2 - €I
DIPTERAI 8 7 I I BF A S C I C U L E

GENERAI NSECToRTJM
D IR IGES PAR

P .W Y T S M A N

DIPTERAFAM. TIPULID,€
Surpeu. CYLIN DROTOMIN,IE
By Cuenws P. ALEXANDER
WITH

2

PLAIN

PLATES

t927

En ventechezLoursDnsMeI-VeRTENEUTL,
Imprimeur-Editeur,ffi-62, rue T'Kint, Bnuxelles
Prospectusgratis et franco sur demande

DIPTERA
FAM. TIPULIDE
SuereM. CYLINDROTOMIN.lE
Ry Cnanrns P. ALEXANDER
WITH

2 PLAIN PLATES

ffiHeCylindrotominaisthesmallestofthethreesubfamiliesofCrane.Flies
The flies of this group early attracted theconstituting the family Tipulida.
$jnlRpof the larva,
attention of Entomologists because of certain remarkable features
KSrtHil'Iipulida in their curious, spinous, eruciform aPpearance'
which differ from all other
Kdfu[bX
Bryophytic and Spermatophytic
green coloration, and habits of feeding upon the leaf-tissuesof various
plants.
was createdby Macquart (1834)for the Linnobia
The first genus to be describd., Cylindrotoma,
the elongate'lVleigen. As conceivedby Macquart, the genus was based chiefly on
distiaclissimoof
group two
the
add
to
to
authorthecylindrical antennal segmonts,an artificial characterthat induced
and are now placed with the Hexatoother speciesthat are only distantly related to the Cylindrotomina
Greatconfusion
Macquart and Limno\hila(Lasiomastix')nacrocuaSay).
mine Limoniina (Ulamaeroptr,za
proposed
the newwhenschinerregarding the natural affrnitiesof the group prevailed until 1863(f.or Limnobiatrisulcataschummel),gerreric terms phalacroccro(for Tilulo nllicata Linnaus) and.Triagno(I869) completedthe
at the sametime pointing out the relationshipswith cylind,rolona. osten-sacken
nod'ioornis
osten-sacken,
group of four Holarctic generaby proposingthe name Liogmafor the Americangroup of genera constituting theand the European gl,obrataMeigen. The oriental-Australasianyears.
stibadocere have all beeu describedwithin the past fifteen-fhere is still a great diversity of opirrion as to what value this peculiar grouP of crane-flies
On thethe tribe Cylindrotomini'should receive. Some recent authorities still recognizeonlygroup full family rank' while one student'
other hand, several European stud'ents give to thethe Eruceformia'from which he
Bengtsson (r8sz) has even proposedasectibnofthe.Diptera,
would rlerivethe higherPolyneura aud the Oligoneura'system of wiug-venationis used'
In the present paper, the Comstock-Needham-Tillyard
TheterminologyofthemalehypopygiumisthatofCrampton(1923).
the wing'drawings in thisThe writer would expresshis deep appreciation for help in making
p a p e rt o P r o f e s s o r a n d M r s . J . S p e e d R o g e r s 'DIPTERAG e o g n a p h i c a l d i l t n i b u t i o n . - T h e r e c e n t g e n e r ao f t h e C y l i n d r o t o m i n enumber but
seven of which the typical group (Cylindrotoma),including the four genera, Cylindrolottta,
Phaldcrcccra'Liogma and Triogma. is characteristic of the Holarctic Region. The only
species thatis known to occur south of these limits is Phalacroccra
Alexander,from the high mountains of
formosa
Taiwan, where the conditions are typically Holarctic. Vicarious representativesof theabove four
I'enera occur in the Western Palearctic Region (Europe), the Eastern Palearctic Region (Japan) andthe Eastern Nearctic Region (Eastern North America). But two of the four (Cltitdrotoma
andPhalauocera)have been taken in the lVestern Nearctic Region (Western Nortl-r America;.
The secondgrouPr the Stibadocere, is characteristicof the Oriental and AustralasianRegions,
two of the three genera (Slibadocnaand Stibadocerclla)occurring in the Oriental Region fromlndia to'lhe
two known species of. Stibadoccrodas
occur in the Australasian Region, from New South
Java.
'fhe
Waies to Tasmania.
Cylindrotominae are apparently quite lacking from the Neotropical andE t h i o p i a n R e g i o n s( r ) .
Geologlcal distnibutlon. - The fossil genus Clttaronyia Scudder is the only one of several
rlames proposed that seems to be correctly referable to this subfamily. 'l'he genus is very char.acteristicof the White River (Eocene) and Florissant (Miocene) depositsbrrt was apparently quite absent in thewarmer Oligocene.
Blolosv. -The adult Flies of the Cyiindrotomine

on vegetation in cool, usually shaded spots.

a r e s l u g g i s hi n t h e i r h a b i t s , b e i n g f o u n d
Triogna prefers open mossy nreadowswhere there is no

woody vegetation rvhatsoever. The immature stagesof this subfamily differ from those of all other
Tipulida so far made known in being spent on various Bryophytic and Spermatophyticplants, op theleaves of which the larva feed. 'fhese larva are usually green in color and strongly suggest aI-epidopterous caterpillar in their general form. Most of them simulate their host plants to an
astonishing degree. The immature stages of. Cylindrotomaare terrestrial, occurring o' the leaves ofseveralscatteredgenera of Spermatophyticplants. Liogma,likewise,is terrestrialbut occurs on va'ous
speciesof mosses,especially Hfinum and related genera. Triogma varies frorn semi-aquatic to aquatic,
living on various species of mosses. Phal,acroccra
is strictly aquatic in its habits. f'he habits of the
known Cylindrotomine
N r . 3 8 , p . 9 5 9[ r 9 z o ] ) .

h a v e b e e ns u m m a r i z e db y t h e w r i t e r ( C o r n e l l U n i v . , A g r . E x p t . S t a . , M e m .

Cylitdrotomais the best known genus, the larva feeding on a variety of higher plants, including
both nronocotyledons and dicotyledons. The life-history of C. sllandeusDoane, has been well describedb y C a m e r o n ( A n n . E n t . S o c . A m e r . V o l . r r , p . 6 7 [ r 9 r 8 ] ) , h i s a c c o u n t b e i n g s u m m a r i z e dh, e r . ea s
being representativeof the subfamily.
The adults first appear on the wing about the middle of XIay (in British Columbia). Without
food they do not live longer than five or six days, but in breeding cages,where they were supplied withfood in the nature of a sugar solution, they lived as long as frorn seven to nine days. In nature, theadults were found on the wing during a period of about three weeks.
Soon after emergencethe aduits begin to copulate, and one male nray have intercoursewith morethan one female. Copulatiou often takes place in a vertical position, the female above, the male below,
with the tips of the abdomens interlocked. In nature the act of coptrlation is generally undertaken in the(t) Since the above was written, arrewgenus and species of Cylindrotomine
has been discovered in Southern
Chile by Edwards and Shannon, December, 1926. The fly belongs to the Stibadocere
and. is somewhat intermediate
in its characters bets'een Stibad,occrclla and Stibadocetodcs.

FAM. TIP(TLIDE,
deep shade of the large leaves of the insect's food plants. The eggs are of a dull glistening white,elongate-oval in shape. In nature they are almost invariably found on the under surface of the foodplant, Trautacilcriagrantlis Nuttall (Ranunculacee), inserted beneath the incised epidermis. They ale
usually deposited in series along and just within the margin of the serrate leaf. The eggs are only
partly hidden beneath the epidermis, being exposed dorsally, the margins of the slit overlapping the egg
laterally. In ovipositing, the female rests on the under side of the leaf, with the tip of the abdomen
directed toward the leaf edge. The abdomen is slightly flexed ventrally, and the margin of the leaf isheltl between the bifurcated valves of the ovipositor, which is applied to the upper surface of the leaf,
while the paired cutting valves, with their blades, are applied against the under surface. The blades
are then moved to and fro, cutting a slit in.the leaf in u'hich the egg is deposited.
'Ihe
indoor duratiol of the egg stage is fronr 14 to r8 days but in nature as long as three weeks
may be required. When first deposited,the eggs are translucent, grayish u'hite, spindle-shaped,withthe chorion unornamented, measuring approximately o.84 by o.3o3 mm. The larvae require about
three hours to emerge from the egg and begin to feed upon the leaf tissue at once. The first-stage
larvae are grayish ll'hite in color, transiucent, and measure r'r9 by o.37 mm. They feed on both theupper and lower surfaces of the leaf, burrowing through the epidermal layer with their mandibles andfeeding on the parenchymatoustissue within. The young larvae are very sluggish' At the end of nineclaysthe larvae have increased in length to 5.84 mm., and show all the characteristicbehavior of the
Growth is very slow, and before the first molt the larvae become covered withparticles of their excrement which adheres readily to the skin. The first larval molt occurs after aperiod of from r8 to zr days. The second-stage larvae gradually assume a leaf-green color as they
larvae,continue to feed. Toward the end of July, coincident with the dying-off of their food plant. thewhich have now reacheda length of from 8.32 to 9 mm.,become quiescentand ceaseto feed. Feeding
leaves'ancl movement gradually cease completely and the larvae remain clinging motionless to thefull-grown larve.

the winter in al,ater the larvae generally fail among the dead leaves on the ground and under these passfollowing
dor.mantcondition. The overwintering larva first show signs of activity in March of thepupation inspring, when the food plant sends up its new shoots. Growth then proceeds rapidly until
lastbeing thethethe middle of May. There seems to be one molt before hibernation and two after,
length'casting of the larval skin, previous to pupation. The fully grown larve measule 17 mm' in'fhey
attachlarve
theare invariably found on the upper surface of the leaf. Before pupation
pseudopodia'
themselvesfirmly to the surface of the leaf, or to the leaf petiole, by means of their anal
and thethorax'theThe sliin splits transversely behind but is only partially sloughed off. The head,infirst four abdominal segments of the pupa are exposed, but the apex of the abdomen remains encased
bleedingtheInsurface.
the larval skin, the terminal portion of which remains attached to the leaf
required
cages the pupai duration varies from 6 to ro days. From one and one-half to two hours arefor the adult to emerge from the pupal hull.
(Meigen)'
Liogrnaoccursin woods where its food plants grow. The life-history of L. glabratadays' the
to
rz
rr
from
stage
pupal
the
requires one year, the egg stage lasting from 8 to ro days andlarval stage requiring the greater part of the remaining time.
shade of woody
Triogrta is eminently characteristic of wet mossy meadows, unprotected by theplants. The very sluggish larve live and feed in the moss-cushionsin such habitats''fhemosses' TIle
larva of.phaloorocelaoccur among aquatic plants where they feed chiefly on
vegetation by alternatelyla^,a is extremely sluggish, progressing slowiy through the mats of submerged
ponds in which a moss
grasping with the mandibles and.the large anal hooks. The larval habitat is in
The female deposits
vegetation flourishes and in which currents keep the water in constant motion.
arelaid singly'
number,inthe eggs in the axils of the leavesof submerged mosses. The eggs' about6oDIPTER.,\adhering lightly to the moss. Normally the egg stage requires from 8 to rz days, thelarval
stage
approximately r r months and the pupal duration 7 or 8 days. During the larval d.evelopment
there areat least eight molts. Upon emergence from the egg, the young larva measures from z toz.z5 mm. in
iength and about o.5 mm. in diameter, excluding the body projections. The pupa iscomparatively
active, moving about by flexion of the abdomen. In a position of rest, the body is usually
held in a
vertical position with the tips of the breathing horns just reaching the surface o{ the water.
When theadult emerges, the cast pupal skin is left attached to a moss leaf by means of the dorsal
abdominal
projections.
Nothing is known of the habits of the Oriental and Australasian genera constituting
theStibadocere,
except in the case of Stibodoccrodcs
australiansisAlexander, which the collector,
Mr. G. H. Hardy,

found resting sluggishly near a small waterfall in New South Waies.

Ghlnectenr of thc rdult Fllcr. - The adult Flies of the Cylindrotominechalacteristic form, the relatively long abdomen, with the venational peculiarity of an a!t'annl

have afusion ofvein r?r with Ra f .r back from the wing-margin, being especially characteristic of these flies.
The head is always large, with the vertex between the eyes very broad, wider than the diameterof an eye, the ommatidia of the latter usually d.elicate. Rostrum short to very short,with faintindications of a nasus; paipi short, the terminal segment short but still much longer thanthepenultimate. Antenna r6-segmented, of various forms in the different genera in Cylindrotoma
the;segmentsare cylindrical or nearly so, a condition that reaches its greatest extreme in the three
generaof the Stibadocera where the organ in the male sex is longer than the entire body, with the flagellar
segments provided with an abundant erect pubescence. The flagellar segments in Liogma and,Triogmaare more or less produced on the ventral margin to produce a serrated appearance. pronotumm o d e r a t e l y l a r g en, o t d i s k - l i k e a s i nt h e T i p u l i n e .
I \ { e s o t h o r a x a s i n t h ef a m i l y ; m e r o n v e r y s m a l l , t h e
middle and hind coxa thus being approximated. Surface of the head and thoracic interspaces withconspicuous deep furrows and fovee in the genera Triogma and Stibadocc/a,mote nearly smooth in theother genera. Legs moderately elongate, relatively slender; tibial spurs present, n.ricroscopicallysetiferous;unguessmooth; empodia distinct.
Wings genefally ample, with a characteristic venation. Within the limits of this small subfanrilyof crane'flies, we find a range of venation that is almost, if not quite, as extensive as that found in thegreat subfamily Limoniina.
It is a far cry, indeed, from the venation of. Cyttaromlncto that of themost specialized genus Slibad,occralla.From a study of all of the genera and most of the known species
it is possible to reconstruct a hypothetical type for the subfamily that does not differ strikingly from thefossil genus Cytlaronlia, except in the details of the radiai field. The prime tendency in the subfamilyis the alfarc,rl fusion of vein Rr with vein rRz*.r to produce a long fusion back from the wing-margin.
This can readily be seen from an examination of .virtually all of the figures. As the writer has
demonstratedin an earlier paper (cornell univ., Agr. Expt. sta., Mem. Nr. 25, p. g63[1919]), thisfusion is merely apparent and results from the atrophy of the distal tips of veins Rr and Rz. The
behavior of the various fieids of the wing is now consid.ered(pl. I, Fig. A-H) :Sabcosta(Sc). In the primitive condition, both branches of Sc are preserved, Scr apparently
being the first to atrophy.
Rodius (R). The vein Rr is elongate, with the extreme tip atrophied in the great majority of thespecies. It is preserved in Cylindrotomanigriocnttis Loew (Pl. 2, Fig. 2), Triogna c*cult'ta (OstenSacken)(Pl. I' Fig. E), Stibadocuamctall'icaAlexander (Pl. | Fig. F) and others. The remajnder,of Rr is always in perfect alignment with the radial crossvein (z), bending gently caudad toward vein
Rzl3 which it meets immediately beyond the stigma and usually at the fork of Rz*s.
In almost allcases' the distal section of vein Rz is entirely atrophied. but persists in some speciesof phalaerocera,asFAN{. TIPULIDA;
P. rcllicata (Linnaus), P. neorcta Alexander (P1.2, Fig. 6), and in some specimensof P. nikatlo
Alexander(P|.2,Fig.5). Thisiastcaseispeculiarinthat/connectswithRzfifarbeforetheforkof the latter. From the persistence of the fi'ee tips of Rr and Rz ir-r the above cases, it is fullyu,arranted to add them to the hy'potheticaltype (Pl. I, Fig. A). The radial sector (Rs) is almost
alrvaysvery long, arising before nri.llengthof the rving, gentiy arcuated at origin. The behavior of theposterior branch of the sector (Rl*S) is extremely various. In the more prirnitive condition (Cyttaroand the Stibadocerae), the radial-nredialcrossvein(r+n)is present
most Pholacyocnd,
nyia, Cylintlrotoma,arrd eiongate,bqt in the genera l.iogmaand Tiogna, the tendency is distinctly to obliteratethis crossveinIntbee ( S e eP l . l , F i g . E ; P l , 2 r F i g . 8 - 1 3 ) .
b - v t l r e f u s i o n ot hf e a d j a c e n t v e i nRs+, * S o n M t frnore primitive genera, r-,,t, or its position, is distad of the fork of Rs but in the more specializedAlexander, Pl. l, Fig. F) or before
rnernbersr.-n is at. or close to, the fork of Rs (Stibadoccromctallica(Alexander')(Pl. I, FiS. G) culminating in the very curious condition
auslralicnsis
it, as in Stibadocarodes(detrIeijere) (Pl. l, Fig. H), whele r-m lies before midlength of thealbitarsis
forrrrd in Stibodocerclla
extr.emelylong sectol which forlis beyond the levei of cell rsl Mz; in this latter genus, Rz*.J isperllendicular in position and simuiates in appearance the apparent radial crossvein (rRz) in theies.
Lirnoniine crarre-fl
all four btanchesof media are
Cylindroloma)
In the primitive condition (Cyttarornyia,(M),
Mcttiap r e s e n t ,c e l l M b e i n g v e r v s h o l t - p e t i o l a t e t o s e s s i l e . l n C y l t a r o n y i a , v e i n M i s u n i t e d w i t h v e i n R l * sby a supernumerarycrossveinto form an apparent second rrdiscal cell rr imnrediatelyabove the primaryo n e ( P l . l , F i g . B ) . I n t h i s s u b f a m i l y . t h e r e d u c t i o n i n t h e n u m b e r o f b r a n c h e so f M a p p e a r st ohave been brought about by the atrophy of vein M 3, ^ condition that is well illustrated in the genus
phalacroccro. In P. occiduttalis,nov. sp. (Pl, l, Fig. D), the base of this vein persists but the apex isAlexander, Pl. 2. Fig. 6)' the vein
entir.ely atrophied. In other species of the genus (as P. rtco*ena
has entirely atrophied, the apparent medial crossvein(ar) in this casebeing a linear composite of.m andthe basal deflection of.Ms tn perfect alignment. In the more highly specialized members of the genus
p. ntihatloAlexander (Pl, 2t Fig. 6), P. fonnosaAlexander (Pl. 2, Fig.7)r cell M r has becomebroadly
jssessile. The ccndition obtair.ringin these two latter cases that found in all the more specialized
nrembers of the subfamily where vein MJ is interpreted as being entirely atrophied. Cell rst Mz isaushalicnsisAlexander.
closed in all casesexcePt Stibailocerodcs
Cubitus(Cu.\. The n'redial-cubital(zr-rz)crossvein(the basai deflecttortof Cut of the ComstockNeedhartr system of venation) is ahvays well preserved, placed trefore the fork of M in some speciesof Liogna, beyond the fork and beneath cell rsl M z in the majority of species. The semi-atrophieds e c o n d b r a n c ho f . C n i s a l w a l ' 5 w e l l p r e s e l v e d , b e i n g u s u a i l y p e r s i s t e n t t o o p p o s i t e t l t e n - ccur o s s v e i n .
where a single one
Atnl Vcins(l). Trvo anal veins are present except in the genus Stibadocuclla
is preserved.
elongated, especiaily in the male sex. The male hypopygiunr,
The abdomen is cor-rspicuously
as a rule, is powerfully constructed. the basistvlesbeing verv large, bearing at their outer ends the(Alexander), the basistylesare
australicasrs
single powerful dististyle (Pl. l, Fig. 9). In Stibadocerodcs
very stout. their mesal faces grooved to receive the trvo dististyleswhen in a position of rest. The
outer dististyle is more slender, gradually narrou'ed to the apex. The ilrner dististyle is a broadly
flattened.blade. ,lEdeagusterminating in three points or prongs, a condirion that suggeststhe family
T a n y d e r i d e . T h e o v i p o s i t o r o ft h e C y l i n d r o t o m i n e i s v e l y r e m a r k a b l e( P l . l , F i g . 8 ) . T h e v a h ' e s
are fleshy, the dorsal valve being deeply furcate, tire depth of the fork varying in differentspecies. l'heventral valves each have trvo cutting edges, an internal serrate blade and an external plane biade'a
Ventrad of these and arising apparently frorn the eighth sternite iies a valve that is shaped iil<ebeenploughsharewhich functions in guiding the ernergingegg into a slit in the leavesof plauts that has

DIPTERA
rnade by the cutting valves. For a more complete discussion of the details of structure of the ovipositor
and the methods of oviposition in the typical genus Cytrindroloma,
the following papers are recommended.:
C am e r o n , A. E. Life-history of the Leaf-eating Crane-fy. Cylirdrotomasllcnolcns
Doane (Aqn. Ent. Soc.
Vol.
It.
r-18
America,
P.67rf.
[r9r8]); Lenz, Fr., Die Eiablage von Cylhdrotomadistittcflss6c(Mg).
(Arch. f. Naturg. 87 Jahrg. Abt. A, p. rz8, f. r-7 [r9zr]).
Ghrnectcnr

of thcLlnr-.

-

Body relatively short, the integument covered with tubercles,

leaf-like or filiform appendages; head completely retractile within thebody; body with green tints.
of the Pup-. - Pupa within the last larval skin, which attaches it to the host
Chamctcnr
plant; integument generally with filiform or leaf-like appendages.
-

Phylogcny.

At the prosent time it is virtually impossible to indicate the past relationships of

the Cylindrotomine.
AtthebeginningoftheEocene,thogroupwasalreadyalargeandflourishingone in cold, Northern climates, and the few isolated genera now persisting seem to be scattered relics.
The Stibadocere

are the more specialized group and have apparently been derived directlyfrom Cylhdrotona or a closely related form. Liogna andTriogmo have likeurise been derived from anancestor that was very close to our recent Cylindrotoma. Phalaooccza is much more isolated in both its
larval and imaginal characters but presumably came off the main stem before or almost coincidentlv
with Cylindrotomo.

S u a r r n n .C Y L I N D R O T O M I N . €Lf rnnoblnt

Gyllndnotomrfonmer

Schiner, Fauna Austriaca,Dipt. Vol. z, p. 56o(rS6a).

Gyllndnotomlne

osten-Sacken,Mon. Dipt. N. Amer. vol. 4, p. 289 (r869).

Gyf indnotomlnl

Scudder, Proc. Amer. Philos. Soc. Vol. 32, p, r&g (rSS+).
Bengtsson,Kongl. Fysiogra. Siillskapetsi Lund Handl. (n. s.)Vol.8, p. rro(rSgZ).

Enuc-fonmlr

Cylf ndnotornldr

K6rtesz, Cat. Dipt. Yol. z, p. 3zZ (r9oz).

Gyllndrotornlnl

Brunetti, Fauna Brit. India, Dipt. Nemat. p.358 (l9rz).

Gyttanomini

Meunier, Zeitschr. Deutschen Geol. Ges. Vol.67, p. 22g (1916).

CyllnCnotomln-

Alexander. Cornell Univ. Agr. Expt. Sta. Mem. Nr. 38, p.959 (r9zo).
TABLE

OF GENERA
Adults.

(Nore. -Thefemale sex is not known in the majority of the genera of the Stibadocera,
necessitating the usageof sexual characters.)
r. Antenna of both sexes short, not exceeding the
combined head and thorax

Group Cyrrupnorotra.a.

2.

Group Srraepocsn,E.

6.
3.

Antenne of the male as long as, or longer than,the body, the flagellar segments provided withlong, erect verticils
z. Four branches of.M reach the wing-margin.
Three branches of M reach the wing-margin+.

3 . A supernumerary crossvein connects vein Rlli
with

Mr

near its origin,to produce two

so-called < discal cells rr .
No supernumerary crossvein as above

8. Genus Cyrrenouyrl, Scudder (fossil).
r. Genus Cruupnoroue, Macquart.

FAM. TIPULID,IE
4. Head and intervals of mesonotal prescutum withnumerous deep punctures and fovea; bodycoloration dull
Head and intervals of mesonotal prescutum with.

4. Genus Tnrocve. Schiner.

the punctuation indistinct or ill-defined; body-coloration shiny (Genus Liogma) or dull .
5. Antenne subserrate in both sexes,the ventral faceof the basal and intermediate flagellar segments
more or less produced; body coloration shiny
or subshiny.
Antenna not serrate, the flagellar segments subcylindrical with a conspicuous basal swellingthat is short-verticillate; body coloration, atleast in part, dull.

3. Genus Lrocue, osten-sacken.

2. Genus pnerecnocrnl,

Schiner.

6. Rs of normal length, r-n being at or close to thefork; two Anal veins present
Rs very long, forking near mid-distance between

7.

r'm and the wing-tip ; a single Anal vein present. 7. Genus Srrseoocansne, Brunetti.
Z. Tip of vein Rr presorved; prescutal interspaces
deeply punctured; no macrotrichia in cells of
wing; tarsi concolorous with the remainderof legs.
Tip

5, Genus Srrseoocsne, Endellein.

of vein Rr entirely atrophied; prascutum
entirely impunctate; sparse macrotrichia inapices of cells R5 and M z; tarsi white

6. Genus SrrsepocnnoDrs,nov. gen.
Larve.

r. Body appendages very longand filiform; aquatic.
Body appendagesshort, leaflike or tuberculate
z. Dorsal appendages all sirnple, on the terminal
abdominal segments in a single rowl terrestrial
on spermatophytic plants.

z. Genus Pnat,acnocsne. Schiner.
2.

r. Genus cvr,rNonoroua, Macquart.

Dorsal appendages with teeth on anterior convex
sido

3.

3. Some of the dorsal appendageswith three or fourteeth on anterior face; aquatic or semi.aquaticon mosses

4. Genus Tnrocul, Schiner.

Dorsal appendages with one (nodicornis) or. two(glabrata)teothl terrestrial in mosses .

3. Genus Llocue, osten-Sacken.

Pupe.
r. Basal abdominal tergites without spines .
Basal abdominal tergites with acute spines
z. Mesonotumunarmed; segments6 and8of abdomen
each with two powerful dorsal hooks; abdo-

.

2.
3.

DIPTERA
minal segment 7 with a pair of strong ventral
sprnes; pronotal bleathing hor.ns elongate,
directed backward .
z. Genus PualecRoceRe,Schiner.
Nlesonotum with two flattened erect lobes; abdo_minai segments 6, 7 and 8 unarmed; pronotal
breathinghornssmall, directedslightly forward.. r. Genus Cyrrxpnotorre.
Macquart.
3. .\bdominal spinesbranched .
3. Genus Lrocrvre(pars).; 4. Tnrocve, Schiner.
Abdorninal spinessimple .
3. Genus Lrocve (pars).

t . G e r u u sC Y L T N D R O T O MM
A ecguanrG y f i n d n o t o m a M a c q u a r t ,H i s t . N a t . D i p t . p . r o 7 ( r S 3 a ) .
Remanks. - The adult flies of. Cylittdroto,naoccur on rank vegetation incool. boggy andswarllpYrvoods. The occurrence and habits of the larvaeare discussed.
eisewherein this paper.
Ghanactcns of Adurtr. - Vertex betrveenthe eyes broad, much wider than the
diameter of asingle eye. Rostrum short, highly arched dorsally; parpi rerativelyshort.
Anterrne of moderatelength
but always much shorter than the body, r6-segmented; flagellar segmentssubcylindrical,
narrowed at
the incisures, densely provided with a delicate white pubescenceand witha few longer.,scatteredverticils. Eyes with relativelvsmall onrmatidia.
Tholax as in the subfamily. Legs normal, relatively sler-rder;tibial spurs approximately
equal,
sparselysetiferous. Wings (Pl. 2, Fig. l-3) with venation as in the sublhmily,
distinguishedfrom allother recent genera by the presenceof four branchesof media reachingthe rving-rnargin; cell M r sessile
orveryshort'petiolate.Tipsof veinsRrandRzentirelyatrophied; rr-cnbeneath
cellrstMz.
Abdomen relatively long and slender. trtale hypopygium (Pl. l, Fig. g) relatively
large andprominent; caudal margin of ninth tergite with a broad, deep notch, the lobes
thus formed rounded attheir aPices. Basistyleconspicuous,the inner margin at the tip produced caudad
into a small flattened
blade. the dorsai face with a densely hairy lobe; dististyle single, large. Ninthsternite small, deeply
split on the midventral line by a very acute angle. Aedeagus trifid, the inneredge of the lateral arms
w i t h n u m e r o u s a p p r e s s e d t e e t h .O v i p o s i t o r ( P l . l , F i g . 8 ) a s i n t h e s u b f a m i l y ; v a l v e s
fleshy, with tr,voof the sternal valves provided with inner serrated margins that function in oviposition.
- Body covered with simple tubercles, a med.iandorsai
Ghenactenl of Lerv-.
r.owand a
double veutral row- Spiraculat disk large, surrourrded bv small lobes.
Head capsule compact.
l'Ientum with about r5 teeth.
Ghenactenr of Pupa. - Pronotal breathiug horns short, cylindlical, directed
strongly
ventrad. Vlesonotumwith trvo flattened lobes directed cephalad. Abdomen
unarmed with spines orlobes.
Type spcci es : Cylindrotomalistirtct,issinaMeigen.
Geognaphical dlrtnlbution.
r, C. anuicaaa Osten-Saclcen,Proc. Ent. Soc. Philad. Vol.
4, p, 236 ( r 865). Eastern North .{ merica.
z. C, distinclissimaMeigen, Syst. Beschr. Z w e i f l . l n s . V o l . r , p . r 3 r
Ir8r8] Europe.
(Liwtobia).
C. brevicornis
Zetterstedt,Ins. Lappon.Dipt. p. sa6[re38](Ti!ula).

FAN{.TIPULID,€
3 . C . j o p o n i c aA l e x a n d e r ,A n n . E n t . S o c . A m e r . V o l . r z , p . 3 + + ( t S t g ) .
4. C. nigtiaentrisLoew, Stett. Ent. Zeit.Yol. ro, p. 342 (t8+S).
r so a n e ,J o u r n . N e w Y o r k E n t . S o c . V o i . 8 , P . r 9 7 , p l . 8 , f . z r
5 . C . s l , I c n d aD
(r9oo).

9J apan.

IRegion.
Northern Palaarctic
Western North America.

C.juncta Coquillett, Proc. lVash. Acad. Sc. Vol. z, p.4or (r9oo).

6. C. tarsalisJohnson, Psyche' Vol. r9, P. 2' f. 4 Qgrz).

Eastern United States.

C. anonala Johnson.ibidem, Vol. 19, p' z, f . 3 (tgrz\.

2 . G E r u u sP H A L A C R O C E R AS C N I U C R
P h a f a c n o c e n e S c h i n e r ,W i e n . E n t . M o n a t s c h r .V o l . 7 , p . 2 2 + ( r 8 6 3 ) .
tifulina Osten-Sackett,are most commonly found in
fhe adult flies of Phalacyoccra
close proximity to sphagnum bogs. The habits of the larve of the genotype, P. rcflicala Linneus,
Remanks. -h;tve been discussedelsewherein this paper.
Chanactens of Adults. - Vertex betlveen the eyes broad; head ratirer strongly narroweti
behjnd. Rostrum short, feebl,r'arched dorsally, the apex terminating in a microscopic nasus; palpi
short, the terminai segmentfeebly clavate. Antennc of modetate length, in the male about equal inlength to the combined head and thorax, r6-segmented; flagellar segmentsgenerally similar to thoselbund i11the genus Tipula.,consisting of relatively elongate, subcylindlical segmelrts,the base slightly
eglalged antl set with a whorl of about four stout verticiliate seta that are shorter than the segments'
Eyes rvith snrali, delicateommatidia.
'['horax
as in the subfamily. Legs relatively slender. with delicate appressedsete; tibia withtlvo long, microscopically setifelousspurs. Wings in the prirnitive condition with a venation that is(Compare Pl. l, Fig. C and D). The radial fleld is peculiar
notstrikinglydifferent trom Cylittdrotoma
in that some of the species,as P. rcplicala(Linnreus)and P. ,tclram Alexander, have retained the distal
section of vein Rz (Pl. 2, Fig. 6). In sorne specinrens of.P. nihado Alexander, what appears to be
the free tip of vein Rz is present (Pl. 2, Fig. 5) btrt lies far beyond r so the latter crossveinconnects.l far before its fork. The extreme tip of .Rr is atrophied but in most of the known species
i t s b a s e i s p l e s e r v e da s h o r t d i s t a n c e b e y o n d t h e e n d o f v e i n S c ( P l . 2 , F i g . 4 a n d 6 ) . T h ernedial field presents a transition betlveen the primitive condition found in Cyttarcmyiaand Cllindrotonn( Pl. | , F ig. B, C) and all the remaining membersof the subfamily. Yein M s is more or less atrophied
irr all the speciesknown, so but three branches of the vein (M r, M z and M 4) attain the margin. In

rvith Rzf

some species (P, rellicala Linneus; P. lipulina Osten-sacken; P. ncorcnaAlexander and P. occidentalis,nov. sp.) cell Mr is short-petiolateto nearly sessile; in other cases(P. formoso Alexander, P, mihado
Alexander) the cell becomes broadly sessile,in the extrenre case, the basal section of vein M z being
equal in length to the posterior element that closescell rsl M z (the composite vein M 3 plus the medial
crossvein). This latter caseis similar to that found in all higher members of the subfamily.
Abdomen elongate, the male hypopygium relatively large, the structure not essentially differerrt
from the common type of the subfamily. Ovipositor with four nearlv fleshy valves. approachingthe
normal Linroniine or' Tipuline type.
Chanactens of Lanv-. - Body covered n,ith numerous, elongate, trachea-bearingfilameDts,
the posterior pair on dolsal segments deeply forked, the others simpie. Spiracular disk with dorsal
pair of lobes, formecl by rudimentary posterior branch of branched filaments of eighth abdominais e g m e n t . H e a d c a p s u l ec o m p a c t . ) l e n t u m r v i t h a b o u t I 5 t e e t h ( P l , | , F i g . 3 ) .

DIPTERA
Chenactenl of Pupr. - Cephalic crest low, not setiferous. Pronotai
breathing horns long,almost straight. Dorsal abdominal segmenrsrvith tubercles, those ofsixth and eighth segmentsenlarged
into spinous hooks; two pointed tubercleson seventhsternite (pr.
r, Fig. 4).
Typc species t phalacrocclarcllicata Linnaus.
Oeognrphicel dlstnibution.
r. P.iformosaAlexander, philipp.
Journ. Sc. Vol. 22, p.+7g egz3).
Taiwan.
z. P. nihadoAlexander, Ann. Ent. Soc. Amer. Vol. rz, p.'SiA
GSrS).
Japan.
3 . P . n c o * c nA
a l e x a n d e r , p r o c .A c a d .N a t . S c . p h i l a d . p .O o : , p t . z S , f .
r o l r g r a ; . Eastern North America.
nov. sp. (r).
4. P, occidcnlalis,Western North America.
5. P. rcplicalaLinneus,
Nat. (ed. ro), p. 5g7 [r75g] (Ti/ula).
Europe.
_Syst.
p. breairostr.is
Zetterstedt,
Ins. Lappon.Dipt. p. eaairi.3e1elita),
P. ntulicornis
Schummel,Beitr. Ent. pt t, p.-tzz, pi. ,, i.'a ; pL.5, f .
7-9
It8z9l(Limnob,a).
6. P. tilutrinaOsten-Sacken,proc. Ent. Soc. philad. Vol.4,
p.24r (1365). Eastern North America.

3. G ENUs LIOGMA osTE N-sAcKEN
Liogme Osten-Sacken,Mon. Dipt. N. Amer. Vol.
4, p. 29g (rg69).
Rcrnenkr' - The very sluggish adult Flies of the species
of.Liogma may be swept from rank
vegetation in cool, shaded woodlands. As indicated before,
the immature stagesare spent in relatively
dry moss cushions.
Chenectcnr of Adults. - Vertex between the eyes broad.
Rostrum very short, arched,dorsally; palpi short. Antenne of moderate length,r6-segmented, but appearing r7-segmented
because of the very long terminal segment, which, insome cases, appears to be binodose; flagellar
segments (Pl' | ' Fig. 6) with the lower or ventral facestrongly prod.ucedto give the organ a marked.
serrate appearance; individual segments densely covered
with a pale, erect pubescenceand with a few
powerful black bristles that are somewhat longerandmore slender on the dorsal side of the segments.
Eyes with relatively small ommatidia.
Head and thorax nearly smooth and subshiny. Legs relatively
long and slender, covered withrelatively numerous subappressed sete; tibial spurs relatively
slender, setiferous. Wings with thevenation variable in severarregards. sc rong, end.ingbeyond.
the fork of rRs,scr atrophied or tendir.rg
to atrophy; Rs long, the origin arcuated; tip of .Rr atrophied,in most caseswith no indication of its
point of leaving the rnain vein; tip of Rz atrophied,l r-npresent in some cases,in others obliterated
by the short to long fusion of vein Rt*son Ml jz (pl. 2, Fig. g-lO); m-cuvariablein positio', inthe L' glabrutafigured being nearly its own length before
the fork oI M, inother casesa short distancebeyond the fork of.M and beneath cell rct M z.
Abdomen relatively long and srender (pl. l, Fig. |), as in the subfamily. Mare hypopygium
large, having the general structure of the subfamily;
basist5rlesand dististyles large and massive.
Aedeagus large, in some casesvery powerful, trifid at apex, asin the subfamily. ovipositor with fleshy
valves of the same general type as in the typical genus Cylindroloma.

(t) Phehorocrr.a
oocldcntllll,
nov. sp. - Male. Length about rz mrr.
black, gray pruinose; wings pale brown,
the "iig-"oval, slightly darker brown;
atrophied, as in P. tiltliza osten-sacken; distal
seciion -of vein Mg persistent basa[y.
Holotype, 6f , Mt. Rairier, Washington,
June roth (in Alexander Collection).

Wing rr mm. General coloration
distal section of veiu R9 eutirelyFA\,I. TIPULID,E
Chenectenr of Lanva. - Body covered with elongate, leaf-iike projections, the dorsal ones
with one or two teeth on anterior convex face. Spiracular disk surrounded by four lobes. Head capsulecompact. Mentum with about 14 teeth (Pl. | , Fig' 2)'
Chenactcns of pup-. - Metanotum and abdominal tergites with elongate spines, branched inL. glabrata(Meigen), simple in L. nodicorzrs(Osten-Sacken)'
Osten'sacken (Pl'Type spccl es z Liogma nod'icornis| ' Fig' 6)'

GcognaPhical distni bution.
r4z [r8r8]
r. L. globrata Meigen, Syst. Bescht' Zweifl' Ins' Vol' r, p'(Linnobia).
proc. Ent. Soc. Philad. Vol. 4, p. z3g [r865]
z. L, noilicorzrsosten.Sacken,(Triogmo).
p. r95 (rStS)'za.L. ttodicornisfaacolaAlexander,The Canad. Ent. Vol. 5r,
p. 3+5 (tgrg).
3. L. scrratic,ln;s Alexander, Ann. Ent. Soc. Amer. Vol. rz,

Europe'North-Eastern North
meuca'South'Eastern United
Japan.

[States'

4. GEttus TRIOGMA ScxlNER
T n i o g m a S c h i n e r ,W i e n . E n t . M o n a t s c h r ' V o l ' 7 , p ' z z 3 ( 1 8 6 3 ) 'where they may be
Remenkr. - The adult Flies of. Triogma occur in wet, mossy meadows,
and mosses' The immaturefound walking and fluttering about among the rank growth of sedges
stagesare spent in these moss cushions.
head, as well as the
Chanactent of Adultl. - Vertex between the eyes broad, the entire
very
shortl palpi small,thoracic interspacesdeeply and consptcuouslyroughenedand pitted. Rostrum
penultimate segment. Antenne
the terminal segment short but about twice as long as the very shortstrongly narrowed at base,r6-segmented, of moderate length; flagellar segments (Pl. l, Fig. 7)
the lower or ventral face;widened distally into a clavate structul'e but scarcely more produced onommatidia moderately
shorter and slightly more produced in the female than in the male. Eyes withlarge and coarse.
the tarsi; tibial spurs
Thorax as in the subfamily. l-egs relatively short and stout. especially
opposite or some distance
setiferous, except at the extreme tips. Wings lvith Sc long, ending nearlyshor t e r a n d m o r e a r c u a t e di nb e y o n d t h e f o r k o f . R s ,S a , r t e n d i n g t o a t r o p h y ; R s m o d e r a t e l y e l o n g a t e ,atrophied in the otherT. cxculflaOsten-sacken;tip of veinRr preserved inT. c*cuQtaOsten-Sacken,
vein
R+15 on Mt f z' thisoffusionspeciesl tip of vein ftz entirely atrophied.;z-D obliteratedby theso but three branchesof M
fusion variable and sometimes very extensive; vein M3 entirelv atrophiedattain the wing-margin 1m-cubeyond the fork of M, beneath cell tst Me.
in minor details'
I\Iale hypopygium and ovipositor almost asin Liogma, differing only
projections, some of the
Ghanactcns of Lar.v-. - Body covered with elongate, leaf-likedisk with four primary lobes
dorsal ones with as many as four teeth on anterior convex face. Spiracularand an additional smaller median dorsal lobe'
Chanactcnt of Pup-.

-

Dorsum of abdomen with elongatebranched spines'

Type specl e, . Trio7ntotrisulcataSchummel'
GeoEnaPhical d iltnibution.
(I865).
t, T. ucult'ta Osten-Sacken,Proc. Ent. Soc. Philad. Vol. 4, P.23g

Eastern North America'rz

DIPTERAz . T . h u w a u aAr l e x a n d e r ,T h e C a n a d .E n t . V o l . 4 5 ,p . 3 z r , p l .
+, f.4;pl. ro, Japan.
f. r3-r5 [r9r3] (Liogma).
3 ' T . t r i s u l c a l o S c h u m m e l , B e i t r . E n t . p t . r , p . r o 5 , p l . r r f . 3 [ r g z 9(]L i w t o b i a ) .
Europe.

5. Genus STIBADOCERAEruoeRLEtN
Stf badocena Enderlein, Zool. Jahrb. Syst. Vol. 32, p. g3, f. Az$grz).

Chcnectenr of Adults. - Antennc r6-segmented;in the male longer than theentire body, theflagellar segnrentsprovided rvith long, conspicuous, erect, delicate seta. Headsmooth.
Interspacesand margins of the mesonotal prescutum punctate. Wings (pl.
2, Fig. l4) with
Sc moderately long, ending beyond three'fourths the length of Rs, Scr preserved (s.
buuattsEnderlei')o r i n d i s t i n c t ; t i p o f R / p r e s e r v e d ;r v e r y s h o r t , u s u a l l y n o t l o n g e r t h a n t h e f r e e p a r tof Rr (a littlelonger
inS. mctallicaAlexander); Rs long, the long basal section arcuated. at origin; r-m at (5. quadricclluln
Brunetti) or before the fork of Rs, in S. bullartsEnderlein, a distance before that is
approximately equal
to r-n; cell rsl M z elongate-hexagonal; cell M r lacking; three branches ofM reaching the wing-margin.
Abdomen slender; genitalia relatively small.
Type species: Stibadoccra
bullansEnderlein.
G e o E n e p h i c a ld i s t n i b u t i o n ,S, bullataEnderlein, Zool. Jahr.b.Syst. Vol. 32, p. g4, f. Az (rgrz).
Sumatra.
S . n c l a l l i c aA l e x a n d e r ,P r o c . U . S . N a t . M u s . V o l .
49rp.r7d,pt. aS,f. fo Java.
(r9r5).
3. S. quadricclhilaBrunetti, Rec. Indian, i\Ius. Vol. 6, p.z6g[r9rr] (Cylin- I n d i a .
rlrolona);I.'aunaBrit. India Dipt. Nernatocera,p. 36r, p|.7, f. vz
lryrzf (Cylindrotona).
I.

2.

6 . G e r u u sS T T B A D O C E R O D E SN, o v . c E N .
Chenactenc of Adults. -Characters as in Stibodoccra
Endellein, differing as follows : Headand thorax quite impunctate. Antenne of rnale very elongate, as in this groupof genera. Wings( P f . 2 , F i g . 1 6 , l 6 ) w i t h a f e w m a c r o t r i c h i ei n a p i c e s o f c e l l s R s a n d M z . V e n a t i o n: Crossveinz_rz
at (S' tasmanizszs'Alexander)or before (5. attstralicnsrs
Alexander) the fork of Rs,. Scr and Scz pr.esent
but in sonre casessubobsolete, Sc7 ending before midlength of.Rs (tasmanicnsis)
or beyond (auslraliatsis);
tip of l?^ratrophied; cell rsl Mz closed (tasmaniutsis)
or open by the atrophy of the combiued outerdeflectionof Ms andnt; m'ur'beneathcell rst Mz. Male hypopygium with the tergite
feebly concave
medially; basistylesvery stout, the mesalface longitudinally grooved for the receptionof the dististvles;
dististyles two, the inner more flattened anti blade-like than the slender.,apicallycurved outer stvle;
edeagus terminating in three branches.
Type rpecies= Stibodoccrodcs
auslralicnsisAlexander.

Geognephical distnibution.
t. S, austrahaasas
Alexander, Proc. I_inn. Soc. N. S. Wales, Vol.
47, p. 586 New South Wales.
corc lla).
lr gzzl (Stibad.o
2, S. taynanjaz.srs
Alexander, Rec. S. Austral. Mus. Vol. z, p. z5olrgzzl Tasmania.
(Slibadocerella).

FAM. TIPULID,IE

r3

7. Genus STIBADOCERELLABRurrrETTI
Stlbedoccnella Brunetti, Rec. Indian Mus. Vol. r5, p. 283 (r9r8).
A E a g t o m y l e d e M e i j e r e , B i j d r . D i e r k . A m s t e r d a m ,V o l . z r , p . 1 7 ( r 9 r 9 ) .
Remanks. -The Stibadocerogroup of genera fall in two distinct divisions, depending on theposition of r-n in relation to the fork of fls, and the number of Anal veins. ln Stibad,occraand
this crossvein is at or close to this fork, whereas in Stibadoccrcllathe fork of the sector is
Stibadoauodc.s,
approximately at mid-distance between r-m and the wing-tip, a very unusual character in the family.
Chenrctenr of Adults. - Anterior vertex rnoderately broad, scarcely swollen; front almost
flat, rather short, not produced into a rostrum ; palpi moderately elongate, the terminal segment notprolonged. Antenne of male very long, filiform, r6-segmented; basal segment of scape very short;
flagellum very long, nearly twice the length of the entire body, all flagellar segments strongly elongated,
rod-like, the terminal segmentsshortestbut stiil six times as long as thick; flagellar segmentsprovided
with long, outspreading seta that arise from microscopic tubercles. In the female, the antenna areless than one-half the length of those of the male and are only shortly and inconspicuously hairy. Head
impunctate.
Thorax of normal Cylindrotomine structure. Legs normal, the tibial spurs smail. Wings(P|.2, Fig. l7) with Sc moderately elongate, extending to beyond midlength of the basal section of.Rs,both Scr and Scr preserved and subequal in length; tip of Rz entirely atrophied; ftr, z and distal
section of Rz{3 in direct longitudinal alignment; Rs very long, divided into two sectionsby r-m, thebasal section relatively short, arcuated, less than the long, straight distal section, rn being before
midlength of Rs; R+*S shorter than the distal section of Rs; Rzl3 simulating z in appearance,transverse in position, appearing to connect R.r with the end of rRs; free distal portion of Rz!3
(possibly representing rR.ralone) about three times the basal section; cell rsl Mz large, closed; cell
Mr lacking, there being three simple medial veins issuing from the cell; m-cubefore midlength of cellrsl M z 1 vein znil z{ compietely lacking. Wing-veins extremely distinct and well-defined.
Abdomen relatively slender, the hypopygium somewhat enlarged, with relatively small styli.
Type speci ec ':,Stibadoccrclla
fristitco Brunetti.
Geognephlcal distnlbution.
de Meijere, Bijdr. Vol. ar. p. r7, Pl. z, f , 7 ltgtgf (Agaslonyia). Sumatra.
r. S. albitarsrs
z. S, pristina Brunetti, Rec. Indian Mus. Vol. 15, p. 283 [r9r8] (written Assam.
erroneously Cylinfuotomalatcfurcata; Brunetti, ibidem, p. z8o).
FOSSIL CYLINDROTOMIN.€
Remankr. - The only fossil genus that has been ascribed to this subfamily that wouid appear
to really belong here is CyttaromyiaScudder, disctrssed below. The genus Oryctogna Scudder, wasproposedin r 894(Proc. Amer..Philos.Soc.Vol. 32,p. ry4) for the single species,O. sachcniiScudder, fromthe Florissant. The present writer cannot detect Cylindrotomine features in this fly and would preferto ascribe the group to one of the other subfamilies of the Tipulida. In 1916, Meunier (Zeitschr.
Deutschen Geol. Gesellsch. for tgtl, 67 i z2g-23o; fig.) erected the new genus Cfitaronyclla from theSiebengebirge Aquitanien, based on the new species, C. bastiniMeunier. This genus was based upona misconception on the part of Meunier as to what actuolly constitutes the so-called second discal cellthat is characteristic of.Cyttaronyia. A study of the rather numerous speciesof the latter genus shows

I4

DIPTERA

that the group is very close to the recent
genuscylindroromaMacquart, differing
chiefly in the possessiono f a s u p e r n u m e r a r y c r o s s v e i ni n c e l l R S ,connectingvein Rl*Swithvein Mrashortd i s t a D c eb e y o n dthe origin of the latter' This crossvein is
so consta't tl.rat in some casesit seems
topersisteven after
the atrophy of the basal deflection of'
M t, giving us a case exactly comparable
to that found in the truediscal cell (tst Mz)' where this cell
may be open by the atrophy of eitherof the two veins crosingits
outer end' such a persistenceof the vein
is found in clttaro)yia(cockerell),fusculareferred to under
this genus' Meunier's description andfigure of his new genuscyrtaromlrcila,however,shows a typical
Eriopterine or lfexatomine venation
witl a supernumerary crossvein in celi
R5, connecting with veinMr)-z immediately beyond its origin.
There isa difference between these two
tbat should not be
misunderstood' In the opinion of thewriter. clrttaromycilais a Limoniine crane-fly,possibly an
Eioccra' with an adventitious or supernumerarycl'ossvein in the cell indicated. In thissamepaper,
Meunier erectsthe new tribe cyttaromini(cyttaromyini) for the two genera,cyttaromlia
and.
cyttaromyclla.
As has been indicated above, cyttaromlia
is so ciose to cylindrorona that it is certain
that
no group
higher than the genus is required
and the new tribal name has consequently
beenplaced.
in thesynonymy of the Cylindrotomine.
Besides the new genera mentioned
above, cockerell (proc. u. S. Nat. IVIus.
vor. 57,p. 247,
34'
f' 3 [r9zo]) has described as new aPl'
cltlindrotomaocrcrauawhichmay actually belong
to this genus
or may be an aberrant cyttatotryrr'ain
which the supernumerary crossvein diagnosticofthegenus hasbeen lost by atrophy or obscured in thetype specimen. The writer has included
this species in thegenuscyttaronyiauntiTthepointcanbedecided.
Ifthespeciesshouldprovetobeatrue
clrlindrotoma,
the status of the genus cyttaromyiarvould
appear to the writer to be very questionable.

8 . G e r u u sC Y T T A R O M Y | AS c u o o e n
Cyttanornyia Scudder, Bull. U. S. Geol.

Surv. Vol. 3, p. 75t (rglil,A r f f o p r i r C o c k e r e l l ,p r o c . U . S . N a t .
M u s . V o l . 5 7 ,p . z 5 o ( r 9 z o ) .
Rcmenks. - The genus Cyttaromyiais
very close to Cylindrotoma Macquart. Scudderstatesthat the tibial spurs are lacking, and if
this is true, it would constitute a marked difierence
betweenthisg e n u sa n d t h e r e c e n tC y l i n d r o t o m i n a .
The venation is so ver)r closeto that ofcylindrotona in allregalds that the only point ofdifference
that can be indicatedis the supernumerarycrossveinin cell Ri, connecting posteriorlywith
vein Mr not
far from the o'igin of the latter, and producing
the so-called n secondary discal cell , above theprirr:ary
one that is characteristicof the genus (pt.
l, Fig. B).
Ty pe s pcc i et : Cyttaronyia
ata Scudder.
fcn cstr
Geologicel dirtnlbution.
r. C. cancdllala
Scudder, proc. Amer. philos.

(,8s+).

Soc. Vol. 32, p, rg3, pl. r, f.

C. clathrataScudder, i!id.1m_,Vol. 32, p. rg4,

f. g (rgga).

!1.-r,
3 . c.fauslrata Scudder, Bull. U. S. Geoi. .Surv._vor.
3, p.'7s-r'(rgzil.
proc.
C.
Cockerell,
4 . fuscula
U. S. Nat. Mus. Vol. 57,,p.'zSo, t"*t.ng.

7

Miocene (Florissant).
Miocene (Florissant).
Eocene (white River).
Eocene.

7,
pl. 35, f. 3 jgzol (Asitofsis).
_
5 . c. oligocena
Scudder.Proc. Amer. philos. soc.Vol.3z, p.
r93, pl. r, f. z ( rg94). Miocene (Florissant).
6 c' princctoniara,
scudder, ilidem, vol. 32, p.-r?":pl. r,-f. i
Miocene (Florissant).
1ra9a;.
C. rcclusaCockereil, proc, U. S. Nat.
Mus.-Vol. Oi,p.l, pt. ), fl'r
Eocene.
6g"a,1,?C. vctcrana
Cockerell,ibidem,Vol. 57,p..47,pl. 34,i. i7ig"ol1Cym|drotomo). Eocene.

r5

FA\[. TIPULID.IE

INDEX(genus), de Meijere.

AgettomYla

albitarsis, de Meijere(Stibailoccrolla:1.

Prger
. 1 3
. r 3

L rM N o B | N I E c y l t N D R o r o M E F o R M E s ,Schiner'

.

""t'"
6
to

(genus;, Osten-Sackm

8

Llogma9r+12

metallica, Alexander (Stibailoceru) .

. 1 3

. l o

8

neoxena. Alerander (P halaeroccta)'
nigriventris, Loew lCylindrotona) .

.

9

br n iro str i s, Zetterstedt (P halaotoccra)'bullata, Enderlein (Stibailoccra) ' .

. r onodicornis. Osten-Sacken \L iogma)

.

tt

. 1 2

nodicornis flaveola, Aiexander {Liogma) '(Phalacroeera)
tttilicornis,Schummel

cancellata. Scudder (Cyttaronyia)'
clathrata, Scudder (Cyttarotny ia\, t 4

americana. Ostea-Sacken (Cfiiniltotoma).

(Clliniltotona)
anomalo,J<-rhnson'(genus),Cockerell
ArlloPrlr' "(Stibaitrocerciles\
Alexander
atrstraliensis,
bastini, Metrnier (Clttarcmyclla).
brawcontis. Zetterstedt (Cytrinilrotoma)'.

mikado.Alexander(Phalaetocera\. 1 4

Cylladrotoma(genus)' Macquart.

8

CVLI N DROTO

ill I DE'

K6rtesz

,

6
6

.

o

CYLTN DROTOM

I N lr Scudder, Brunetti.

6

CYTTAROMINI,

Meunier.

,

6

M I NE,Gy(tencmYclle(genus), Meunier. 1 3

GyttanomYle(genus), Scudder. 1 4

CYTTAFOMYI

N l, Alexander. 1 4

tlistinctissima, Meigen (Cylinilrotona)'

ERUCEFOR.

8

M lA, Bengtsson
II

excnlpta, Osten-sacken (Triogma\'.

fenestrata, Scndder (Cyttar ony ia)
formose, Alexander (P halatocera) .

.

fuscula. Cockerell (C1thttotnyi a)'glabrata,

1

4

. r o
. r +

(genus), Schiner .

Phalacnoocne

princetoniana, Scudder (Clttaromyia)'
pristina,

Brunetti

9

. 1 4
r?

I4

sackenii, Scudder (Oryetogma)
serraticornis,Alexander(Liogma)'s1:lendens,Doate (CYlinilrotoma)
Stlbadoa.Fa (genus), Enderlein'
Stlbedoccnclla (geutrs),Brunetti
Stl badocenodel (genus),Alexander

r3

9

TriogmaTnzolc, Coquill ett (Cyli*lrotonal'.

9

trisulcata,

Schummel (Triogma)

, 1 2

.

reclusa, Cockerell lCyttarongia)'
replicata.Linneus (Phaloaoccra\'

.

'

lo

. 1 1

12

Brunetti

japonica, Alexander (Ciiailrotoma)(Triognd),(Stibailocna)

quadricellula,(geaus), Schiner 'kuwanai. Alerander, r o(stibailoocrclla)tarsalis, Johnson l.Cylinibotona)''tasmaniensis, Alexander (stiboiloac/odts)'
tipulina, Osten-sacken (Phalacroccra)

Meigen (Liogmal. r o(genus), Scudder'

Onyotogna.

DROTO m lNA, Osten'Sacken .

CYLI N DFOTO

Alexander (P halanocera)

occidentalis,oligocena, Scudder (CYttarorylia)

Alexarder

CVLIN

.

'
veterara. Cockerell \?C1tllaro41ia\

IO

II

T2
r3
I2
9
'

12
ro

'

tr
tz

'

14

I6

DIPTERAEXPLANATION OF PLATES

tr,r#l"E;L"$.i0"
R :Prerp rcvlindrotomina. (Interpreted
accordingto the comstock-Needham-

Symbols : A : Anal vein; C,, :eu5itu., M - Mediai ,r_cu:
Medial-cubital crossvein;
Radius; Rs : Radial sector j S: Supernumerarycrossveini Sc _ Subcosta.
Fig. A. Ilypothetical- type of Cylindrotomiue
wing.
Cyttoromyiaolig.lun-aScudder (after
Scudderi.
I.
C. CylindtotomasllcndcnsDoane.
D. Phalactoceraoccidcrtalis,nov. sp.
E. Triogmo creulpta Osten-Sacken.
F. Slibadoccramctallica Alexander.
G. Slibadoccrodcs
austral,iensisAlexander.
- H. Stibadoccralla
albitarsisde Meijere (after de Meijere).
Fig. r . Liogma nodicoyaisOsten_Sacken,
lateral aspect of adultmale.
2. Liogma nodiaornisOsten_Sacken,dorsal
aspect of larva.
3. Phalauoccra nllicata Linnaus, iateral
aspect of larva.
4, Phalacroccranltlcqta Linneus, lateral ".i""t of pop".
5. Liogmo nodieornisOsten.Sacken.
dorsal ".p""t "f d;;.
6. Liogma noilicornis-Osten.Sacken, basal .i* .ug*".ni,of antenna of male.
Z. Triogma c*culpla Osten-Sacken, basal six segmentsof antenna of male.
8. Clrlinfuotomosphndas Doane, lateral
aspect of ovipo.
sitor (after Cameron).
_
9. cylhdrototna sprcndcnsDoane, ventral aspoct ofmalehypopygium (after Cameron).
pretr
Wings

Fig. r.
z.
3.

-

Cylindrotona tarcalisJohnson.
Cylindrotomanigriucnlris Loew.
Cyl,ittdtotoma
jaloniaa Alexand.er.
+, Phalacroccraoccidentalis,nov. sD.
5. PhalaeroccromihadoAlexander.
6. PhalaaroccrarrcorcnaAlexander.
7. Phalaeroccrafornose Alexand.er.
8. Liogma glabrota Meigen.
9. Liogma nodiconis Osten-Sacken.

Amherst, Mass. U. S. A. l5
July, r925.

zof Cylindrotomina.
Fig. ro.
r r.
12.
r3.
r4.
r5.
r6.
17.

Liogma scrraticornisAlexander.
Triogna crcueta Osten-Sacken.
TriogtnahuwanaiAlexander.
Tiogmn trisulcataSchummel.
StibodoccramctallicoAlexander.
Stibadocerodes
tasmanicrsisAlexand.er.
Stibadoccrodcs
australicnsisAlexander.
Stibadocuclla albitarsisde Meijere (afterde Meijere).

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