderBrazil.
xanthostomaLoew

Studia Ent., vol. 12, fasc. l-4, novembro1969

20s

Dl,
55

54

(b

S g - . 1 - 9 ,O z o d i c e r a , ( O z o d i c e . r ae)u r y s t l ' / aA l e x a n d e r ;v e n a t i o n .- F i g . 5 0 . P h a c e l o d o c e r aFig. 51. Ozddicera (Dihexaclonus) m.acraiaitha
tlabellilera (Loew); venation._ Alexander; male antenna. _- F-lg. 52.- Ozodicera (Ozodice'ra) cinereipeinis
nteiinoer;'Alexander;
male antenna. =Fig.. 53. Ozodicela (Dihcxaclonus) biacuteatamaltihypopygiu_m._- Fig. 54. Ozodicera,(Dihexaclonus) teleityta sp. n.; male hypof
ygium.
'-Syin-Uots:- Fig. 55. Ozodicera (Ozo-dicera)thaumasta Alexlnder; male hypogiygium.
b , b a s i s t y l e ; d , d i s t i s t y l e s ; p , p h a l l o s o m e ;t , i i r g i i e - .

t4

2ro

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

Ozodicera(Dihexaclonus)telestylasp. n.
Ceneralcolorationof mesonotalpraescutumobscureyellow, with four
redtlishbrown stripes; flagellar branchessubequalin length to the segments; wings strongly tinged with brown, prearcular field and stigma
slightly darker brown; male hypopygiumwith apex of basistyleproduced
caudatl into a long flatteneclblade, its apex slightly expanded.

L e n g t h ,a b o u t2 l m m . ; w i n g , 1 6 m m . ; a n t e n n a ,M a le.
of headelongate,nearlyequal
about5 mm. Frontalprolongation
to the remainder,chestnutbrown, nasus short; palpi black.
Antennaewith scapeelongate,dark brown, pedicelyellow; branched f lagellarsegmentsbrownishyellow with dark brown branelongate,black; branchesof segches;terminalsimplesegments
ments short, equal, subequalin length to the segments.Headgrayish brown, orbits vaguely more yellowed.
Pronotumbrownishyellow, vaguely patternedwith brown,
especiallythe lateralends and anteriorborder of scutum.Mesoobscureyellow with four reddishbrown stripes
notal praescutumthat are vaguelyborderedby pale brown, the centralmore darkonly at anteriorend,humeralborderbrown;
enedvitta represented
scutal lobes cinnamonbrown; posteriorscleritesof notum dark
regionbrown,gray pruinose.Pleurayellowishgra!, dorsopleurallight brown. Haltereslight brown, knobs slightly darker. Legs
with coxae pale grayish yellow; trochantersbrowriishyellow;
remainderof legs brown, tarsi passinginto black; claws ofmale toothed.Wings stronglytingedwith brown, prearcularfieldand stigmaslightlydarkerbrown; a small more darkenedcloud; Rs
on Ru-r abover-m; veins brown. Veins behindR glabrous
Venation:punctures.
setigerous
and R+-s with sparsemicroscopic
cell Ml narrowlysessile;m-cu on M"-n shortlybeforefork.
brown, lateral bordersof
Abdominaltergitesclark reclclish
by blackish,sternites
beyondthe first narrowlyrnargined
sesnlents
slightly paler reddishbrown; outer segmentsand hypopygium
more darkened.Male hypopygium( Fig. 54) with tergite,t, narrowed posteriorly,apical border with two slenderlobesthat are
Basistyle,b, atseparatedby a narrow U-shapedemargination.
apex of mesaledge producedinto a long flattenedblade thatis directedcaudad,stightly expandedoutwardly,apex gently
emarginate,the blade about one-halfthe length of the style.
Outer dististyle,d, flattened,pale, length about twice the greatest
wiclth; inner style appearingas a broad-basedcurved sclerotizedStudia Ent., vol. 72, fasc. l-4, novembro 19692tl

rod, apex shallowlybidentate,before tip with a small tubercle.
p, includingthe stout gently curved aedeagusand
Phallosome,their tips acute.
slendcrsinuous.rodlikegonapophyses,
Habitat: Colombia.
Holotype,6, Latitude4" l0' North,Longitude73" l0' West,
January20, 1959 (Juan Foerster).The localityapparentlyis in
Meta, east of PuertoLopez.
Ozodicera(Dihexacloruts)tetestylais readily told from other generally
blade
similar speciesby the hypopygial structure, where the elongate_of the baiistyle is quite different from that found in other species.
_Species(D.)
includeO.
ofbasistylepointsapex
athaving spines or spinous
marricantha, O. (O:) exten.sa,O. (O.) piatrix, and O. (O.) septemtrionis,
while O. (O.) biaculeatahas two such spines.

Enderlein
Phacelodocera
P h a c e l o d o c e r aE n r J e r l e i n ;Z o o l . J a h r b . , S y s t . , 3 2 : 2 6 ; l 9 l 2 ;o r i g i n a l d e s i g n a t i o na n d m o n o t y p i c ) .

(type flabellilera (Loew);

Enderlein based this genus on the only known American
speciesso far. assignedto the group. Three other species are
Australasian,including PhacelodoceratasmaniensisAlexander,ofsoutheastern
Australia,and two f rom New Caledonia,P. herroni
Alexanderand P. ntargaritaeAlexander.It shouldbe emphasizedthat the group is very closeto the AustralianPtilogynaWestwoodand may perhapsbe treateclas a subgenusor even fall into thesynonymy.
List of Species
flabellifera (Loew), as Ptilogyna - Brazil.
References:
Linnaea
L o e w, H., Beschreibungeiniger neuen Tipularia terricola
E n t o m o l . , 5 : 3 9 2 - 3 9 3 p, l . 2 , f i g s . l - 3 , w i n g , a n t e n n ao f m a l e ; 1 8 5 1 .
A l e x a n d e r , C . P . , N e w s p e c i eos f c r a n e - f l i ef sr o m S o u t hA m e r i c aP. a r t
XV. - Ann. Ent. Soc. America, 46: 600-602,pl. l, fig. 2, antennaof
female; 1953.

Loew's type, a ntale, was front Brazil, taken by Beske.
It is uncertainwhether the collectorwas C. F. C. Beske whocollectedinsectmaterialsin the vicinity of Rio de Janeiroprior
to 1832,or his son, Carl Heinrich Beske,who made collectionsin Brazil in 1834-35.The only specimenthat I have seen is afemale taken at Campo Bello, Rio de Janeiro,March 16, 1946,
by J. F.' Zikin. As tar as is known to me these are the only
l{+

2t2

A I e x a n d e r, Tropical AmericanTipulidae(VIII)

specimens
taken to date but there may be othersin variouscollections.The wing of the allotypefemaleis shown (Fig. 50).
This differs from Loew'sfiguresof the type male in the venationof the medialfield, includingthe very long medialcrossvein
which
is locatedat the fork of Ms-n Whether this representsan abnormalconditionor the characteris a normalhighlyvariableone
renrainsin question.
The chief characterthat separatesthe f ly f rom other Neotropicalgroupsis the renrarkably
branchedantennaof the male,in conjunctionwith hypopygialcharacters.Loew describedthemale as having l3-segmented
antennae,with the f irst f lagellar
segmentbearing a single very long branch,segments2 to 8
inclusiveeachwith threesimilar branches,arisingclosetogether
near lhe base of the segntent,with the outer three segments
simple.The fernaleantenna,?s describedand figured by thewriter, has 14 segments,
the first flagellarsegmentwith a single
branch,two to eighteachwith two long branches,outerfour segmentssirnple.It nlay be noted that the malesof the two species
f rom New Caledonia,above mentioned,have the largest andmost consipcuously
branchedantennae,in relationto the size ofthe insect,of any known speciesof Diptera.In comparison
withthesethe antennaeof the type of Ptilogyna,ramicornisWalker,
is relativelysmall and inconspicuous.
Tipula Linnaeus
TipulttLinnaeus;
SystemaNaturae,Ed. l0: 5g5; l75g; (typeoleraceaLinnaeus;designatedby Latreille,I g l 0) . It should be
notedthat recentstudiesby Dr. Bo Tjeder,of Lund, haveshownthat the speciesoleracea,?S recognizedby Linnaeus,has been
mis-determinedby later studentsand actually pertainsto thespeciescalledpoludosaby Meigenin 1830;the misnamed,oleracea
was designatedas Tipula submendosaby Tjeder in l94l (consult,
Tjeder, Bo. The identity of TiputctoleraceaL. OpusculaEnt o m o l o g i c1a8, : l l l - 1 1 5 , 3 m a p s ;l g 5 3 ) .
I had earlier discussedthe vast genus Tipula in a seriesof four reportspublishedin the Revistade Entomologiabetween
1945 and 1951
, zS f ollows: Part I, vol. l6 : 330-356,g f igs.;
1945.
SubgenusMicrotipula.Part I I, vol. I 6 : 4ls-440, 2s
figs.; 1945.- SubgenusMicrotipulo.Part III, vol. tT : 172-201,
7 figs.; 1946.- SubgenusEumicrotipula.Part IV, vol. 22: 26s314, 24 f igs.; l95l . - SubgeneraBellardina,Trichotipula,Tipula,Studia Ent., vol. 12, fasc. l-4, novembro1969

2r3

Yamatotipula,Oreomyza, and Lunatipula. In latter years certain
importantchangeshave been adoptedin the latter group of subgenera, as f ollows: Subgenus Tipula now in yamatotipula;
Oreomyza in Pterelacftlsus;Lunatipula in Triplicitipula, as thesenames pertain to the local speciesand are so listed under thepresent account of the genus. Neotipula Alexander, formerty
placed in Tipula, now is included in IschnotomaSkuse and has
been consideredunder that genus in this report. Recentcomprehensivepapers that discuss this re-organizationwithin thegenus Tipula include the following.
A l e x a n d e r , C . P . N e w o r l i t t l e - k n o w nT i p u l i d a ef r o m E a s t e r nA s i a(Diptera), LIII. PhilippineJour. Sci., 93: 77-130,b2 figs.; 1964.
- The same, LVI. Ibid., 94: 235-286,50 figs.; 1965.
- New subgeneraand speciesof crane-fliesfrom California (Diptera:
Tipulidae). Pacific Insects,7: 333-386,33 figs.; l gO5.
Variousother recentpapersconcerningthe taxonomyand biology ofthe genus Tipula includethe following.
Taxonomyol the adult flies
A l e x a n d e r , C . P . , T h e c r a n ef l i e s o f C a l i f o r n i aB. u l l . C a l i f o r n i aI n s e c t
Survey,8: l-269, I plate, 106 maps, 524 figs.; 1967.
M a n h e i m s, Bernharcl,In Erwin Lindner, Die Fliegen der PalaearktischenRegion,Lief. 170: 65-112,figs. 33-65,plate; 1952.Lief. 173-.
113-136,figs. 66-83; 1953.Lief. 238 137-176,
figs. 84-127;1963.Lief.
256: 177-212,figs. 128.172;l!)65. Lief. 267: 213-25e,,
figs. 173-219;
1966.Lief. 270 257-288,figs. 220-255;1967 (to be continued).
S a v c h e n k o , E u g e n ( S a v t s c h e n k oF)a, u n a U S S R ,D i p t e r aI I , n o . 3 .
Fam. Tipuliclae,subfamily Tipulinae, Genus 'tipula Linn. (part t )l-487, 295 figs.; 1961. The same, (part 2) l-503, 382 figs.; 1964
(two further parts to appear).
Biology of the genus Tiputa
B r i n c ll e , A l l a n ,T h e l a r v a ea n d p u p a eo f t h e B r i t i s hT i p u l i n a e( D i p t e r a :_ Tipulidae).- Trans. Soc. British Ent., l4: 63-114, t'7g figs.; 1960.
C h ! s w e l l , J . R . , A t a x o n o m iacc c o u not f t h e l a s t i n s t a rl a r v i e o f s o m e
B r i t i s h T i p u l i n a e( D i p t e r a :T i p u l i c t a e )- . T r a n s . R . E n t . S o c .L o n c l o n ,108: 4W-484,I l0 f igs.; 1956.
T h e o w a I d , B r . ( T h . H . V a n L e e u w e n ) ,D i e E n r v i c k l u n g s s t a d ideenr
Tipuliden (Diptera, Nematocera) , insbesonderecler V/est-PalaearktischenArten.
Tijd. v. Entomologie,lo0: l-XI, lg5-30g,332 figs.;
I 957.
- Bgstimmungsbticher
zur Boclenfauna
Europas.Lief. Z. FamilieTipulidae(Diptera,Nematocera)Larven uncl Puppen,l-100, 344 figs.; 1967.
The general structureof the adults ancl the immaturestages havebeen treated in such detail in the referencesgiven that they need not
be discussedf urther here.

214

A I e x a n d e r , T r o p i c a l A m e r i c a nT i p u l i d a e ( V I I I )List of Species

SubgenusBellardinaEdwards
T l p u l a ( B e l l a r d i n a )E d w a r d s ; A n n . M a g . N a t .
Bellardr, as cravieri; original designation).

Hist., (10) 8:82;(type craverli

Mexico.
craverii Bellardi.
Guatemala,Venezuela.
cydippe Alexander.
Mexico.
edwardsi Bellardi.
Mexico.
larga Alexander.
Colombia, Venezuela,Ecuador, Bolivia.
obliquefasciataMacquart.
parrai Alexande
Mexico.
United States.
schizomeraAlexander.- Mexico; south'westerntheobrominaEdwards. - Venezuela,Ecuador, Peru, Bolivia.
wetmoreanaAlexander.- Guatemala.
SubgenusEumicrotipula Alexander
M i c r o t i p u l a ( E u m i c r o t i p q l . a )A l e x a n d e r ; B u l t . i ! 1 u 9 . d ' H i s t . n a t . , P a r i s t 9 2 2 : 7 1 ;
1 9 2 2 ; ( t y p e m a c r o t r i c h i a t a A l e x a n d e r ; o r i g i n a l d e s i g n a t i o)n.

abortiva Alexander.
Peru.
absonaAlexander. Ecuador.
accipitrina Alexander.
Ecuador.
accumulatrix Alexander.
Guatemala.
aedon Alexander.
Venezuela.
aglossa Alexander.
Bolivia.
agrippina Alexander.
Ecuador.
albifasciata Macquart.
Chile.
amblythrix Alexander.
Peru.
amphion Alexander.
Chile.
andalgala Alexander.
Argentina.
Argentina.
andina Brdthes.
andromacheAlexander.
Venezuela.
angolensisAlexander.
Chile.
antarcticaAlexander.
Argentina,Chile.
anthonymphaAlexander. Argentina, Chile.
apterogynePhilippi.
Chile.
araguensisAlexander.
Venezuela.
araucania Alexander.
Chile.
arecuna Alexander.
Venezuela.
armillata Alexander. Colombia.
Argentina.
asaroton Alexander.
Peru.
asteria Alexander.
Bolivia.
atacama Alexander.
Peru.
atamelesAlexander.
atroscapa Alexander.
Brazil.
Paraguay.
atrovelutina (Alexander), as Microtipula.
Peru.
auricomata Alexander.
Argentina.
austroandinaAlexander.
azteca Alexander.
Mexico.
Chile.
btickstrtimiAlexander.
Costa Rica.
balloui Alexander.
Argentina.
borretoi Alexander.
Bolivia.
bathromecesAlexander.
bigotiana Alexander.
Chile.

Studia Ent., vol. 12, fasc. l-4, novembro 1969
bogotana Alexander.- Colombia.
brithesiana Alexander.- Argentina, Chilebrevicoma Alexander. - Chile.
Panama.
browniana Alexander.
Argentina.
hruchi Alexander.
Peru.
callisto Alexander.
Peru.
callithrix Alexander.
Peru.
campo Alexander.
Venezuela.
Alexander.
capucina
Colombia, Ecuador, Peru.
ca-rizonaAlexander.
sphaeruliferaEdwards. - Ecuador.
Syn.
chacopatachacopata Alexander.- Venezuela.
Peru.
c. chanca Alexander.
sp. n. (Fig. 57, 6 hypopygium).- Peru'
chanchanensis
charmosyneAlexander. - Peru.
Ecuador.
chicana Alexander.
Chile.
chilensisAlexander.
Chil€.
chitlanica Alexander.
Chile.
Alexandel.
chilotaclarkiana Alexarider.- Chile.
Peru.
clavaria Alexander.
Ecuador.
consonata Alexander.
Bolivia, Peru.
conspiciltataAlexander.
Ecuador.
coronaria Alexander.
(Alexander),as Microtipula.- Costa Rica'
costaricensis
Chile.
crepero Alexander.
Venezuela.
cristata Alexander.
c r o s s o s p i l aA l e x a n c l e r . C h i l e .
Peru.
curinao Alexander.
Peru.
c),clomeraAlexantler.
Hispaniola: Dominican Republic'iarlingtoniana Alexander.
Ecuador.
delectato Alexander.
diardis sp. n. (Fig. 59, venation). - Argentina'
Bolivia.
dictyophora Alexander.
Argentina, Chile.
dimorpha Alexander.
(Fig.
58,
6 hypnpygium). - Chile'
diodonta sp. n.
Venezuela.
duidae Alexander.
Chile.
duseni Alexander.
Ecuador.
efficax Alexander.
Ecuador.
emerita Alexander.
Argentina, Chile.
enderleinanaAlexander.
Peru.
exilis Alexander.
Bolivia.
expleta Alexander.
fatidica Alexandet.- Ecuador.
Chile.
fazi Alexander.
ftaviduta Alexancler. Ecuadtlr, Peru, Bolivia.
- Argentina.
flavoannulataJacobs.
Colombia.
loersterianaAlexander.
Bolivia.
forsteri Alexander.
- Bolivia.
lortior lortior Alexander.
Bolivia.
Alexander.
f. arepuchoensls
Alexander.
f. klausma Alexander.
Ecuador.
fraudulenta Alexander.
Chile.
fuegiensisAlexander.
Chile.
glaphyroptera PhiliPPi.
Bolivia.
phora Alexander.
glosso2t5

2t6

A I e x a n d e r, Tropical AmericanTiputidae(VIII)

graphica Schiner.
<South America>.
guarani Alexander.
Brazil.
hedymopaAlexander. Ecuador.
hostifica Alexander.
Brazil.
huancaAlexander. Peru.
hylonymphaAlexander. Chile.
iguazensis(Alexander),as Microtipula.
Argentina.
immerensAlexander.
Venezuela.
immorsaAlexander.- Peru.
inaequiarmataAlexander.
Bolivia.
inaequidensAlexander.
Ecuador.
inca Alexander. Peru.
inconditaAlexander. Bolivia.
inlidelis Alexander.- Peru.
infinita Alexander.
Venezuela.
innubensAlexander.
Peru.
invigilans Alexander.
Peru.
itatiayensisAlexander.
Brazil.
jacobsianaAlexander. Argentina.
jaennickeanaAlexander. Chile.
jivaro Alexander.
Ecuador.
jubilans Alexander. Ecuador.
juventa Alexander. Ecuador.
kathemaAlexander. Peru.
kuehlhorniAlexander.- Bolivia.
kuscheliAlexander. Chile.
lanigera lanigera Alexander. Argentina.
l. tau Alexander. Chile.
laterosetosaAlexander. Colombia.
Iatifolia Alexander. Chile.
legitima (Alexander),as Microtipula.
Paraguay.
Alexander.
ligulata
Argentina,Chile.
ligulipenicillataAlexander.- Mexico.
longibasisAlexander.
Mexico.
Iongurioides
Alexander. Bolivia.
macintyreana
Alexander. Ecuaclor.
macrotrichiata(Alexander),as ilicrotipula.
Brazil.
Alexander. Argentina,Chile.
magellanica- Chile.
Syn.- magellanicola
Alexander.
marmoripennisRondani
Venezuela.
martin-browniAlexander. Ecuador.
mecoglossaAlexander. Colombia.
mediodentataAlexander. Peru.
meridiana Edwards
Ecuador.
microspilotaAlexander.- Chile.
miranhaAlexander. Colombia.
mithrodatesAlexander. Peru.
mitua Alexander. Colombia.
mocoa Alexander. Colombia.
moctezumae
Alexander. Mexico.
monilifera Loew.
Brazil.
moniliferoidesAlexander.
Argentina, Brazil.
moniliformisRdder.- Colombia.
mordax Alexander. Brazil.
morphaeaAlexander. Ecuador.
navarinoen.srs
Alexander.
Chile.

Studia Ent" vol. 12, fasc. 14, novembro1969
neivai Alexander.
Venezuela.
Peru.
nethis Alexander.
nigriscapa Alexander.
Surinam.
Mexico.
nimbinervis Alexander.
Alexander.
Ecuador.
nolensAlexander.
Argentina, Chile.
nordenskjoldi
Argentina.
nothofagetorumAlexander.
Ecuador.
notoria Alexander.
Alexander.- Mexico.
novaleonensis
novatrix Alexander.r- Peru.
Argentina.
nubilera van der Wulp.
Ecuador.
obirata Alexander.
obscuricinctaAlexander.- Mexico.
Peru.
odontomeraAlexander.
Venezuela.
olssonianaAlexander.
Peru.
omniluteaAlexander.
ona Alexander.
Chile.
Argentina.
oreonymphaAlexander.
Alexander.
Mexico.
orizabensis
Colombia.
ornaticornisvan der Wulp.
osculataAlexander. Ecuador.
palencaAlexander. Venezuela.
Peru.
palitans Alexander.
Chile.
pallidisignata pallidisignata Alexander.
p. salutatoria Alexander.
Chile.
pallidineuris Macquart (pallidinervis of authors)
Venezuela.
pantherina'paranensisAlexander.
Brazil.
Alexander.
Argentina, Chile.
parviloba Alexander.
Argentina, Chile.
patagoniccAlexander.
Bolivia.
pediformis Alexander.
Peru.
perjovialis Alexander.
perstudiosaAlexander.- Brazil
petolura Alexancler. Chile.
petaluroidessp. n. (Fig. 60, 3 hypopygium)
Ecuador.
petiolaris Alexander.
Ecuador.
phalangioidesAlexander.
Argentina, Chile.
philippiana Alexander.
Chile.
pictipennisWalker.
Ecuador.
Edwards
pilulifera
Chile.
pirioni Alexander.
Peru.
piro Alexander.
platytergata Alexander. Bolivia.
Ecuador.
procericornisEctwards
Brazil.
profuga Alexander.
Argentina.
protrudensAlexander.
psittacinaAlexander. Argentina,Bolivia.
quadrisetosaAlexancler.
Peru.
quichuaAlexander. Peru.
Mexico.
reciproca Alexander.
redunca Alexander.
Colombia.
reedi Alexander. Chile.
resplendensAlexancler. Ecuador.
Ecuador.
riveti riveti Edwards
r. tolimensisAlexander. Colombia.
rucane Alexander. Peru.

Colombia.

Chile.

2t7

2t8

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

rufirostris Bigot.
Chile.
runtunensisAlexander.
Ecuador.
Alexander.
sariapampae
Peru.
Alexander.
satrapa
Chile.
schachovskoyiAlexander.
Argentina.
Panama.
scriptella Alexander.
Peru.
semivulpinaAlexander.
serrilobata Alexander.
Chile.
sefval Alexander.
'
songoanaAlexander.
Bolivia.
spatuliferaAlexander.- Chile.
spilota Wiedemann.
Brazil.
steinbachiAlexander.
Bolivia.
stenoglossa
slenoglossaAlexander.- Peru.
Peru.
s. propitia Alexander.
Peru.
suavissimaAlexander.
Philippi.
subandina
Chile.
Edwards
Ecuador.
subcana
subglobrataAlexander.- Peru.
subligulata Alexander.
Chile.
Bolivia.
tanymetra Alexander.
Argentina. Chile.
tehuelcheAlexander.
tephronota Alexander. Peru.
tersa Alexander.
Chile.
tersoidesAlexander.
Chile.
Ecuador.
thalia Alexander.
tovarensisAlexander.
Venezuela.
triemarginata Alexander.
Chile.
trispilota Alexander. Peru.
Argentina, Chile.
tristillata Alexander.
Ecuador.
tunguraguanaAlexander.
Alexander.
Mexico.
tyrannaPeru.
unistriata Alexander.
Peru.
ursula Alexander.
Chile.
valdivianaPhilippi.
varineura (Bigot), as Nephrotoma; (variinervis of authors)
Mexico.
virgulata Williston
Syn.
oreomyzoidesAlexander.- Mexico.
yotiva Alexander.- Bolivia.
willinki Alexander.- Argentina.
Argentina.
wittei Alexander.
Peru.
woytkowskianaAlexander.
yanamonteanaAlexander.
Peru.
yungasensisAlexander.
Bolivia.
Mexico.
zeltale Alexander.
Alexander.
Mexico.
zeugmata
Mexico.
zotzil Alexander.

Chile.

SubgenusLunatipulaEdwards- dwards; Ann. Mag. Nat. Hist., (t0) g: gl;
T i p u l a ( L u n a t i p u . l a lE
L i n n a e u s ;o r i g i n a l d e s i g n a t i o n ) .

abscfssaAlexander.
Mexico.
dampfiana Alexander.
Mexico.
Iagunicolasp. n. (Fig. 61, A hypopygium)michoacanaAlexander.- Mexico.

Mexico.

l93t; (type tunata

Studia Ent., vol. 12, fasc. 14, novembro1969

2r9SubgenusMicrotiPulaAlexander
tllcrotipala Alexander; Ann. Ent. Soc. Amerlca, 5: 360; l9l2;
Alexinder; orlginal deslgnatlon)-(type amazonlca(MexlcoD.. f.),Cien.Pio-logicas
svri.--'ri'pnr-o7'oiiibei_Aiiiiind#;Anal. Esc. Nac.
-'i;zzij--lgtil"'iJ--rlinlittt'epirotomodes;(type smitodon Alexander,origlnal
deslgnatlon).

aequitorialis Alexander.- Ecuador.
Syn. - insolabilis Alexandet. - Peru.
Brazil.
aflabilis Alexander.
Peru.
akestra Alexander.
Peru.
.
alectro Alexander.
amara Alexander.- Peru.
Brazil, Surinam.
amazonica (Alexander), as MicrotipulaamoenicornisAlexander. - Argentina.
Bolivia.
apoltyon Alexander.
ippendrns (Enderlein), as Macromastix. - Ecuador'ariranhae Alexander. - Brazil, Paraguay.
ir:matipennis armatipennisAlexander. - Argentina, Brazila. napoensisAlexander. - Peru.
Bolivia.
austrovolensAlexander.
Bolivia.
aymara Alexander.
Mexico.
bitimeki Alexander.
biprolata Alexander. - Brazil.
bitributa Alexander.- Peru.
Brazilblaseri Alexander.
as Megistocera.- Brazil.
(wiedemann),brasiliensis
bruisi Alexander. - Lesser Antilles: Grenada'
Ecuador.
cerogama Alexander.
Ecuador.
cithiriformis Alexander.
Colombia.
colomiicota Alexander.
Guatemala'contemplafaAlexander.
(Alexander), as Microtipula. - Costa Rica.
co,stariiensis
crassisfYtaAlexander..- Peru.
ctenopyga Alexander. - Panama.
Peru.
decensAlexander.
decorata (Alexander), as Habromastix. - Brazil, Paraguay.
Venezuela.
detecta Alexander.
Peru.
diacanthos Alexander.
Syn. - eftera Alexander.- Peru.
Peru.
diadexia Alexander.
Peru.
didactyla Alexander.
Argentina.
didolos Alexander.
dirhabdophora Alexander.- Panama.
Peru.
discophora Alexander.
Btazil.
efferox Alexander.
Peru.
effeta Alexander.
Ecuador.
effulta Alexander.
epione Atexander.- Bolivia.
erostrata Alexander. - Colombia.
Brazil.
eurymera eurymera Alexander.
Brazit.
e. goyazicola Atexander.
Paraguay.
e. paraguayicola Alexander.
Ecuador.
Ialcifer Alexander.
Btazil.
feticiana Alexander.
Brazil.
ferocia Alexander.

220

A I e x a n d e r, TropicarAmericanTipulidae(vlll)

Paraguay.
liebrigi Alexander.
Brazil.
flavopolita Alexander.
gladiator Alexander.
Brazil.
guato Alexander.
Brazil.
guerreroensis Alexander.
Mexico.
,gutticellula Alexander.
Brazil.
heterodactyla Alexander.
Ecuador.
hexamelania Alexander.
Cuatemala.
histrionica Alexander.
Costa Rica.
horribilis Alexander.
Bolivia.
icasta Alexander.
Ecuador.
peru.
impatiens Alexander.
inaequilobata Alexander.
Ecuador.
inarmata Alexander.
Argentina.
infida Alexander.
Ecuador.
intemperata Alexander.
Ecuador.
jivaronis Alexander.
Ecuador.
jordanensis Alexander.
Brazil.
juquiella Alexander.
Brazil.
klagesi Alexander.
Brazil.
Iagotis Alexander.
Brazillanguidula Alexander.
Peru.
laticostata Alexander.
Brazil.
Ietalis Alexander.
Brazil.
Iichyana Alexander.
Venezuela.
Iuctilica Alexander.
Ecuaclor.
luteilimbata Alexander.
Ecuador.
Iyriformis Alexander.
Peru.
macrosterna (Alexander), as Pachyrhina.
Guatemala,Honduras.
mandator Alexander.
Costa Rica.
manniana Alexander.
Bolivia.
mediocompressaAlexander.
Ecuaclor, peru.
megalyra Alexander.
Ecuador, peru.
monocera Alexander.
Peru.
mulfordi Alexander.
Bolivia.
multimoda Alexander.
Colombia.
myriatricha Alexander.
Peru.
neolenta Alexander.
Ecuaclor.
Syn.
Ienta Alexancler.
nicoya Alexander.
Costa Rica.
nigroabdominalis (Alexander), as Habromastix.
Brazil.
nigrovariegata Alexander.
Ecuaclor.
niobe Alexander.
Venezuela.
opipara Alexander.
Bolivia.
ortoni Alexander.
Bolivia.
pala Alexander.
Brazil.
palaeogamaAlexander.
Ecuaclor.
paloides Alexander.
Brazil.
paralenta Alexander.
Costa Rica.
pararia Alexander.
Peru.
parishi Alexander.
Brazil.
penana Alexander.
Bolivia.
perangustulaAlexander.
ccllombia, venezuela.
percompressaAlexander.
Ecuador.
percomptaria Alexander.
Ecuador.
peru.
perdelectaAlexander.

Studia Ent., vol. 12, fasc. 14, novembro 1969I

2Zl

perlaticosta Alexander.
Brazil.
plaumannianaAlexander.
Brazil.
plumbeithoraxAlexander.
Peru.
pontifex Alexander.
Peru.
porrecta Alexander.
Peru.
pretiosa Alexander.
Peru.
pritchardi pritchardi Alexander.
Mexico.
p. glabristyla Alexander.
Mexico.
proctotricho Alexander.
Ecuador.
prolixisternaAlexander.
Ecuador.
quadricollis Alexander.
Bolivia.
rectangulusAlexander.
Peru.
regre.ssaAlexander.
Venezuela.
retrorsa Alexander.
Bolivia.
scaphula Alexander.
Panama.
Alexander.
scelesta
Bolivia.
schildeanaAlexander.
Costa Rica.
schwarzmaieriAlexander.
BrazilseptemhastataAlexander.
Bolivia.
sexcincta Alexander.
Brazil.
Alexander.
smilodonEcuador.
smithi Alexander.
Brazil.
sparsipila Alexander.
Bolivia.
spinicaudoAlexander.
Panama.
sternohirsutdAlexander.
Ecuador.
subarmata Alexander.
Paraguay.
subaymaraAlexander.
Bolivia.
subeffeta Alexander.
Peru.
subeffultasp. n. (Fig. 63, 6 hypopygium)Peru.
subinfuscataWilliston
Lesser Antilles: St. Vincent.
subtecta Alexander.
Venezuela.
Syn.
auricularis Alexander.
Ecuaclor.
tancitaro Alexander.
Mexico.
temperatoAlexancler. Mexico.
tenuiculaEnderlein
Colombia.
terutilobataAlexrncler.
Ecuador, Peru.
terpischoreAlexe.itder. Guatemala.
terribilis Alexander.
Ecuador.
tijucensis Alexander.
Brazil.
topoensisAlexander.
Ecuador, Peru.
trichoprocta Alexander.
Peru.
trihastata Alexander.
Peru.
trinidadensis(Alexander),as Pachyrhina.
Trinidad.
trinitatis Alexander.
Trinidadtucumanensis
Alexander.
Argentina.
urophora Alexander.
Colombia,Ecuador.
virgilia Alexander.
Peru.
volen.sAlexander.
Peru.
zeteki Alexander.
Panama.
zonalis Alexander.
Venezuela.
SubgenusPectinotipulaAlexander
Pectino-t-i-pulaAlexander; .Jour, N. y.
der Wulp; originaldeslgnation).

(.
l.

t

Ent. Soc., 2g: g; lg20; (type argenlna van

222

Alexander,T r o p i c a lA m e r i c a nT i p u l i d a e( V I l l )argentina (van der Wulp), ?s Ozodicera. Argentina.
boliviensis(Alexander),as Pectinotipula. Bolivia.
titicacae(Alexander),as Pectinotipula. Bolivia, Peru.
tucumana(Alexander),as Pectinotipula. Argentina.
SubgenusPterelacftisusRondani
PterelachisusRondani; Rev. Zoolog. (Soc. Cuv.) 5: 243; 1842; (type Bertei Rondani;
not berteii of authors; monotypic).

bellardiana Alexander.

Mexico.

Subgenus Trichotipula Alexande
T i p u l a ( T r i c h o t i p u t a ) A l e x a n d e r ; P r o c . A c a d . N a t . S c i . P h i l a d e l p h i al g l 5 : 4 6 8 ; 1 9 l 5 ;i t y p e o r o p e z o i d e sJ o h n s o n , N e a r c t i c , o r i g i n a l d e s i g n a t i o n ) .

Mexico. .
aplecta Alexander.
Mexico.
kraussi Alexander.
Mexico.
religiosa Alexander.
Mexico.
selanderi Alexander.
Mexico.
uxoria Alexander.
Mexico.
vultuosa Alexander.
Subgenus Triplicitipula Alexander
Tipula (Tripticitiputa) Alexander; Pacific Insects,7:354;
Nearctic, original designation).

hoogstraaliAlexander.

1965; (type triplex Walker,

Mexico.

SubgenusYamatotipulaMatsumura
Y a m a t o t i p u l aM a t s u n t u r a ; T h o u s a n d I n s e c t s o f J a p a n , .4 d d . . . 2 : . t 9 1 6 ; ( t y p e n o v a
W a l k e f , a s N o h i r a e M a t s u m u r a , P a l a e a r c t i c ,o r i g i n a l d e s i g n a t i o n ) .

jamaicensisAlexander.
Jamaica.
'ludoviciana
Alexander.
SoutheasternUnited States; Cuba.
Eastern United States; Bermuda.
sayi Alexander.
Mexico.
subvirgo Alexander.
Mexico.
virgo Osten Sacken.
Tipula species,subgenusor genus uncertain<<SouthAmerica>>.
filigera Walker.
quadrimoculata Bellardi.
Mexico.
Colombia.
seticornisMacquart.
Peru. Apparently atabida Enderlein.
suggests Zelandotipula.
Chile.
trimaculata Macquart.
Brazil.
trina Wiedemann.

Miuotipula;

Vane-Wright

Tipula (Acutipula) gaboonensis Alexander
T i p u l a g a b o o n e n s i sA l e x a n d e r ; B u l l . l l t u s . d ' H i s t . n a t . , P a r i s 1 9 2 0 : 4 0 3 - 4 0 5 ; 1 9 2 0 .
Tipula (Acutipulal Alexander; Arkiv fiir Zoologi, 16, no. l8: ll-12; 1924.
T i p u l u ( A c u t i p u l a ) g a b o o n e n s i sA l e x a n d e r ; P h i l i p p i n e J o u r . S c i . , 5 7 : 1 0 8 ; 1 9 3 5 .
T ' i p u l a ( A c u t i p u l a l S a v t s c h e n k o ;F a u n a U . S . S . R u s s i a 7 9 , D i p t e r a I I , 3 : 3 6 0 - 4 2,7
f igs. 214-262; 1961.

Studia Ent., vol. 12, fasc. l-4, novembro1969

223

A nraleof this specieswas received,labelledTela Progresso,
Honduras,March 5, 1923,collectedby T. H. Hubbell (Universityof Michigan) . The specimenwas included in a collection ot
Honduran crane flies taken by Hubbell and there is a question
as to how this record is to be construed.Presumablyit representsa ntis-labellingof the specimenbut the questionof accidental
introduction of the species into Tropical Anrerica cannot be
overlooked.The present species is the type of the subgenus
AcutipulaAlexander,vastly developedin the Old World, including
the Palaearctic,Oriental and Ethiopian regions, but with no
known f ront the New World.
representative

sp. n.
Tipula (Eumicrotipula)chanchanensis
A l l i e d t o s u b l i g u l a t a ;m e s o n o t a lp r a e s c u t u mg r a y w i t h f o u r s o l i d l yrfark brown stripes, pleura light gray, patternedwith darker; antennae
with outer end of scape and the peciicelyellow, f lagellum black; femora
light brown basally, outer half bro'wnishblack, encltlsinga yellow subterminal ring, claws simple; wings marbled with cream yello'w andbrown, including four darker subctlstal areas; male hypopygium withposterior border of tergite emarginate,outer end darker and nearly
glabrous; posterior encl of inner clististyleenlargeclancl rounded; eightha small flattenedplate, apex shallowlyemarginate,
iternite with appenclaget h e . r o u n d e dl o b e s w i t h s h o r t s e t a e .
M a I e.

L en g t h , a b o u t I I

n l l l ' t;. w i n g , 1 7 n l n l .; a n t e n n a ,a b o u t 5 . 2 n l n ' I . F r o n t a l p r o l o n e a t i o nt l t h e a c l r e l a t i v e l y l o n g ,n e a r l ye q u a l t o t h e r e n r a i n c l elri,e h t b r o r v n i s hg r a y a b o v e ,c l e a r e rb r o w n o n s i d e s ; n a s L l sel o n g a t e ,t u ft e d w i t h l o n g y e l l o w s e t a e ;p a l p i b r o w n , t e r n r i n a ls e g n l e n tb r o w n i s h b l a c k . A n t e n n a ew i t hscape light yellow, basal third pale brown, pedicel light yellow,
with small basalenlargements,
flagellunrblack; flage'llarsegnrents
krngestverticilsa trifle less than the segments.Front pale brown,
gray pruinose;posteriorvertex Sray,weakly patternedwith lightb r o w n , i n c l u d i n ga d i f f u s ec e n t r a ll i n e a n d b r o a d e rl a t e r a la r e a s ,orbits narrowly gray.
Pronotal scuturnbrownish graf, anterior nrargin variegatecl
rvith darker brown, scutellurnyellowed. Mesonotal praescutunt
Sray, with f our solidy dark brown stripes, interntc'diatepair
ltroader,virtually contiguousat ccphalichorcler,lateral praescutal
rnargins broadly darkened; scuturn light gray, each lobe withtwo dark brown areas,the postc'riorone rrruchlarger; scutellum
g r s / , w i t h a b r o w n c e n t r a ll i n e , p a r a s c u t c l l lai g h t b r o w n ; m e d i o tergitewith centralthird bron'n,siclespalcr brown, Sray pruinose,

224

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

suture and posterioranapleurotergite
brownishyellow, remainderof pleurotergite
dark brown,the lowerelevatedpart graypruinose.
Pleura light gray, conspicuously
patternedwith darker, including
a brownish gray dorsal stripe extendingfrom the propleurato
ttt. pleurotergite,ventral sternopleuritepaler gray; dorsopleural
membraneconspicuously
tight yellow. Haltereswith stem lightbrown,baseyellow,knob darker,apex restrictedlywhitened.Legs
with coxaelight brown, gray pruinose;trochanters
yellow;femora
Iight brown, yellowedbasally,outer half brownishblack, witha conspicuous
yellow subterminalring that is subequalto thebrownish black apex; tibiae and tarsi light brown, outer tarsal
segmentsdark brown; claws small, simple. Wings with therestrictedgroundpalecreamyellow,with an extensive
pale brownpattern,with darkerbrown subcostalareas,includingfour in cellSc, thc'semore extensivethan the ground interspaces;stigma
slightly paler; the more extensiveyellow ground areas include
the prearcularfield, basesof Anal cells,a transverseband acrosscellsR and M beyondmidlength,and a broad subbasalarea inouter radial field, extendingfrom veinsC to Mt-z; veinsbrown,darkerin the subcostalnrarkings,yellow in the brightenedcostal
interspaces.
: R.snc.arlytwice m-cu; petioleof cell M L
Venation
about three times m.
Abdomenelongate;basal tergite brownish gray,succeeding
tergitesyellow, each with a sublateraldark brown stripe thatis narrowlyinterruptedat posteriorborder,lateralmarginsmore
broadlyyellow; sternitesyellow; outer segments,
includinghypopygium, more extensivelydarkened.Male hypopygium(Fig. sr)with tergite,t, transverse,
broadestat midlength,posteriorbordervery shallowlyentarginate,
darkenedancl alnrostglabrous.Appendagesof basistyle,b, includinga larger setiferouslobe ancl
a inicroscopic
papillosearm, its tip truncate.Outer dististyle,d,moderatelydilatedon outer half, with long but relativelysparsesetae;inner style long and narrow, apex of the slenderlower
beak obliquelytruncated;dorsal crest with a row of about 30
strong yellow setae,those at ends of row smaller,the central
group powerful; posteriorend of style in region of the outerbasal lobe enlarged and rounded,with relatively few setae.
Gonapophysis
blackened,shapedas figured, g, outer third h?rrowed. Eighth sternite,8 s,. with appendagea small flattened
dark-coloredplate, its length about twice the breadth,apex witha shallowemarginationto form paler roundedlobes with shortsetae.

StudiaEnt., vol. 12, fasc. l-4, novembro1969

225

59

rIllllr...lI

6)

Fig. 56. Tipula (Eumicrotipula) diardis sp. n.; venatio!: - _Fig.
-Ti.puta.
.57. Tipula .(Eumicr-o(E_umicrotipula)Fig. 58.
ttpnn) chdnchai,ensissp. n.;-male hypripyglqm. - Fig.
diodohta sp. n.; male h-ypopygium.
- 59. Tipula (Eumicrotiptllg) diardis sp. n.;
Fig. 60. Tipula (Eumicrotipula) petaluroides- sp-. n.; male
male hyp<ipygldm.
Figs. 61, 62. Tipula (Lunatipula) Iagunicola sp. n.; .male hypqpygiu{n.
hypopygium.
Fig. 63. Tipula (Microtipula) subelfulta sp. n.; male hypopygium. -. Symbols:
D ,a a s i s t y l e ; 'd , d i i t i s t y t e ; ' g , b o n a p o f i h y s i sp; , p h a l l o s o m e ;s , s t e r n i t e ; l , t e r g i t e .

l5

A I e x a n d e r, TropicalAmericanTipulidae(Vlll)

226

Habitat: Chile.
Holotype,d, Chanchan,of Lake
Valdivia,in forestsouthwest
Panguipulli,
altitude120-600meters,March2, 1955 (Pena).
'f

is most similar to T. (8.)
ipula (Eumicrotipula) chanchanen.srs
subligulafaAlexander,with the wing pattern much the same,the whitened
band in outer cells of raclial field more clearly defined. The details of
the hypopygium are distinctive,incluclingthe tergite, lobes of basistyle,inner dististyle, gonapophyses,and the eighth sternite. In subligulata theinner dististyle has the lower beak broad and the posterior end of stylemore narrowed, virtually glabrous.

Tipula (Eumicrotipula)diardis sp. n.
Belongs to the macrotrichiatagroup; size meclium (wing of maleI I mm.); general coloration of mesonotumlight brown, patterned withdarker, including four praescutalstripes; outer flagellar segmentsblackenecl; femora yellow, tips narrowly brownish black; wings light brown,
patterneclwith darker brown and whitened areas, the former including
four major areas in cell Sc; sparse macrotrichiain outer ends of cetts
f r a n d M t , n o s t i g m a l t r i c h i a ; p e t i o l e o f c e l l M l s u b e q u a lt o m ; m a l ehypopygium with posterior margin of tergite with an oval notch, theangles produceclinto slender blacles.

M a I e.
L e n g t h ,a b o u t 1 0 . 5n l r n . ;w i n g , I I n l m . ; a n t e n n a ,about 4 nllll. Frontal prolongationof heacl light brown, above
with a narrow nlore clarkenecl
vitta; nasusslender,more yellowed,
with long yellow setae;palpi brownishblack. Antennaerelatively
long, slightly exceedingone-thirdthe wings; scapeobscureyellow,
pedicelpaler; first flagellarsegnrentbrownishyellow, baseweakly
darkened;segnrentstwo and three bicolored,the very small basal
enlargements
black, renrainderlight brown; succeedingsegments
black, about one-half longer than their verticils. Head brownish
SraI, posterior vertex darker; vertical tultcrcle very low, witha n i n r p r e s s e cd e n t r a l l i n e .
Pronotunrbrownish gray, inclistinctlypatternedwith brown.
Mesonotalpracscutunrwith ground light brown, with four darker
stripes, including an internrediatepair that are gray in front,
borderedby darker brown, incluclingthe centralvitta, the stripes
strongly narrowed behind at the suture, lateral stripes narrow,dark brown; scutal lobeschiefly dark brown, centralarea brownish yellow, immediatelybehind suture with two parallel darkened
lines; posteriorscleritesof notunr light brown, rnediotergitewith
indicationsof a darker centralline. Pleuraanclpleurotergitechiefly
yellowed,indistinctlypatternedwith very pale brown, especiallyon the anapleurotergite,dorsal pteropleurite,anepisternumand

Studia Ent., vol. 12, fasc. 14, novembrol969

227

ventral sternopleurite;mesonotalvestitureshort and sparse,mostevident on the rnediotergiteand praescutalinterspaces.Haltereswith stent pale yellow, knob infuscated.Legs with coxae brownish yellow, trochantersclearer yellow; femora yellow, tips narrowly brownish black; tibiae brownish yellow, tips similarly
infuscated; tarsi brownish yellow, outwardly passing into darkbrown; claws simple.Wings (Fig. 56) chiefly light brown, patterned with darker brown and pale areas; a seriesof f our large
darker brown marks in cell Sc, restrictingthe deeperyellow interspaces;stigma paler brown; on disk the restrictedwhitenedareas
include an oblique band before cord, extendingf rom cell Rl tovein Cu; a broad post-stigmalband and a comparableareaacrosscell /s/ A along the vein, includingthe subterminalpart of cellCu; a darkenedseam along vein Cu, nlore intensebetween the
branches;veins brown, more yellowed in the brightened areas.
Macrotrichiaon outer third of cells Rs and M r, in f igure theirposition indicated by stippling; most longitudinal veins beyond
generallevel of origin of Rs with trichia, lacking on basal sectionoI Ms; trichia of vein lst A chiefly at outer end, of 2nd A scattered over outer half; a few trichia on prearcular section of
1s/ A. Venation: R, -, atrophied;Rs long, nearly straight,about
twice m-cu; r-m short; petiole of cell M I subequalto m ; m-cu
shortly before fork of Ms-+.
Basal abdominal segnrentsobscure yellow, lateral tergalbordersblackened,outer segmentsdarker, including central partsof tergites and more than outer half of sternites; hypopygium
light brown. Male hypopygiunr( Fig. 59) with tergite,/, transverse,
posterior border broadly emarginate,with a further small oval
nrediannotch,the adjoininganglesproducedinto slenderglabrous
blades.Outer dististyle,d, snrall, nearly cylindrical;inner stylewith apex of lower beak and lower margin of beak blackened;outer crest weakly corrugated; all setae clelicate and inconspicuous.Gonapophysis,g', with outer blacle terrttinatingina small point, separatedby a U-shapednotch.
H abitat.'Argentina.
Holotype, J, Parque Aconquija, Tucumdn, altitude 80ornetcrs,January I I , 1953 ( PeclroW. Wygodzinsky).
The most similar speciesis 'l'ipulu (Eumicrotiputa)tegitima (Alexander), of Paraguay,which lacks macrotrichiain the wing cells and has
the leg pattern quite distinct. The structure of the hypopygiumof legitima
has not been describeclancl is unknown to me. In the present fly thetergal spines that have suggestedthe specific name are particularly
notewclrthy.
!.tr*

228

A I e x a fl d e r, Tropical AmericanTipulidae (VIII)

Tipula (Eumicrotipula)diodontasp. n.
Size large (wing over 17 mm.); generalcolorationof thoracic notum
gray, patterned with brown; antennaerelatively long, scape and pedicel
yellow, flagellum black; legs brownish yellow to light brown; wings
cream-yellow,with an extensive marbled light brown pattern, 'with aseries of four darker brown subcostalareas that are extendedbackward
across cells R and M; basal abdominal segmentsyellow, tergites withnarrow sublateral lines, outer segmentsmore uniformly darkened; malehypopygium with posterior border of tergite with two acute inner teeth
and broader more lateral lobes; outer dististyleterminatingin a broadened spatula, inner style narrow, posterior border near base with aroundedknob, beak slender;eighth sterniteunusuallynarrow, appendage
simple, about two and one-half times as long as broad, apex truncate.

L e n g t h ,a b o u t l 7 . 5 m n l . ; w i n g , 1 7. 5 m m . ; a n t c ' n M a I e.
fla, about 5 mm.
F e n ra I e .
L e n g t h ,a b o u t 1 9 n t m . ; w i n g , 1 8 n l n t .
Frontal prolongationof head abovebrownishyellow, sparsely
pruinose,cleareryellow below, nasus conspicuous;palpi brownish black. Antennaeof rnale relativelylong; scape and pedicel
yellow, f lagellum black; f lagellar segnrentslonger than their
verticils. Head light gray, vertex with a narrow brown central
vitta, sides of genaeslightly infuscated.
Pronotuntlight brownish gray. Mesonotunrgray, praescutunr
with four stripes,the narrow intermediatepair light brown, nruch
narrower than the central gray area,lateral stripes broader,darkbrown, humeral and lateral borders dark brown; scutum gra!,each lobe with two separateddark brown areas; scutellumlightbrown, heavily light gray pruinose,with a central brown darkening. Pleura reddish brown, variegated with darker brown,
especiallyon anepisternum;
dorsopleuralntenrbraneobscureyellow. Haltereswith stenr pale brown, base narrowly yellow, knob
dark brown, apex yellowed. Legs with coxae brownish yellow,
sparsely pruinose; trochantersyellow, lower face at apex witha brown spot; femora brownish yellow, tips darker brown,
precededby a subequalclear yellow ring; tibiae and tarsi lightbrown, tip of f ormer narrowly darker. Wings of rnale creanl
yellow, heavily patternedand marbledwith light brown, the darkcoloration much exceedingthe yellow ground; a series of fourdarker brown areas in cell Sc, the basal one including also costa
at h and the basesof cells R and M; remainingsubcostalareas
not involving cell C but continuedposteriorlyacrosscells R andM, greatly restricting the yellow ground at midlength of cells;
remainder of the ground patterned with interrupted spots inS t u d i a E n t . , v o l . 1 2 , f a s c . l - ' 1 , n o v e m b r ol ( ) 6 ! )229

virtually all cells, including a marginal series in all cells, mostextensivein cell Rr, very reducedin Mr and 2nd Mr, darkened
area in cells Cu and /sf A more zigzag; in the female,groundareas broader, especiallythe band at near midlength of cells
R and M which continuevirtually uninterruptedin cells Cu andl st A, outerband in cellsR2 to R5 broad; veinsbrown,interspacesof C, Sc, R and Cu yellowed.Venation: Rs lon$, more than
twice Rr-s; petioleof cell Mr about one-halflonger than m.
Abdominal tergites yellorv, basal segmentswith a narrowbroken brown sublateralline, becomingobsoleteat about segment five; sterniteslight yellow, in female posteriorborders ofouter segmentsnarrowly darkened; outer segments,including
hypopygium,darker brown, restrictedlypatternedwith yellowOvipositorwith cerci slencler,straight. Male hypopygium ( Fig'
58) with tergite, t, broad, scooplike,narrowedoutwardly; Posterior margin with four points,the outer pair larger, obtuse,thesmall inner pair appearingas acute points. Basistyle,b, withsclerotizedarmatureat apex, includinga flattenedblade shaped
like an arrowhead,as f igured. Outer dististyle,d, dilated onouter third; inner style narrow, posteriorborder near base with
before base on margin
4 rouncledglabrousknob; beak slender,
with a row of yellow setae. Phallosonteapparently including
slender sirnple spines; aedeagusrelatively long, black. Eighthsternite,Ss, with body unusuallynarrow' strongly arcuated'theirnpressedlines shortly behind nridlength;a sinrplemedian appendage,about two ancl one-half tirttesas long as the breadth
acrossthe truncateclapex, outer half rvith allundantsmall setae
setulae.
ancl densenticroscoPic
Habitat; Chile.
Holotype,6 , Butarnalal,Nahuelbuta,Arauco,altitude 1,4001,600 meters,February 12-20, I 953 ( L. E. Pefra)' Allotopotype'
9 , F e b r u a r y2 5 , 1 9 5 3 ( L . E . P e f r a ) ''tiputa (Eumicrotipula)bigot.a-na
The most similar regionalspeciesare
Alexander,which similarlyhave.marbled
Alexanderand T. (E.) magellanica
male hypowings and a simpie upp.n,lug. on the eighth sternite of thepfrgiur Theie, and other species, differ in hvpopygial structure,piiticularly the tergite, basistyleand dististyles.

Tipula (Eumicrotipula)petaluroidessp. n.
Alliect and similar to petalura; wings with darkened areas lessextensive,includingthose in cell R; a completewhitenedcrossbandbeyond
r'oftl; male hypopygium with gonapophysesand appentlageof eigh.ths t e r n i t er l i s t i n c t i v e .

230

A l e x a n d e r , T r o p i c aAmericanl
Tipuliclae(VI I I )M a I e.
Length,about 1 7 m m . ; w i n g , 1 5 m m . ; a n t e n n a ,about 4.4 mnl.
F e m a I e.
Length, about 20 nlm.; wing, lS-lT mm.;
antenna,about 2.5-2.9 mm.
Frontal prolongationof head relatively long, subequal toremainder, castaneousabove, darker below, nasus conspicuous,
brown; palpi brownish black. Antennaebrownish black, apexof pedicelobscureyellow; f lagellar segrnentsfeebly incised,theintermediateones subequalto their longestverticils.Head chiefly
infuscated,orbits gray, posterior vertex with a capillary darker
central line; vertical tuberclelow, rounded.
PronotuntgraY,patternedwith brown. Mesonotalpraescuturrr
gray with four narrow brownish btack stripes, the long intermediatepair separatedby a narrowerground line, the ends nlore
approxinlated;scutumlight gray, lobesslightly darker; scutellunr
and mediotergite graY, with a poorly indicated darker central
line, parascutellabrown. Pleura and pleurotergitegray, vaguely
patterned with darker, more evident on ventral anepisternunr,
sternopleuriteand meron; dorsopleuralmembraneconspicuously
buffy yellow. Haltereswith stem yellowed,knob infuscated,paling
to brownish yellow. Legs with coxae gray; trochantersobscureyellow, inner f aces darkened;femora obscureyellow, tip narrowly brownish black, preceded by a slighily clearer yellow
area; tibiae and basitarsi brownish yellow, tips vaguely darkened, remainderof tarsi dark brown; claws of male with basal
tooth. wings with ground whitened,with a conspicuousmore
extensivebrown pattern;prearculararea and interspacesof costalf i e l d l i g h t y e l l o w ; f o u r c o s t a l d a r k e n i n g sa, l l r e a c h i n gv e i n c ,the first at arculus,attainingCu hehind; secondand third areas
larger, reachingvein M; fourth clarkeningincludingstigma anda large cloud over anteriorcord and outer end of cell R; wing
tip broadly darkened,with barely indicatedbrighteningsin outerends of cells R; and 2nd M2, cell M+ ancl outer half of M uniformly lighter brown; f urther pale brown darkeningsin cell Cu
except at end and extensivelyin outer ends of both Anal cells;
the pale ground areas include three in cell R and a complete
band beyondcord, crossingwing from C into cell M7,; veins tightbrown, more yellowed in the ground areas.Venation: Rs long,
about two and one-half times m-cui Rr _: strongly elevated;petiole of cell M, nearly twice m; m-cu on Mr, the basal sectionof the latter long, transverse.

S t r r d i aE n t . . v o l . 1 2 , f a s c . l - 4 , n o v e m b r o1 9 6 9231with basal tergite dark brown, secondchiefly yelAbdornc.nlow, succeedingtergites infuscated,the lateral ntargins broadly
p a l e , i n t e r n a l l yw i t h a d a r k e n e dl i n e; b a s a l s t e r n i t ey e l l o w e d ,t h erenrainderdarkened;outer segnrentslnore unifornrlydark brown,of hypopygiunrmore yellowed.Male hypopygiuntthe.appendages( Fig. 60) generallyas in petalura,diftering in certain importantand appendage
cletails,especiallyof the basistyle,gonapophyses,of the eighth sternite.Ninth tergite,t, with posteriorentargination
cleep and narrow, lateral lobe's longer and narrower than inpetalura.Basistylewith dorsal rttargin of lobe producedinto af lattened glabrous blade. Gonaptlpltysis,.{, with both arms
the larger one long and narrow, inner arm outwardly
clistinctive,
ternrinatingin a truncatedhead. Eighth sternite,8S,
expanc'lecl,with appenclaecpale, nearly parallel-sided,narrower than inpetalura.
Hahitat: Chile.
H o l o t y p e , 6 , P i c h i n a h u e l ,N a l t u e l b u t a ,A r a u c o , a l t i t u d e
n t e t e r s ,F e b r u a r y l 2 - 2 O , 1 9 5 3 ( L . E . P e n a ) . A l l o 1,400-1,600
type, 9 , Neltunte,Valdivia, altitude 480 meters,March 3, 1955
( L. E. Pefra). Paratype, I , Puerto Fui, Valdivia, altitude 600, arch 4, 1955 (L. E. Pefra).
8 0 0 r n e t e r sM
Although it is very similar to 7'ipula (Eumirrotipula) pctaluraA l e x a n r l e r ,f r o m C h i l r t e I s l a n d , C h i l e , t h e p r e s e n t f l y i s d i s t i n c t i v ei n
h y p o p y g i a ls t n r c t u r e ,e s p e c i a l l yt h e g o n a p o p h y s easn d a p p e n d a g eo f t h eeighth-'sternite.The hypopygiumof petalura was discussedand figured
i n t h e o r i g i n a l c l e s c r i j t i o n( A g r . T e c . C l t i l e , 1 3 . N o . I : 8 - 9 , f i g s . l - 2 :I953).

Tipula (Lunatipula) lagunicolasp. n.
Belongs to the tt,rgata group; mesonotal praescuturngrayish 1'ellou',w i t h f o u r 1 ' r o o r l f i n c l i c a t e dn a r r o w h r o u ' n s t r i p e s ; a n t e n n a e r e l a t i v e l v
long, scape anrl perlicel light yellow, flagellum black; wings stronglv
velltrwecl,stigma pale brown; abclominaltergites recldishyellow with threeh l a c k e n e csl t r i p e s ; m a l e h y p o p y g i u m l a r g e , o u t e r l a t e r a l a n g l e s o f t e r g i t eb r o a c l a n d f l a t t e n e c l ,v i r t u a l l y b i c l e n t a t e ;i n n e r c l i s t i s t y l ew i t h b o t h b e a k
a n c l l o w e r b e a k h e a v i l y b l a c k e n e c l ,o u t e r b a s a l l o b e l a r g e , u n e q u a l l yhilohed; eighth sternite'with lateral angles hearing large irregular darkenecl lobes, without setae, meclian area with tu'o brushes of long recldish
vellow setae.

L c n g t h ,a b o u t | 4 . 5 - 1 5 n l n l .; w i n g , I 7 - I 8 . 5 n l n l .;, \ 1a I e .
a n t e n n a a, b o u t 5 . 3 - 5 . 5n l n r . F r o n t a l p r o l o n g a t i o no f h e a d y e l l r l u , ,r c l a t i v c l yl o n g , o n l y a l i t t l c s h o r t e r t h a n t h e r e n t a i n c l c rn, a s u st rown.Antcnnae
c l i s t i n c tp
; a l p i o b s c u r ey e l l o w ,t e r n t i n a ls e g n l e n b

232

A I e x a n c le r , T r o ' p i c a lA m e r i c a nT i 1 - . u t i 6 3(.V I I I )relativelyIong; scapeand pedicellight yellow, f lagellumblack;f lagellarsegrnents
with basal enlargements
distinct,longestverticils subequalto or slightlyexceedingthe segments.
Head lightgrayish brown, sides of vertex clearer gray; vertical tubercle
low and inconspicuous.
Pronotum brownish yellow. Mesonotalpraescutumgrayish
yellow with f our poorly indicated narrow brown stripes, theintermediatepair separatedby a broader more grayish central
line, posteriorscleritesof notum brownishgray. Pleura brownish yellow, sparsely pruinose,more distinct in the holotype.
Haltereswith stem obscureyellow, brighterat base,knob darkbrown.Legswith coxaebrownishgray; trochanters
yellow; femora
and tibiae light brown, outer tarsal segmentsdarker; claws ofmale toothed. Wings strongly yellowed, costal region slightly
more brownishyellow; stigma pale brown; veins slightly darker
brown. Macrotrichiaon outer radial veins, sparseron medial
branches,with relativelyfew on outerend of M; Cu, with relatively
numeroustrichia,includingalso a few on posteriorend of m-cu;
vein 2nd A with numeroustrichiaover most of the length.Venation: Petioleof cell Ml and m subequal.
Abdominal tergites reddish yellow with three blackened
stripes,lateralpair beginningat posteriorend of secondsegnrent,
widenedbehind through segmentf ive; sternitesand outer segments more uniformly reddish brown, outer segmentssparsely
pruinose;in the holotypesternitesthreeand f our more blackened,
evidentlycausedby discoloration;ninth tergite darker brown.
Male hypopygium(Figs. 61, 62) large, as in the tergata group.
Ninth tergite, t, with outer lateral angles broad, appearingas
flattenedvirtually bidentateblades,the irregular nrarginsnarrowly blackened,
centralemargination
extensive.
Outer dististyle,d, slender,apical dilation very small; inner style with both beakand lower beak heavily blackened,especiallythe former; outerbasal lobe large, pale brown, unequallybilobed, outer lobule
smaller and more obtuse.Phallosomeincluding two pairs ofunequal gonapophyses,
the shorter lower pair narrowed andstrongly curved outwardly,tips acute. Eighth sternite,8s, withlateral angles bearing large darkenedirregular lobes without
setae,median area with two brushesof long reddish yellow
setae. Habitat.' Mexico.
Holotype, d , La Laguna, Sierra Laguna, Baja California,
October 14, lg4l ( E. S. Ross & G. E. Bohart); California
Academyof Sciences.Paratopotype,6 , in AlexanderCollection.

StucliaEnt., vol. 12, fasc. l-4, novembro1969

233. Related speciesin the tergata group occurring in the Southwestern
United States include Tipula (Lunatipula) flavocauda Doane, T. (L-)
kirkwoodiana Alexander, T. (L.) mesotergata Alexander, T. (L.)
T. (L.\ raysmithi Alexander, T. (L.) sternata
mohavensisAlexander,'t. (L.\tergata Doane. All of these have been discussed,
Doane, andkeyed and figured in the author's recent Crane llies of California (BullCatifornia,Inslct Survey,8: l-269, I plate, 106 maps, 524 figs.; November
1967; referenceon page 33; Berkeley,California).The male hypopygiumof the present fly is distinct f rom those of all of the above named
species.

Tipula (Microtipula) subelfulta sp. n.
Allied to effutta; flagellar segmentsmore uniformly blackened;praescutum orange, the intermediatestripes clearer orange; male hypopygiumwith tergal 5lud.r narrow, blackened;lower arm of inner dististyle witha strong recurved blackened point.

i
i

i

I

I

Length,about 14-15mm.; win$, 13-13'5mm';
M a I e.
antenna,about4.4-4.5mm. Frontalprolongationof headyellow,
nasusrelativelyshort; palpi brownishyellow,the long terminal
segmentcleareryellow.Antennaewith Pt?Iittl,.lht.. segments
two or threebicolored,black,the narrow tips
y.ilo*, succeeding
yellowed,outer segmentsuniformlyblack.'Head buffy orange.
praeThoracicdorsumalmostuniformlyorange,intermediateclearerorange.Pleurayellow.
scutalstripesfaintly differentiated,
Haltereswith stem light brown, knob brownishblack. Legs withcoxae and trochantersyellow; femora brown, more yellowedbasally; tibiae brown, tarsi passinginto black; claws of male
toothed.Wings faintly tinged with brown, cells C and Sc darkbrown, prearcularf ield and stigma slightly paler brown, wingtip, especiallyin outer radial f ield, still paler brown; narrowbrown seamsover cord and outer enf of cell /sf
inconspicuous
Mzi veinsbrown.
Abdomenorange,lateralbordersof the more proximalsegblack subterminalring,mentsnarrowlydarkened;a conspicuous
includingsegmentssix to eight; hypopygiumpaler brown, dististylesycllow.Male hypopygium(Fig. 63) generallyas in effulta,
especiallyclifferingin details.Tergal blades,t, narrow,blackened,
along mesalmarginswhich are f ringed with short setae.Dististyles,d, slightly different,the lower arm of inner style with astrongslightlyrecurvedblackenedpoint.
Habitat.' Peru.
Holotype,6 , Divisoria,Cordillera Azul, Huanuco,altitude1,600nteters,August 18, 1947 (J. M. Schunke).Paratopotype,234

A I e x a n d e r, TropicalAmericanTipulidae(Vlll)

6 , altitudeI ,800 nreters,July 3l , lg47. Paratype,d , Fundo
Sinchono,
Huanuco,altitude1,500meters,
August10, lg47 (J. M.
Schunke).
Although the presentfly is closely related to Tipula (Microtipula)eflulta Alexander,of Ecuador,I considerthe two flies to be distinct in
the characterslisted in the diagnosis.
lValdiviana Alexander
V a l d i v i a n aA l e x a n d e r ; D i p t e r a P a t a g o n i a a n d S o u t h C h i l e , l : 1 7 , f i g s . l , 2 ( w i n g ) ,f i g . l l 2 ( d a n t e n n a ) , f i g . l l 3 ( d h y p o p y g i u m ) ; 1 9 2 9 ; ( t y p e e d w a r d s i n aA l e i a n d [ i ioriginal designationl.

Valdivionaincludesonly the specieslisted below.The nearest
refatives appear to be the genera Elnoretta Alexander andAcracanthaSkuse.Antennaeof nlale with scapevery long, subequalto the succeeding
four or five segments
combined;proximal
six or sevenflagellarsegmentswith lower face stronglyproduced,
rounded,without verticils, upper surface with numerousshortsetae; outer f lagellar segmentscylindrical,sinrple,with long
verticilsthat exceedthe segments.Wings ( Fig. 64) with cell
ML broadlysessile.Male hypopygiunrwith the basistyleproduced
at apex into a strong spine.Dististylesconrplicatedin structure.
List of Species
edwardsina Alexancler.
Argentina.
Alexander.
neuquenensis
Argentina, Chile.
shannoninaAlexander.
Chile.
synempora Alexander.
Argentina.

ZelandotipulaAlexander
H o l o r u s i a ( Z e l a n d o t i p u l a )A l e x a n d e r ; A n n . A l a g . N a t . H i s t . . (9) 9: 522-523; 1922;_ ( t y p - e N o v a r a e S c _ h i n e r;.t s T i p u l a ; o r i g i n a l t i e s i g n a t i o)n.
Z t , l a n d o t i p u l aV a n e - \ [ ' r i g l r t ;J o u r . n a t . H i i t . , l ; 5 + Z - 5 1 5 ,f i g s . , n r a p ; 1 9 6 7 .

The genus Zelandotipulawas describedas a subgenusof
HolorusiaLoew, being basedon a few speciesfrom New Zealand.
In the recentsignificantpaperby Vane-Wright,cliscussed
earlierunder H olorusictand I schnotome,numerousNeotropicalspecies
that had beenreferredby the presentwriter to the genusHolorusia
were transferredto Zelandotipulaand this arrangelllent
is adopted
here.The variousspeciesstronglysuggestHolorusiain antennat
structureand in venationbut differ in importantfeaturesof the
malehypopygiunr.
For a summaryof characters
the Vane-Wrightpapershould
be consulted.The structureof the male hypopygium,particularlyStudia Ent., vol. 12, fasc. l-4, novembro1969

235

of the inner dististylewith its armatureof blackenedpeglike
spinoidsetae,is of specialsignificancein the genus,particularlyin the numberand distributionof thesesetae.
In the Neotropicalregion speciesoccur as f ar north as
southernMexico, thencesouthwardto Paraguayand northernvan der
Argentina.A speciesthat was describedas microcephalasee parvicepsSpeiser,in list) from
Wulp (name preoccupied,island in the LesserAntillesremainsunrecognized.
Guadeloupe
It seemscertain that this locality as given is erroneoussinceno memberof the genusis known f rom the West Indies andthe speciesscarcelyappearsto be regional.Nothing is known
concerningthe biology of the Neotropicalspeciesbut the pupaof Zelandotipulonovaroe,in New Zealand.was mentionedbriefly
by Vane-Wright.
List of Species;
I

I

Ecuador.
acutistyla (Alexander), as Holorusia(Walker), as Tipula
Mexico, Guatemala.
a.ssocian.s
Syn.guatemalensis(Alexander),as Holorusia
Guatemala.
Colombia.
austrofurciferasp. n.
(Alexander),Bolivia.
as Holorusia
hisatraEcuador.
calvicornis (Edwards), as Prionocera
Ecuador.
calypso (Alexander), as Holorusia(Alexander), as Holorusia
Argentina.
catamarcen.si.s(Alexander),Peru.
as Holorusia
celestissima(Alexander),as
Ecuador.
Holorusia
chimborazon.
sp.
Colombia.
corynostyla
Bolivia.
cristalta (Alexander), as Holorusia
Ecuador.
cristifera (Alexander),as Holorusia
Bolivia.
cristobtusa (Alexander), as Holorusia
Ecuador.
diducta sp. n.
(eluta Schiner,see schineri Alexander)
fassliana (Alexander), as Holorusia - Bolivia.
llavicornis (Alexander),as Holorusia - Venezuela.
Peru.
flavogenualis(Alexander), as Holorusia(Alexander),Bolivia.
as Holorusia
forsteriana(Alexander),Costa
as
Holorusia
Rica.
furcifera
Brazil.
gracilipes (Walker), dS Tipula
Brazil.
hornii (Alexander), as Holorusia
Venezuela.
infernalis (Alexander), as Holorusia
juturna (Alexander),as Holorusia
Paraguay,Peru.
(Atexander),Brazil,Paraguay.
Holorusia
aslaevis-'Ecuador.
lassula(Alexander),as Holorusia
Colombia,Venezuela.
longitarsis(Macquart), as Tipula(Alexander),Bolivia..
as Holorusia
luteivena(microcephalavan der Wulp, see parvicepsSpeiser)monostictula(Alexander),as Holorusia - Ecuador,Peru.
Bolivia.
neurotrichia (Alexander), as Holorusia
Peru.
nigrosetosa(Alexander),as Holorusia
Colombia.
orophila (Alexancler),as Holorusia

236

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

parviceps(Speiser),as Tipula - LesserAntilles: Guadeloupe.
Syn. microcephala(van der Wulp, l88l) as Tipula, preoccupied
by microcephala(Bigot, 1858); vanderwulpi(Alexander,l9l2)
as Tipula.
parvimacula
Brazil.
-perobtusa (Alexander), as Holorusia
Peru.
sp. n.
(Alexander),Bolivia.
perstrangalia
as Holorusia
peruviana (Alexander),as Holorusia - Peru.
plagilera (Alexander), as Holorusia - Venezuela.
Colombia.
retrorsa sp. n.
Brazil, Paraguay.
ringens (Alexander), as Holorusia
British Guiana, Brazil.
schineri (Alexander), as Holorusia, 1934
Syn. eluta Schiner,as Tipula, 1868,preoccupiedby eluta Loew,
as Tipula, 1863).
(Alexander),Peru,
as Holorusia
sinuosa
Bolivia.
songoensis(Alexander), as Holorusia
strangalia (Alexander),as Holorusia (Fig. 65, venation) - Venezuela.
Honduras.
subfurcifera (Alexander), as Holorusia
Brazil.
sublaevis(Alexander), as Holorusia(Alexander),Paraguay.
as Holorusia
subtarda
tarda (Alexander), as Holorusia - Brazil, Paraguay.
triatra (Alexander),as Holorusia - Bolivia.
Bolivia.
trichoneura (Alexander), as Holorusia
Peru.
tuberculilera(Alexander),as Holorusia(Alexander),Peru.
as Holorusia
uniatra(Alexander),Ecuador.
as Holorusia
vividaBrazilvulpes (Alexander), as Holorusia
yungasicola(Alexander),as Holorusia - Bolivia.
zamorae (Alexander),as Holontsia - Ecuador.

Zelandotipulaaustrofurciferasp. n.
Allied to furcifera,'mesonotalpraescutumbrown, intermediatestripes
more yellowed; antennal f lagellum bicolored; wings conspicuouslypatterned with brown, inclutlinga postarculardarkeningand a major spotat midlengthof cell M; male hypopygiumwith posteriormargin of tergite
trilobed, vestiture of lateral krbes relatively sparse; branch of outerdististyleglabrous,tip obtuse; spiculesof disk of inner style relatively
few in number-

M a le.
L e n g t h ,a b o u t 1 5 m m . ; w i n g , t O m m . ; a n t e n n a ,about 3.8 mm. Frontal prolongationof head about two-thirds
the remainder,dorsalhalf obscureyellow,ventralhalf dark brown,nasus elongate; palpi with basal three segmentsdark brown,
terminal segmentobscureyellow. Antennaewith proximal threesegmentsobscureyellow, succeedingsegmentsbicolored,basal
and ventral surfaces brown, tips yellowed, the amount of thelatter decreasingon outer scgments,outer three uniformly darkened; terminalsegmentnearlyone-halfas long as the penultimate,
slender.Head light brown; anteriorvertex nearly four times thediameter of the scape.

Studia Ent., vol. 12, fasc. l-4, novembro1969

I
I

237

Pronotunt brown. Mesonotal praescutum chiefly brown,
including the usual lateral stripes, intermediatestripes moreyellowed, the capillary tnedian vitta gray; scutal lobes brown,
scutellunrand nrediotergiterllore brownish yellow; pleurotergite
ligttt yellow. Pleura
weakly clarkeneddorsally, katapleurotergite
light yellow. Halteres infuscated. Legs with all coxae andtrochantcrsclear light yellow; f enrora and tibiae brownish yellow, tips blackened,those of the latter narrower; tarsi brown,
claws of male simple. Wings with the ground light brown,
conspicuouslypatternedrvith darker brown and cream colored
areas; the dark pattern includes the prearcular f ield, cells C
and Sc, stigma,an extensivepost-arculararea in cells R and M,
a large spot at ntidlengthof cell M, and seams at origin ofRs; f urther darkeningsincludetlie f ork of M, m-cu and beyond
the corcl in bases of cells Re and Rr-,i the chief cream colored
areas are at outer end of cell R and before and beyond thedarkenedspot in cell M; veins brown, obliterativeareas conspicuous.Veins behindR essentiallyglabrous,with very sparseweaktrichia on R+-s and petiole of ceil Mr Venation: Distal sectionof Rn-, strongly sinuous,narrorvingcell Rs; petiole of cell M I
nearly twice m; m-cu beyond nridlengthof Ms-q.
Abclorninaltergites medium brown, basal ring of second
segmentmore yellowed, sternitesyellow; subternlinalsegntents
dark brown to form a narrow ring; hypopygiumyellowed.Malehypopygium ( Fig. 66) with the tergite, t, trilobed, the central
lateral lobes longer with relativelysparse
projectionconspicuous,vestiture.Outer dististyle,d, elongate,relativelyslender,setiferous,
the conspicuousbranch at midlength of inner f ace glabrous,
tip obtuse; inner style with blackenedspinoid setaeof disk much
fewer than in either furcifera or subfurcifera.
Habitat; Colombia.
Holotype, 6 , Monteredondo,Cundinamarca'altitude 1,42O
meters,January5, 1959 (Juan Foerster).
Other speciesthat have the outer dististyleof the male hypopygittm
furcate include Zelandotipula furcilera (Alexander) and Z. subfurcifera(Alexander), both differing from the fly in hypoPYgialstructure, 8s
discussecl.

Zelandotipulacorynostylasp. n.
Mesonotalpraescutumlight brown with four more cinnamonbrown
stripesthat are narrowly marginedwith darker brown; posteriorsclerites
of notum and the pleura yellow; wings light brown, prearcularand

238

A I e x a n d e r , T r o p i c a l A m e r i c a nT i p u l i d a e ( V I l l )costalfieldsdarker,a small brown spot in cell M; abdomen
reddishbrown,
subterminal
segments
black; male hypopygiumwith posteriorface of eachtergal lobe long-oval,with abundant blackenetlspinoid setae; innerclististylewith outer plate broacllysubtriangular,
clenselyset with small
blackenedspinoidsetae.
M a I e.
Length, about l6- l7 mnl.; wing, l6- l7 mm.;
antenna,about 3.5-3.7 mnl. Frontal prolongation of head yellow-

ish brown, nasus concolorous,long and slender; palpi black,terminalsegmentbrownishyellow.Antennaerelativelyshort; scapeclongate, brownish yellow, pedicel and first flagellar segment
light yellow, f lagellar segmentstwo to eight bicolored,brownbasally and along lower f ace, outer end yellowed, outer threesegmentsbrownish black; f lagellar segmentsrelativelyshort, theintermediateones about twice as long as broad, terminalsegment
slender,about one-thirdthe penultimate.
Head brown,very vaguely
patternedwith paler.
Pronotuntlight brown nredially,paline to yellow on sides.
Mesonotalpraescutunr
light brown with four scarcelydifferentiatedmore cinnantonbrown stripes that are best indicatedby narrowdarker brown ntargins; scuturn alnrost unifornrly brown, withclelicateyellow setae; scutellurnand postnoturnyellow. Pleura,
including the dorsopleural nrembrane,yellow. Halteres darkbrown, base of stem narrowly yellowed. Legs with coxae andtrochantersyellow; fentora and tibiae brownishyellclw,tips narrowly darker brown; tarsi light brown; claws of nrale bispinous,
outer spine obtuse. Wings light brown, paler in outer end ofcell R, nluch of M, and basesof cubital and anal cells; prearcular
anclcostalregions,with the stigma,brown; two very small brownspots, at base ancl at near nridlenethof ccll M adjoining vein
C u ; v e i n s b r o w n . V e i n s b e h i n d v e i n R g l a b r o u s .V e n a t i o n : R srelativelylong, about trvo ancl one-halftirnesRr-*; cell Rr modc'ratelyconstrictedat near rnicllength;nt-cu at near two-thirds
Mv+.
Basal abdorninaltersitesrecldishbrown, sidesslightly darker,
sternitesand hypopygiurnyellowed, segnrentsseven ancl eight
blackenedto form a ring, tergitesfive and six narrowly darkened
medially.Male hypopygiunr(Fig. 67) with tergite, t, as viewed
from above,terminatingin two low transverselobes lvith slendersetae, the posterior faces long-oval, with abunclantblackened
spinoid setae. Outer dististyle,d, slender,narrowed outwardly,
apex glabrous,obtuse; inner style with outer plate broadly subtriangular in outline, with dense snrall blackenedspinoid setae,
lower blade rvith relativelyfew nrarginalpoints.

StudiaEnt., vol. 12, fasc. l-4, novembrol969

239

.--.=
-----..*\t--__---\

_
-

69

flg. 6l. Valdivia shannonina Alexand^er;'. venation, - Fig. BS. Zelandotiputa strangatia(Alexander); venation.
Fig.. .OO. Zetandotiputa
dusiitiiifer"'ni'ale- sp. n.; maleig, 67. zetandotipura cor1,:noit1,ta
rp. n.;
lypopygium.
rriFopvgiirm. .f
Fie. 68. Zetandotiput_ldllucta ?p. h.; Tlte Jrlnirpvsirilo,I Fii.- og."i'niiiiol'iputa
perobtusa-sp. n.;. male hypopygium.
_ . . F i . g -z o . ' 2 e f t i a o t i p u t a - i F i-r o r r o s p . n . ; m a l ehypopygium.
s y r i r b o r s :d , t r i s t i i t y r e s ; p , p h a i l 6 s o m"i t, tergite.

Habitat; Colombia.
Holotype,6 , Monteredondo,
cundinamarca,
altitude 1,42ometers,January-Februaryt g5g (Juan Foerster).
.

From the other relatively numerous speciesof the genus that havea small darkened brown spot at near midlength of cell M of the wings,
the present fly is best distinguishedby the length ancl coloration of themale antennaeancl especiallyby the hypopygial structure,including thetergite ancl both clististyles.

240

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

Zelandotipuladiducta sp. n.
i V l e s o n o t apl r a e s c u t u nct i n n a m o nb r o w n , w i t h f o u r v a g u e l y d i f f e r e n t iated yellowishbrown stripes,pleura yellowed;antennaeof male relatively
long, exceedingone-fourththe 'wing, outer sevenor eight segmentsblackened; femora yellow, tips narrowly darker brown; wings fulvous brown,r e s t r i c t e r l l yp a t t e r n e dw i t h d a r k e r b r o w n ; a b d o m i n a lt e r g i t e sl i g h t b r o w n ,s t e r n i t e sa n d h y p o p y g i u ml i g h t y e l l r r w ;m a l e h y p o p y g i u m' w i t h t h e t e r g i t ebroad, outer lobes obtuse, with black setae, the broad emarginationn e a r l y t r u n c a t e ,u ' i t h s p a r s es e t a e t h a t a r e l a c k i n g a t t h e m i d l i n e ; o u t e r
d i s t i s t y l ep a l e , g e n t t y c u r v e c l ,a p e x w i t h f i v e o r s i x u n e q u a lp o i n t s .

L e n g t h ,a b o u t 1 5 -1 6 n u n . ; w i n g , a b o u t2 l - 2 1 . 5
M a I e.
m m . ; a n t e n n aa, b o u t5 . 7 - 5 . 8m m .
F e nt a I e.

L e n g t h ,a b r : u t2 l l t ' l n l . ;w i n g , 2 3 n l n l .

Frontal prolongation of hcad light brown, sliglttly nlore
yellowed at base above; nasus long and slender,apex narrowly
b l a c k e n e dp
; a l p i b l a c k , t h i r d s e g n r e n st l i g h t l y p a l e r . A n t e n n a eo t
m a l e r e l a t i v e l yl o n g , e x c e e d i n go n e - f o u r t l tt h c w i n g ; s c a p e a n dpedicel obscureyeilo'uv,the fontter shrtrt ancl stttut, sliehtly lcss
t h a n t r v i c e t h e l e n g t h o f t h e l a t t e r ; p r o x i n r a lf l a g e l l a r s e g n t e n t s
obscure1lellour,thc outcr oncs brownishblack; segmentsgradually decreasingin length outwardly,terminalsegmentvery reduced,
slender, about one-sixth the penultimate.Antennae of fentale
shorter. Heacl above chestnut brown, with a capillary yellow
c e n t r a ll i n e .
Pronotulttttarrowly dark brown nrediallyand on sides, theinterveningspacc light 1'ellotv.Mesonotal praescutunrcinnanron
brown with f our vagucly diff erentiatedyellowish brown stripes,
Itunteral and lateral bordcrs stlnrcwhatdarkcr brorvn; scutumd a r k b r o w n , s c u t c l l u n rev e n d a r k c r ; r n e d i o t e r g i t gerayishbrown,
posterior f ourth yellowed, n'ith a capillary central brown vitta;
pleurotergitelight brown with a snrall darker brown discal spot;
scutellumand nrediotergite
with conspicuouspale setae,the latter
porrect. Pleura yellowed, with a short brown dorsal stripe on
propleura and dorsal anepisternum;dorsopleuralregion conspicuously light yellorv. Halteres with stem light brown, base ofknob darker, apex vaguely pale. Legs with coxae and trochanters
yellow; femora light brown, tips narrowly darker brown, involving
about one-sixth of the segnrent; tibiae brownish black, tarsi
brownishyellow; claws of male bidentate.Wings strongly fulvous

patternedwith darkerbrown,includinga postbrown,restrictedly
arcular area in basesof cells R and M, and a smallerspot at
origin of Rs,'stigrnalarge,brown;cell C yellow,outerend weakly

Studia Ent., vol. 12, fasc. l-4, novembro1969

241

darkened,cell R1 before stigma extensivelylight yellow; very
snlall and indistinctyellowed areas near bases of cells M and2nd M2 and at rnarginof /s/ A; veins light brown, basal fourthof R.sdarker.Veins behindR glabrous,Rn-s with very few microscopicpunctures.Venation:Cell R3 moderatelyconstricted;petioleof cell Ml subequalto m; m-cu at from about one-halfto threefourths Mz-+; cell 2nd A relativelynarrow.
Abdomen relativelyshort in both sexes, especiallyin themale; tergites light brown, incisuresof intermediatesegments
narrowly yellowed, sternitesand hypopygium light yellow. Ovipositor with cerci slender,straight, narrowed gradually to thesubacutetips. Male hypopygium ( Fig. 68) with the tergite, t,slightly narrowed outwardly, outer angles produced into low
obtuselobesthat are denselyprovidedwith black setae,the broadinterveningenrarginationnearly truncate,with sparsesetae thatarc lacking at and near the midline. Outer dististyle,d, pale,nroderatelybroad, narrower than in lassulo,gently curved, apex
truncated,with five or six unequal pale points to produce anerose appearance;innc'r style with apex broadly obtuse, theouter hoodlikepart with snrallblackenedspinoidsetaethat extend
to the summit, lnore basally with two other nearly contiguouss n r a l l e ra r e a sw i t h s i n r i l a rs p i c u l e s .
H abitat.' Ecuador.
Holotype, d , san Francisco, Rio pastaza, artitude I ,300
rneters,October l, 1938 (Wm. Clarke Macintyre).Allotopotype,
9 , Paratopotype,d , pinned with the allotype.
Zelandotipuladiducta is generally similar to Z. lassula (Alexancler)
ancl to Z. triatra (Alexander),differing in the length and structure of
the male antennaeancl especiallyin the hypopygium,includingthe tergite
ancl both dististyles.The ninth tergite in all three speciesis distirictive,in lassulabeing trilobed, all lobes bearing conspicuoussetae; in triatra
with only two lobes,as in the presentfly, thesemuch larger and separated
bv a narrower emargination.

Zelandotipulaperobtusa sp. n.
General coloration brown, praescutum 'with four brownish yellow
stripes that are marginecl by slightly darker brown, pleura yellow; tipsof femora clark brown; wings pale brown, heavily patterned with darker
bror+'nanrl whiteneclareas; male hypopygium with beak of inner dististylet?ry" short ancl stout; gonapophysis unusually broacl, the tip incurved.
t6

212

A I e x a n c le r , T r o p i c a l A m e r i c a nT i p u l i d a e ( V I I I )L e n g t h ,a b o u t 1 6 l l l m . ; w i n g , 1 9 n l m . ; a n t e n n a ,M a I e.
about 4.8 mnl. Frontal prolongationof head above dark brown,
includingthe long nasus,the base beneathmore yellowed; palpi
clarkbrown, terminalsegmentobscureyellow.Antennaemoderatety long; proximal three segnrentsyellow, succeedingsegntents
weakly bicolored,brown basally, the outer half obscureyellow,
the anlount of the latter decreasingon outer segments,the outer
three or f our uniformly darkened;terminalsegmentslender,morethan one-fourththe penultimate.Head light brown, with a narrow
clarkerbrown central line, posterior orbits very narrowly gray.
Pronotunrmediunrbrown. Mesonotalpraescututttwith groundlight brown, clisk with four brownish yellow stripes that are
rnargined with slightly darker brown; scutunt and scutellum
nrediurrrbrown; postnotuntpaler brown, anterior pleurotergite
and pleura light yellow. Halteresbrown, base of stem light yellow. Legs with coxae and trochantersyellow; femora and tibiae
obscureyellow,tips of fenrorarather narrowly dark brown, of thetibiae more narrowlyso; tarsi brown, claws bidentate,outer toothlarger. Wings with ground pale brown, heavily patternedwith
clarkerbrown and whitened areas,generallyas in strangalia andallies; cells C and Sc unifornrlybrown; further major darkenings
in basesof cells R and M, at midlengthof cell M, and in outerraclialf ield with the exceptionof cell Rsi nlore restrictedbrown
seamsat origin of Rs and m-cu; the pale pattern includesa prestignralbrightening,major areasbeforeand beyondthe darkening
in cell M, and most of cetl R; i veins brown. venation: vein
R*-r, rather strongly sinuous,constrictingcell R:li m-cu at nearn.,i,ll.ngt5of My+i cells Ma and 2ntt A broaderthan in acutistyla.
Abdonrinaltergite'slight brown, sliehtlypatternedwith darker
brown on sides, outer segtnentsnlore extensivelydarkened,sternitesand hypopygiurnyellowecl.Male hypopygium( Fig. 69)
with the tergite,r, trilobed,nredianlobe shorter,pale and glabrous,
apex truncated,lateral lobes triangular,their mesal marginswith
abunclantshort subcultrateblackenedspinoidsetae.Outer dististyle,styled, slender,outer third very narrow' apex subacute;inner
broad, beak very short and stout, disk with three groups of small
subblackenedspiculesor modifieclspinoid setae,outer groupwith
circular in outline, the points very numerous.Phallosonleunusuallybroad, apicesincurved.
gonapophyses
Habitat.' Peru.
Holotype, 6 , satipo, Jauja, Junin, altitude 800-900 meters,
June24, 1940 (P. PaPrzYcki).

Studia Ent., vol. lZ, fasc. l-4, novembrot969

243

The most similar species Zerandotiputaacutistl,ls(Arexander),of.is
Ecuador,which differs in slightdetailsnf wing coloiaiionand venation
ancJespeciallyin .the hypopygialstructure,incl"uding
the inner oististylephallosome.
andOther species,as Z. per.strangalia(Alexander)and Z.
.strangalia(Alexander)are less ckrselvallied.

Zelandotipularetrorsa sp. n.
Mesonotalpraesc'utum
tight brown with four brownishyellow
stripes,
pleura testaceousyeilo'w; uit.nnu.
-rhori; femora yeilrw, tips brownish
black; wings with disk whiteneJ, ;.r;ir;
patternecrwith dark brown,veins glabrous,
strongly.
constyi.t.,ri
ior border of ,ttll .fs prodticecl,rrrulru*ry' mare 'ypopygium with posteremarginate,clenselyset with.tergite"short blackened
cuttrati bristlei; outer ,t'i.ti.tyd u.t'
bloao, the length
about one-halfgreater than the'brealil,l'"inn.,disk with two longituctinarrows of microscopicstyle with beak slenderpegs or spinoid setae.

Male.

Length, about I 5- l6 mm.; wing 20-21
nrm.;,
antenna,about 2.4-2.5 mm.
F e m , I ..
L e n g t h ,a b o u t 2 2 m n r . ; w i n g , 2 3 n r m . ;antenfra, about 2.3 mnl.
Frontal prolongationof heacl shorter than
the remaincler,
dark cinnamonbrown; nasus very rong ancr
srencrer,exceeding
one-thirdthe prolongation;proxinralthree palpat
segnrents
black,
the elongateterminalone clarkenedbasally,
ort.l. two-thirclstightyellow' Antennaerelativelyshort; scapeand
pedicelobscureyetlow, f lagellumbrown,outwarcilypassinginto
dark brown; terminarsegmentabout one-thircllonger than the penultimate.
Head darkbrown, vertex with a capillarydarker central
line, front and narrow orbits vaguely paler.
Pronotumbrown medially, broacllytestaceouson sides.MesoItotal praescutuntIight brown with f our lightbrownish yellow
stripes that are very narrowry marginedby
darker; scutar robes
brownish yellow, patterned with biown; scuteilunr
obscureyer_
low; postnotumbrown, centrar area of preurotergite
more y.rlowed. Pleura testaceousyeilow, anepisternum
anct adjoining
dorsopleuralregion weakly infuscated.Halteresdark brown,
baseof stenr yellowed. Legs with coxae and trochanters
yeilow;renrainderof legs very long ancl slender; fenrora
yellow, withsreenishtints, tips brownishbrack; tibiae tight brown, tips
nar_
rowlv more darkenecr;tarsi brown, craws sirnpte.
wings withdisk whitened,including the prearcularf ielcl, heavily
pitternecl
with dark brown, Anal cells chiefly Iight brown, bases
whitened;brown areasincludea postarcularctarkeningthat entirelycrosses
the wing, more extensivein bases of cells R and
M; a second
l6f

214

A I e x a n d e r, Tropical AmericanTipulidae (VIII)

arca at ncar two-thirdsthe lengthsof cells R and M, not connectedwith the basal band; other extensivedarkenedmarkingsat origin of Rs, cord, expandedoutwardlyinto the radial cells
and basallyinto cell M; outer radial field extensively
darkened,leavingthe centersof the cells and all but base of cell Rr
paler; cell C uniformlylight yellow; stigmadarkerbrown; veinsdark brown, in vicinity of the stigmawith greenishtints. Veins
punctures.
behindR glabrous,Rn-u with very few microscopic
Venation:Outer sectionof Rn-u stronglysinuous,narrowingcell
Rs at near midlength;cell /sf M2 small,pentagonal;nt-cu close
to fork of Me-+.
Abdomenof male relativelyshort; tergitesbrownishyellow,
patternedwith darker brown, sternitesyellow; segnlentssix toeight blackenedto f ornr a conspicuousring; extrenlebases ofhypopygiurtt
segmentspale,not apparentin unnlountedspecimens;
small,brownishyellow.Malc hypopygiunr(FiS. 70) with posteriorborder of tergite, t, broadly producedinto a lorv lobe, its bordershallowly emarginate,denselyset with short blackenedcultrate
bristles,those back fronr nlargin longer and more norntal.Outcr
dististyle,d, pale, very broad, the length about one-half greaterthan the breadth; inner style about as f igured, beak slender,tip
obtuse;dorsal crest erect,with severalsetae; disk of style with
two separateareas of nricroscopicblackenedpegs or spinoid
setae,outer group extendedltackward along the lower marginof a slenderextension.
Habitat; Colombia.

Holotype,6 , Monterc'-londo,
Cundinanrarca,
altitude 1,420
nleters, January 10, 1959 (Juan Foerster). Allotopotype, ? ,d , DecemberI l, 1958.
January30, 1959.Paratopotype,

Other generally similar regional speciesof Zelandotipulathat havethe samegeneraltype of wing patternand 'rvithcell R, stronglyconstricted
at near midlength,includeZ. acutistyla(Alexander),Z. crislifera (Alexander), Z. infernalis (Alexander), Z. sinuosa (Alexancler),Z. strangalia(Alexander),Z. vivida (Alexander),and some others. All of these cliffer
among themselveschiefly in hypopygial structure, incluclingthe tergite
and both dististyles.The very broad outer style of the present fly is
distinctive.