70 B
29

Notes on the Tipulidoe of Ecuodor(Order Diptera)Part IIIBy (hrb

P. Atcnndcr U*"-*yotMrffinmrUfnre+m.
The precedingarticle under this generaltitle $as publishedin Aprtl 1953(Rev. Ecuat.Ent. Par. I Q)z 6 - 15, figs. I - 8).
In the ltresentpaper I wish to discussthe local speciesof the genus
Ef&fnagma Osten Sacken,one of the most characteristicgeneraof
Tlptfldae'in the Neottopics.
Epiphngnr Osrcn Sacken
Limnophila @piphragma)Osten Sacken;Proe. Acad. Nat.
Sci. Philadelphiafor 1859: 238, 1859; (typelascipennis
San as pavonina Osten Sacken)' 1 t' EpiphragmaOutonSacken;Mon. Dipt. N. Amoriqu 4: 193194,1869.
'ii ,Epiphgma Abrander; Rev. dsEntomologie,19; 168 - 175,
fig. 5 (venation),1948.
"ttti.ttt;() ,.

. . r, rRev. Ecurl. Ent. Per. f €) tr[o 1953

30
The genusEpiphragmaincludesflies of mediumsize,invariably
with a brownpatternon the wing, usuallyin the form of ocelli or annuli, thesefrequentlybroken into abundantspotsand dots. The genus
is characterized
by the presenceof a strongcrossveinin cell C of thewing (Fig. 1); in the Oriental- Australasiansubgenus
Eupolyphragma Alexander,with approximately 25 speciesrangingfrom the Philippines southeastward
to New Guinea, there are severalsuch crossveinsin this cell of the wing.
\
The typical subgenusis wide-spreadthroughoutthe Holarctic
Regionbut here with relativelyfew species. In Tropical Americait
by u hostof forms,of whichno fewerthan 14 havebeen
is represented
found in Ecuador and are discussedin this report. A further concentrationof speciesis found in the Oriental-AustralasianRegion,includingseveralspeciesin Australia. No membersof the genushave
been found in the Ethiopian Region,includingMadagascar,nor in
New Zealand. In theselatter areasand elsewherein the World, thoughprincipally in the SouthernHemisphere,the genusis replacedby theevenlarger and more diversifiedgroup AustrolimnophilaAlexander.
The immaturestagesof theseprimitive HexatomineTipulidae,
including both Epihragmaand Austrolimnophila,occur in wet decaying wood,sometimes
temporarilysubnrerged. Variousspeciesof Epiphragmahave been reared from such a habitat in Europe (ocellaris
Linnaeus),North America(iascipennisSay and solatrixOstenSacken),and in Argentinaand Uruguay(imitansAlexander).
Key to the Species
Legs with the femora and the broad tibial bases black,
the rernainderof legs yellow.
Legs coloredotherwise.

2.

Wings pale yellow, with all cells abundantlydotted withbrown.
hirtistylataAlexander
Wings pale, crossbandedwith darker but without isolated
brown dots in the cells.
varia (Wiedemann)

3.

Mesonotal praescutumobscue yellow, with two maior
polishedblack areason either side.

2
3

31
nigroplagiataAlexander
Mesonotalpraescutumnot paternedas above.

4

4.

Wingswith a singledarkenedareain cell 2nd A, additional
to the markingsat baseand apexof cell.
subsolatrixAlexander
Wingswith two or moredark marginalspotsin cell 2nd A. 5

5.

Antennae (male) moderately elongate, approximately onethird the length of wing.
Antennae (male) shorter, approximately one-fourth thelength of wing or less.

6.

Antennaewith scapeand pedicel pale brown, the flagellar
verticils small and inconspicuous;no isolateddark costal
spot basadof the annulusat origin of Rs; dark spots incell 2nd A rounded or oval in outline.
parviseta Alexander
Antennae with scape and pedicel pale brown, the flagellar
verticils long and conspicuous;an isolated dark spot incells C and Sc basad of the annulus at origin of Rs;
dark spots in cell 2nd A elongate,narrow, crossingthe
cell.
filiformis Alexander

7.

Tibiae uniformly vellow.
Tibiae in large part darkened or blackenned.

8.

Pleura uniform in color; antennal scape and pedicelgblackened.
Pleura striped longitudinallywith vellow and black; antennal
scape pedicel yellow or onlv slightly infusticated
.
l0

9.

Femora ringed with black: ground areasof wing uniformly
pale.
subenixaAlexander
Femora virtually unpatterned:ground areasof wing dusky,
narrowly marginedrvith yellow.
interspersasp. n .

10.

Cell Cu of wingswith a seriesof darkenedareas.
Cell Cu with the darkenedareassparse,usually only one
or two near outer end.

8
l3

11

t2

32
11.

Femorawith two narrowclearlydefineddark rings;halteresyellow,the baseof knob darkened.
enixa Alexander
Femorawith a singlenarow brown subterminalring; halteres infuscated.
oxyphallus Alexander

12.

Size larger (rving, male, 12 mm.); halterespale, the baseof knob infuscated;darkenedannuli of wing at arculus
and at origin of Rs widely separated.
diademaAlexander
Size small (rving, male, 7 .5 mm .); halteres yellow;
darkened annuli of wing at arculus and at origin of Rs
phaeoxatha Alexander
approximated.

13.

Terminal segmentof antennayellow: thorax dull black,
including the mesonotum;ground color of wings, especiallv in the basalcells,weaklvinfuscated,cleareryellow
inmediatelyadjoiningthe darkenedareas.
deliberataAlexander
Antennae black, only the first or fusion segment offlagellum orange;mesonotumlight brown, pleura black;ground color of win.gsclear light yellow.
xanthomelaAlexander

Certain of the species,especiallyhirtistylata and oxyphalus,
show distinct hypopygialcharacters. Others have thesestructuresmore uniform and are evidently closelv allied. The great majority ofthe specieshere consideredwere taken at and near Abitagua, NapoPastaza,which thus showsa surprisingconcentrationof speciesin thisgenus. There can be little question but the many further membersof the group remain to be discovered.
As was indicated in the preceding part, the various species
consideredthroughout the entire series of papers will be numberedconsecutively,the present treatment including species 1? to 26, inclusive.
Epiphragma (Epiphragma) deliberets Alexander, 1939.
Abitagua, altitude 1200 meters, April to September 1937
(Macintyre); types.
13 .

33
Epiphragma (Epiphragma)disdema Alexander, 1939.
Abitagua, altitude 1200 meters, April to September1937(Macintyre); types.
14.

Epiphragma @piphragma) enixa Alexander, 1939Abitagua, aititude 1200 meters, April to September1937(Macintyre);types.
15.

Epiphragma (Epiphragms) filifonnis Alexander, 1940.
San Francisco, Rio Pastaza,altitude 1300 meters, October 1.,
1938 (Macintyre);type.

16.

Antenna (Fig. 2) l6-segmented; basal segmentof flagellum
aboui two-thirds the second segmentand a little stouter; succeeding
segmentslong<ylindrical, with long conspicuousverticils, the longest(on the intermediatesegments)up to two-thirds the length of the segment; terminal segmentabout two-thirds as long as the penultimate.
The figure showsthe pedicel and first four flagellar segmentsof type.
Male hypopygium(Fig. 3) with the tergal lobes, 9t, unusuallysclerotized,raddish brown, relatively broad, their apices obliquely
truncated;lobesseparatedby u U-shaped nocht. Interbase,i, with
its lobe slender,the apex a long recurvedcrooklike spine. Outer dististyle, d, small, abruptly terminatingin a decurvedapical spine, theouter margin near base with a low setuliferousswelling. Aedeagus,
a, unusuallylong and sclerotized.
Epiphragma (Epiphragma) hirtistylata Alexander, t939 .
Abitagua, altitude 1200 meters, April to September 1937
(Macintyre);types.
17.

Male hypopygium(Fig. 4) with the 9th tergite,9t, producedinto
iwo very low obtuselobesthat are provided with very delicatesetulae;
surfaceof tergal plate back from the margin with a transverserow ofsetae,the more central ones smaller,the outer ones strong and powerful. Basistyle,b, stout, the outer face of the distal half with unusually long and powerful setae,the longest only a little shorter than thestyle itself; mesal face of style on proximal half with a further row ofstrong setae,the outer ones smaller and more slender. Interbase,i,with the lobe a slender spine, the apex produced into an acute deflec-

34
ted point. outer dististyle, d, pale, terminating in an acute curved
spine; surfaceof style with long conspicuoussetae,those of the disk
longer, deflectedventrad and mesad; inner dististyle a little longer,
obtuseat tip, with setaeon the lower margin and with a small concentrationon the disk. Aedeagus,a, small and weak, approximately
equal in sizeto the lobe of the interbase.
18.

Epiphragna (Epiphragma)intercpersasp. n .

size medium(wing,male,over l l mm. ); mesonotalpraescutumwith the sidesdarkened,the disk with confluentpaler brown stripes;
antennalscaPeandpedicelblack,the fusionsegment
light yellow;femora brownishyellow, the tip clearer yellow; wings pale brown, crossbandedwith darker brown, the bandsneurowlymarginedby light yellorv; male hypopygiumwith the posteriorborder of the ninth tergite
producedinto two small submediansetuliferouslobes;outer distisiyle
curvedat apexinto an acutepoint; aedeagus
small and simple,shorierthan the lobe of the interbase.
Male.- Lengthabout 100 mm. ; wing I 1.5 mm.; anrenna
about2.2 mm.
Rostrum and palpi dark brown. Antennaewith scapeand
pedicelblack, the former pruinose;fusion segmentof flagellumlightyellow, the remainderof organ black; fusion segmentinvolving two
segments,
the remainingflagellarsegmentswith long conspicuous
verticils. Head fulvous brown, the vertex with a narrow darker medianvitta.
Pronotum brown medially, thc anterior part more yellowed.
Mesonotalpraescutumwith the sidesextensivelydark brown, the disk
occupiedby confluentpaler brown stripes;scutumreddishbrown, thecentral area darker; scutellumand postnotumsomewhatmore pruinose. Pleuraalmostuniformly dark brown. Halteresobscureyellow,
the baseof knob darker. I*gr with the coxaedark brown; trochanters
yellow; femorabrownishyellow,the tip palingto cleareryellow; tibiae
and tarsi yellorv. Wings (Fig. l) with the ground pale brown, withmore extensivedarker brown areas,appearingas almost entire crossbands, the paler ground areas narrowly bordered by pale yellow
marginsthat separatethem from the darkenedcrossbands;costal re-35gion with the brown bands still darker, becoming paler behind, thethird such area lying at the supernumerarycrossveinin cell C; cell
2nd A with about six dark areasthat are more extensivethe pale interspaces;
veinsobscureyellow, somewhatdarker in the patternedareas,
dark brown in the costal darkenings. Venation: Cells R2 subequalin extent at margin; cell R2 long and narrow; m-cu at near one-third
the length of cell lst M2.
Abdomen chiefly dark brown, the basal sternites more yelowed; hypopygium brownish yellow. Male hypopygium (Fig. 5)
with the posteriorborder of the 9th tergite,9t, transverse,producedinto
two small submedianlobes, their surface with rnicroscopic setulae.
Lobe of the interbase,i, unusuallv slender, its iip broken in type.
Outer dististyle,d, simple, curved at apex into an acute point, thesurface and especiallythe enlargedbase densely setuliferous,with a
few larger setae;inner dististyle longer, flattened, the apex obtuse.
Aedeagus,a, small and simDle,shorter than the lobe of the interbase.
i{olot''pe, nrale,Abltar:u", Napo-Pastaza,
altituCe 1800 meters, April 15, 1940 (Macintyre).
This fly is most similar in its wing pattern to Epiphragma(Epiphragma)persanctaAlexander (Venezuela,Brazil), differing evidently the virtually unpatterned legs. The present fly is apparently
the speciesthat I had called E. (E.) fabricii Alexander(Proc. U. S.
Nat. Mus., 44: 537 - 538, pl. 68, fig. 33 (wing), 1913),a re-naming of the preocupied. E . (E'.) maculata (Fabricius).
It now
appearsthat the Fabrician speciesis not the same as the present fly
and the latter has remained undescribedto this time. The identityof the Fabrician speciesstill remains in question and his identity canbe settledonly by an examinaiion of the type. Concerningthe present species,I am uncertain at this time as to whether the Ecuadoreanmaterial is identical with other similar materials from various partsof South America and due to this uncertainty prefer to restrict thedescription to the actual holotype specimen.
19 .

Epiphragma (Eprphragme) nigroptagiata Alexander,

1939.

Abitagua, altitude 1200 meters, March to September 1937
(Macintyre); types.

36
Male hypopygium (Fig. 6) with the basistyle, b, provided
rvith setaeof normal length. Interbase,i, with its lobe unusually
slender,at apex bent at a right angle into an acute point. Outerdististyie,d, slender,straight,at apex curvedinto a long acutepoint;
surfacewith pale inconspicuous
setae. Inner dististylea little longer, narrowestnear base,thenceslightly expanded,the apex obtusely rounded. Aedeagus,a, very small and slender,about one-halfas
long as the lobe of the interbase.
20 .

Epiphragma (Epiphragma)oxytrlhallusAlexander, 1939.

Abitagua, 1200 to 1800 meters, March to September 1937
(Macintyre);types.
Male hypopygium(Fig. 9) unusuallydistinct within the genus. Ninth tergite,9t, rvith the posteriorborderproducedinto trvo slender submedianlobesthat are separated
by a U-shaped notch,the lcbes glabrouswith narrowly obtusetips. Basistyle,b, with surface
very denselysetuliferous,with a relativelyfew long scatteredsetae,
smallerbut more abundantalong mesalface near apex, again longer
and more abundantnear base. lnterbase,i, slender,at apex narrorvedinto a recurvedspine or hook. Outer dististyle,d, broadest
just beyondbase,the outer basalangleproducedinto a small curved
hook; beyondthis point the style narrowsgraduallyinto a long gently curvedterminal spine;inner dististylelarger,nearly parallel-sided,p, very distinctive,appearing
very broadlyobtuseat tip. Phallosome,as a con'rpressed
flattenedblade that narrowsinto a long slenderterminal point, the entire structnrein a positionof rest decurved;at baseof phallosomervith a flattenedplate with its margin microscopically
serrulate.
Epiphragma (Epiphragma)parvisetaAlexander, l94l .
Mount Tungurahua,Tungurahua,altitude 2600 meters, april
1939 (Macintyre).

2l .

basalflaAntenna(Fig. 7) apparentlyonly 15- segmented;
gellar segmentabout three times as long as thick and not appearing
to representa fusion element;succeedingsegmentslong-cylindrical,
sevenor eight times as long as thick, with a denseerect pubescenceand a few slightly longer scatteredverticils, the longestof these about

37
one-third the segment. The figure showsthe pedicel and the first
three flagellar segmentsof type.
Male hypopygium @ig. 8) with the tergal lobes, 9t, relatively
naxrow, obtuse at tips, separatedby a U-shaped notch. Setae ofouter face of basistyle,b, strong but not unusually abundanteor conspicuous setae;inner dististylelonger, obtuse at tip, with small weak
the apex a long recurved spine. Outer dististyle,d, pale, gradually
narrowed and curved into a long spine; surfacewith small pale incons:
picuous setae;inner dististylelonger, obtuse at tip, with small weakmarginal setae,especiallyat apex, and a small group of longer setaeon disk near base. Aedeagusrelatively long, only a little shorter
than the lobe of the interbase.
22.

Epiphragma@piphragma)phaeoxanthaAlexander, 1944.

Cunibunda,Abitagua, altitude 1100 meters,March 1940 (Macintyre). The hitherto unknown female was taken above Abitagua,
arrrtude1800 meters,April 15, 1940, by Macintyre, and is here cha
racterized as allotype.
Female.- Length about 9 mm.; wing 9 mm.
As was the case rvith the holotype male, the outermost flagellar segmentsof the antennaeare broken and it cannot be stated as towhether the organ is uniformly blackened or possibly pale at op€x,
as in some membersof the genus. Fusion segmentwhitened, paler
than the scapeor pedicel; flagellar segmentslong and slender,the entire organ fully as long as in the male. Darkenedfemoral rings relatively pale and narrow, narrower than the outer yellow rings. Wings
colored much as in the male but with no darkened spot in cell Cu
connectingthe ocellatering at origin of Rs with the solidly darkened
spot at end of vein 2nd A. Venation: Cell lst M2 longer and relatively narrower than in male, with m-cu more than its orvn length beyond
the fork of M. Abdominal tergitesdark brown, paler laterally; sternites almost uniformly light yellorv,the extremelateral parts vaguelydarkened. Genital shlelCfulvousyellow; cerci slender,upcurveCto theacutetips.
2 3 . Epiplragrna (Epiphra$na) subenixa Alexander, 1939.

38
Abitagua, altitude 1200 meters, September 1937 (Macintyre);
Type.
24.

Epiphragma @pipnragns) subsolatrix Alexander, 1939.

Abitagua, altitude L2O0meters,August to November 1937 (Macintyre); tlpes.
25 .

Epiphragna @piphragma)varia (Wiedemann,1828).

At present known form Panama,Ecuador, Peru, Brazil and
Venezuela.
26.

Epiphragma @piphraglra) xanthomelaAlexander, 1939.

Abitagua, altitude l2O0 meters,April lg37 (Macintyre);type.
Still known only from the unique t-vpefemale. As indicatedat timeof descriptionof the species,it is probablethat the male of this species
will be found to havethe antennaemore or lesslengthened.
EXPLANATIONOF PLATE

Fig. 1.

Epiphragma@piphragma)interspersasp. n.; venation.

Fig. 2 .

Eplphragma(Epiphragma)filiformis Alexnnder;antenna.

Fig. 3 .

Epiphragma @piphragma) filiformis Alexander; malehypopygium.

Fig. 4 .

Eniphraqma(Epiphragma)hirtistylata Alexander; malehypopygium.

Fig. 5 .

Epiphragma@piphragma)interspersasp. n. ; rnale hypoPvgium

Fig. 6.

Epiphragma(Epiphragma) nigroplagiata A.lexander;rnale
hypopygium.

Fig. 7 .

Epiphragma@piphragma)parvisetaAlexander;antenna.

39
Fig. I .

Epiphragma(Epiphragma)parvisetaAlexander; male hlpoPygum

Fig. 9 .

Epiphragma@piphragna) oxypallusAlexander; male hpoPygrum.
(Symbols: ant, antenna;b, basistyle;d. dististyles;
i, interbase;p, phallosome;t, tergite).

6.:'.frr-:t9t

bnt