D IP T E R A

I 8 8 ' " F A S C I C UL E

2e?

GENERAI NSECTORLJM
ornrcEs PAR

P. \(/YTSMANDIPTERAI .€F A M . P T Y C H O P T E RD

WITH

I

PLAIN

PLATE

1927

60-62,rue T'Kint, BnuxeLLes
Imprimeur-Editeur,
En ventechezLours Desmnr-VeRTENEUtL,
Prospectusgratis et franco sur demande

Directionscientifique..P. WyTsmln, Quatre-Bras,TenvUenen(Belgique)DIPTERA
FAM. PTYCHOPTERID,€-DIPTERAF A M . P T Y C H O P T E RI D A
By Cnenres P. ALEXANDER
WITH

I

PI.AINPLATE

S now restricted, the family Ptychopterida
Bittaconorfhella and Bittoconorfha. Until

includes only the three genera, Plycholtuo,
r9r9, the Flies that are now recognized aswere included in theconstituting the distinct and primitive family Tanyderida'fhePtychopa group to which they are not very closely allied.
Ptychopteride,
as thus restricted, form a compact group of flies that are widely distributed throughout the
The recent studiesby C rampton (Entomological News,
temperate and tropical regions of the World.
teride,Vol.37,

p. 33-39. 65-7o 119z6))indicate that the true affinities of the Ptychopterida

lie with the

Psychodoid Diptera.
Through the studies of R6aumur on the remarkable lalva and pupa of the group, the typicaland long before its diagnosis as a genus
genus Ptycholtera was known even before the time of Linneus
by Meigen in r8o3. In 1835, Westwood erected the genus BittaconoQha,based upon the so-calleda phantom crane-fly t, Tilula clavilesof Fablicius, and correctly indicated its affinitres with Ptychoftcra.
is very different fron'r that of the
Although the general appeararlceof the Flies of the Ptychopterinet h e r e i s n o ' q u e s t i o nc o n c e r n i n g t h e i r c l o s e r e l a t i o n s h i p .
system of wing-venation is used.
In the present paper, the Comstock-Needhanr-Tiliyard

Bittacomorphina,

The terminology of the male hypopygium is that of Crampton (1923).
T h e w r i t e r i s g r e a t l y i n d e b t e d t 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 f o r c o - o p e r a t i o ni nmaking the figures of wings used in this paper.
GeoEnaFhicel distnibution. - Species of the typical genus Ptycholtcraoccur throughout theHolarctic Region, being especially characteristic of the Ettropean fauna, where no fewer than ninedistinct species are found. Rather numerous species have been discovered in the Eastern Palaarctic
Region (Japan) and at higher altitudes in the Oriental Region. The species in the Nearctic Regionone
are more restricted ir-rnumber. Other species of.Ptychoplcraoccur throughout tropical Africa andtoappears
thus
genus
The
form has been recorded.. but not described, from temperate South America.
be lacking only from the Australasian Region.
the
The r Phantom Crane-Flies tt, Bittacomorphcllaand Bittacomorfha, were long knownonlyfron
Very
Nearctic Region where a single species of each genus occurs on each side of the continent'
recentlya species of.Bittacomorlhellohas been discovered in the mountains of Central Japan.

DIPTERA
Geofogicel dlstnibution.

-

The only described fossil Ptychopterida

are the PtychoPtera

tlelclaNovak (Upper Oligocene), whose strict generic position is doubtful, and. Bittacomorlhcllamioccnica(Cockerell) from the Florissant Miocene, described as a Bittacomorlha but, from its comparison with
Bittacomorfhcl,lasachcnii(Roder), almost certainly a member of thjs latter genus.
The adults of the species of. Ptychoftcla, are especially characteristic of wet swales
and open wet meadows. Bittacomor\hclla is a shade lover, the Flies being found in cold, darkened
Biology.

-lvoodlands, usually near running water or springs, in small darkened ravines or along densely shaded
runs. Bittacomorlhc frequents open pond margins and alder swamps. The larva and pupa of thePtychopteride

are very characteristic.

The larva

have the body eucephalous and metapneustic, thecautlal end being prolonged into a more or less retractile breathing-tube that is protruded throughthe surface-film while the insect feeds beneath the water. A similar adaptation is found in the pupa
where one of the two prothoracic breathing horns is enormously prolo:lged into a breathing-tube, theother being greatly atrophied.

The larve and pupe dwell in the water and saturated earth in the same

habitats in which the adults occur.
As typical of the family, the iife-history of Ptychot'tua albimana (Fabricius), as studied byTopsent, may be briefly summarized. The eggs range iu number from'5eo-587, averaging 554.
They are pale yellow, slightly arcuated, the surface curiously ornaurented, the dimensions being o.Bz5
The duration of the egg stage is seven days. The newly hatched larva measures
3.85 mn., the respiratory tube o.99 nrm. The adult lart'a measutes 77 tnm', of which the respiratory
tube constitutes zo mm. The growth of the larva is rapid. When fifteen days old, the extended
When sixty-seven days old it measures 45 mm. The pupal stage is from ten
larva measures z5 nrp.
to twelve days, averaging eleven days. There are normally two generations per year'
l-tyo.264 millimeters.

The life-histories of the American Ptycholteva rufocincta (Osten-Sacken) and Bittaconorpha cl'attilcs(t abricius) are less completely known, but the pupal duration is very much shorter. In the case of theis sumformer species it is slightly less than five days. The known biology of the Ptychopteride
nrarized in a paper by the writer (Cornell Univ. Agr' Expt. Sta. l'Iem. Nr. 38, p' 772lrgzo])'
dilated.
The species of.Bittacomort'hahave the basitalsi of all the legs, in both sexes, curiously
as theyswollen tarsi are almost completely filled by the trachere and serve to buoy the insects
fhe flies of. B. claailespresent a very curious aPpearance as they drift about indrift in the wind.
the breezes rvith all the legs extended like the radii of a circle.
'l'heseThe aduit flies of the two included subfamilies are ver)'thefrom one auother in their general appearance but the relationship existing is ver-y close,

Gheractens
differentof theAdult

Flies.

-

flies having many characters in coulmon.
triangular I
Head transverse. very closely applied to the thorax ; fronto-clypeus produced, rotrghly
equal to the others
maxillary palpi elongate, the terminal segment especialiy elongate, approximately
ventralor caudal facetaken together. Labial palpi conspicuous. labelliform, the theca at the base on the
2 o - s e g m e n t e di n t h e B i t t a c o m o r p h i n r 'c h i t i n i z e d . A n t e n n a r 6 - s e g r n e n t e idn t h e P t y c h o p t e r i n r ,pronoturn very reduced. \4lesonotum usually highly gibbous. the trausverse suture bent strongly
prascutum with parallei
caudad.at the region of the scutum to form a deep u scutal loop I ; mesortotal
furrows of the scutal
loggitutlinal furr.ows on either side of the median area, these in alignment with the
Nletathoracic spiracle close to base of halter. Mesothoracic meron fused
hairy in thervith the mesepirneron. Legs moderately eiongate, longer aud more conspicuously
in Bilqcomorlha with the basitarsi dilated; tibial spurs present; empodia shott,
Bittaconrorphinr,
cells in Plychot'tcro,
transverse. Wings rvith rnacrotrichie in the outer ends of the radial and medialloop.

prehaltere present.

FAVI. PTYCHOPTERID,€
nrore reduced in number in Bittacomorfhclla,lacking in Bittacomorfha. Venation (see Plate, Fig. A. B.)
rvitl-rSc, lacking; R.susually short; Rr.,., thickened and running close to Rr, connected with the latterby r; Rna, delicate. cell Rn always presentl cell M, present in the Ptychopterina,
lacking in the; p o s t e r i o r s e c t i o n o f v e i n C z , s t r o n g l y s i n u o u s ; a c o n s p i c u o u sl o n g i t u d i n a l f u r r o w
in cell Cz, extending from vein zod A to near the bend in the distal section of Cur. Abdomen moderately
Bittacomorphineelongate,more so in the Bittacomorphina.
Male hypopygium with the lateral lobes of the ninth
tergite and the dististyles produced into more or less elongate digitiform lobes. Ovipositor with thetergal valves strongly compressed, bent slightly dorsad, so the ventral margin is straight or even feebly
concave.
-

Ghanactens of the Lanv-.

Body eucephalous, metapneustic, long and slender. the caudal
end prolonged into a more or less completely retractile breathing tube that bears thespiracles at the tip.
Lobes surroundingthe spiracular disk small or indistinct.

Anal gills two, elongate-cylindrical, siprple.

Integument with tiny hairs (Plychollera, Plate, Fig. 2), or with slight, warty protuberances (Bittacomorfho), or with conspicuous elongate tubercles (Bittacomorlhclla, Plate, Fig. 6). pseudopods on
abdominal segments one to three each bearing a curved claw. Head complete, eye-spots distinct.
llandibles opposed.
Ghenactens

Mentum many-toothed (Pt1'chofterhrc)ot merely bilobed (Bittacomorphina).
of the-Pup-.

Usually with one of the two pronotal breathing horns greatly

elongated (Plate, Fig. 3), much longer than the body (this may not be true in the still insufficiently
known pupa of Bittaconorlhclla); in Plycholtcra and Bittacomorpha it is the light horn that is normallyelongated, the ieft being degenerated.
a l l p a r a l l e li n t h e P t y c h o p t e r i n e ,ln BillacomoQhellathe right horn is degenelated. Tarsal sheaths
the fore pair overlyingthe middle pair in the Bittacomorphine.

Abdonreu covered with setiferous tubercles arranged in transverse rows on tergites and sternites, a1d
more or iess in longitudinal rows on the pleurites.

Cauda with a powerful dorsal median iobe near thebase of segment.eight.
Phylogeny.

-

It has generaliy been assumed that our recent Ptychopterida

derived more or less directly from the family Eoptychopteridae

have been(Proptychopterida)of theNlecklenburg l.ias (lowest Jurassic), where they are represented by three monotypical genera (Eoftychoftcra
sitnflcr Geinitz, Proplycholtcra liasino Handlirsch

and Eolimnobiagcittilzi Handlirsch).

The writer is not

fully convinced of this relationship and, if it is correct, some intermediate types must have existed
that have not yet been re-discovered.

Unquestioned Ptychopteridae

Oligoceneand undoubted members of the Bittacomorphinre

were in existence in the upperin the Miocene.

Among the recent genera, Ptychopltro,from its adult structures, is the most primitive, followed
Based upon the stiucture of the larva, however, Bittacomorlhclla

by Bittaconorphella and Bittacomorlha.

is the most generallized and is very distinct from Bitlacomorpho and PQchoftara, which are generally
similar to one another. It is probable that the two recent subfamilies, Ptychopterine
andBittacomorphina,

were derived from some common ancestor in pre-Tertiary times, and the direct

derivation of Bittacomorphcllafrom Plchol>tcra is improbable.

FArtN.PTYC H OPTE R I D,IE
Ptychoptenin-

Schiner, Fauna Austriaca, Dipt. Vol. 2, p. +gS (r86a).

Ptychoptenina

Osten-Sacken, il'Ion. Dipt. N. Amer. Vol. 4, p. 3o9 (1869); Berl. Ent. Zeitschr.

Vol. 3r, p. zz6 (r887).
Ptychoptenid-

Handlirsch, Fossiien Insekt. p. 967(r9o8).

DTPTERA
p. 74 (t9to)'Griinberg, Si.isswasserfaunaDeutschlands, Dipt. zweifliigler,'I'he
ptychoptenidCrane-Flies of New York, Biology and Phylogeny, Cornell Univ'
Alexander,
Agr. Expt. Sta. Mem. Nr. 38, p.772 (r9zo)'
Lf niopida

Ptychoptenid-

Crampton, Ent' News, Yol' 37, p' 33-39, 65-7o (tgz6)'
TABLE

OF SUBFAMILIES
Adults.

Antenna r6-segmented; wings with cell Mr presetrt 'Antenne 2o-segmented;wirlgs with cell M, iacking 'r' Subfam' PTYCHOPTERIN,E.
z' Subfarn' BITTACOI\{ORPHINAi.

Larve.
Mentum with outer margin finely serrated; mandibles u'ith
three iarge outer teethl pseudopods small; eoloration
yellow or brown
Mentum bilobed, untoothed; mandibles with a single iarge
outer tooth; pseudopods prominent, each with a conspi'
cuous curved claw; coloration rusty red or black 'r. Subfam. PTYCHOPTERTN,E.

2' Subfam. tsITTACOMORPHIN,lE.

Pupe.
'
All tarsi lying parallel; wing-pads with cell Mt Present
Fore tarsi lying over middle tarsi ; wing-pads with cell M ,Iackingr' Srtbfam. PTYCHOPTERIN,'8.
z. Subfam. BITTACOMORP H IN.|E.

l . S u e r a n r .P T Y C H O P T E R I N . €Ptychoptenin-

Schiner, Fauna Austriaca, f)ipt' Vol' 2'P' 4g5 (r86a)'

ptychopterina
all present a generally similar appearance'
Rcmanks. - The adult flies of thecolors. usually contrasting strongly withbeing glabrous or nearly so, with shiny black or blue-black'lhewings are often cross-banded with brown, in such casesthere
fulvous or rufous spots and bands.
band occupying the forks of cells
being a nearly complete lascia at the cord and an outer broken
Rn and Mr.
the presence ofThe essential characters of the adult flies of this subfamily are
The larve and pupe do not differ
sixteen antennal segments and the retention of cell M, of. the wing.
does from Bittacomorphcl'la'str.ikingly from those of Bitlaeomorfla, less so than the latter Senus
Ghanactens.

-

Theonlyvalidgenusinthesubfamill'isPtycholteraMeigen.

l . c e r u u s P T Y C H O P T E R AM e t c E r ' t
Llniope NIeigen, Nouv. Class. Mouches, p. r4 [rEoo] (nomen nudem)'
Ptychoptene Meigen, Illiger's Mag. Ins' Yol' z, p' z6z (r8o3)'
Gtonocenia Rondani, Dipt. Ital. Prodr' Vol' r, p' r87 (t856)'(tgtg)'
Paneptychoptena Tonnoir, Ann. Soc. Ent' Belg' Vol' 59, p' r15

FA[,I. PTYCHOPTERID,€
Rcmanks.

-

The adult flies of. Ptycholtero may be swept from rank vegetation in and nearswales and in open swamps. The larve occur in decaying vegetable matter in rich organic mud at themargins of swamps and ponds.
Ghtnactens

of Adults.

-

Body relatively smooth, shiny or subshiny, the general appearanceof the Flies suggesting somewhat a large fungus-gnat (Mycetophilidr),
heightened by the usual black and rufotts coloration and the banded wings.

this resemblance being

Head transverse; vertex between eyes broad; fronto-clypeus large, roughly triangular, bearing
the tiny labrum at its apex.' Maxillary palpi long, the terminal segment very long and flexible,
exceeding in length the preceding three segments taken together. Labial palpi large and conspicuous,labelliform.

Antenne (Plate, Fig.8)

inserted on the anterior portion of vertex, closely approximatedat origin, r6-segmented; scapal segments modetately large; in the male, flagellar segments cylindrical,
becoming more flattened when dried, terminal segment very small; segments with a few short verticils
and with a dense covering of short, subappressed silvery pubescence. In the female, the antenne arevery noticeably shorter. the flagellar segments beyond the first short-cylindrical, the ends slightly
narrowed.

Eyes relatively large, with delicate ommatidia.

Pronotum narro$', the anterior and posterior divisions evident; head closely applied to the
cephalic nrargin of the mesonotal prescutum. Mesonotum of moderate sizel suture between theprascutum and scufum subtransverse at lateral margins but near the median area deflected very strongly
caudad, near the scutellum forming a posterior scutal loop; more or less distinct longitudinal furrows
extend from this suture onto the prescutum, occupying the region of the interspaces and being inalignment with the scutal loop of the suture. Legs of moderate length onlyl coxa elongate, especiallythe fore coxe; vestiture of legs reiatively short and delicate, appressed or subappressed; tibial spurs
large and conspicuous. hairv, except for the acute glabrous tips; ungues very small; empodia small,transvelse. Wings moderately broad, the distal fifth or more with nracrotrichia in all the cells.
Venation (Plate, Fig. A) : Sc ending in Costa a moderate distance beyond the cord, Sc, lacking; Rsvariable irr length. longest in P. contaninata(Linnr;us), short and oblique in most speciesof the genus,
in a few casesarcuated or weakly angulatedi Rz+s arising at or just before z-rz,running close to ft,,connecring with this latter vein by r which lies close to the tip of .rR,: there is a very strong probability
that this ollarcut radial crossvein is, in reality, the basal section of vein R, a condition that obtains intlre Pecliciine Tipulidei Rr*s forked, the fork a littie longerthan or subequal to the petiole; M forking
at the cord, the fork of M ,,r, shorter and smaller than that of Ra15, usually one-haif or less the lengthof its petiole i M s,+r subequal to m-cu; Cu, very stlongiy sinuate on its distal section, the semi-atrophiedCz" relatively well-defi,ned;a single persistent Anal vein. A strorrg longitudinal foid or furrow in cellCa, extending fron-rnear midlength of vein tst A to near the beud in the sinuate distal section of vein Ca ,Abdomen relatively short, the male hypopygium of moderate to large size. f'he structure of themale hypopygium differs in its details in the various species, but the tundamental plan retnaitts
generaily the same. P. japonicaAlex. (Plaie, Fig.5) rnay be taken as typical : 8th tergite short andNinth tergite with a very deep median incision, the laterai lobes thus formed being produced
caudad into digitiform, setiferous arms. Easistyle roughly triangular, with the apex rounded, beartrrg
at its tip the slender, digitiform ,lististyle. Ninth sternite verv high at base, the caudal ventral margin
narrow.

stronglv chitinized, produced caudad anti dorsad in a long, slender arm, imnrediatelv dorsad of wirich
is a shorter, more strongly chitinized arm rvith five or six blunt denticles along its ventral face',:Edeagustlouble, the parts separateexcept at base. divergent, chitinized, the outer angle produced distad
In many speciesof the genus, the lateral arms of the ninth tergite areshorter and stouterthan in the case described. The ovipositor (Plate, FiS' 4) is strongly compressed,
i11toa siender, chitinized arm.

UIPTERA
nearly glabrous, bent gently dorsad, a condition
Tri choceride.

approaching that found in

the related familv

Ghanectens of Lanv-. - Head oval to subpyriform, broadest behind, narrowed in front,
notheavily patterned abovel sete of head branched or plumose. Submentum large, usually but
a little
narrowed posteriorly, the anterior lateral angles slightly produced; menturn broader than long,theouter margin with numerous (r8-zz) teeth. Prementum with palpi rather small, lying parallel,
densely
hairy on outer or ventral face. Mandible with three powerful outer teeth and about six smaller
inner
teeth. \'Iaxilla with the cardines eiongate-triangular',with three setiferous punctures maxillarypalpi;
antenniform, cylindrical. Labrum broad, transverse, with dense tufts of sete beneath. Integument
almost smooth, without prominent warty elevations. Pseudopods low, with small claws.
Chrnactenr

of Pup-.

-Flead small, with a reduced cephalic crest. Sheaths of maxillary
palpi elongate, the tips projecting around knee joints of fore legs. Leg sheaths with all thetarsi
parallel, not overlapping one another. Wing sheathswith I\{edia branched. Pronotal breathinghorns
very unequal, one (usually the right) exceedingly elongate, longer than remainder of body, theotherhorn tiny, degenerate. Abdomen set with transverseand longitudinal rows of small setiferoustuber.cleson the segments.
On the ventral surface of the fourth abdominal sternite of the adult males of certain species of
Ptycho4tua, is found a setiferous glandular pouch. Tonnoir has proposed the generic term paraptycholtcra
for the European species showing this character (P- hondlirschi Czizel<,P.lacustris Meigen, p.longicaud,oTonnoir and P. lalustris Nleigen). 'lhe function of this pouch is not fully untlerstood.and the group,
being based on a sexual character, is not recognized in this paper, though perhaps worthy of subgenericrank under Pltchoptera. lt should be noted that a very comparable cond.ition is found. in certain
Erioptetine Tipulide(genera Paratuolcsa
and TeucholaDas)
where an entirely similar sternal pouch occurs
in the males of certain specieswhile apparently lacking in others that are seemingly closelv allied.
Type species : Plychot'lcracontaminataLinna:us.
Geognephical dirtnibution.
r . P . a f r i c a n aA l e x a n d e r , T h e E n t o m . V o l . 5 3 , p . r o r ( r 9 z o ) .
z. P. albimaaaFabricius, Mant. Ins. Voi. z, p. 323ItZSil (Tifulo).
3 . P . a n n a n d a l rBi r u n e t t i , R e c . I n d i a n M u s . V o l . t 5 , p . z 9 6 ( r 9 r g ) .
. o l . 9 , p . 1 4 6( r 9 z r ) .
4 . P . c a m u o u n c n sAi sl e x a n d e r , l n s e c . I n s c i t . N ' I e n s t V
5 . P . c a l c n s iAs l e x a n d e r ,A n n . S . A f r . M u s . V o l . r 7 , p . r 3 9 , p l . r o . f . r ( r 9 r 7 ) .
6. P. contaminataLinnaus, Syst. Nat. (ed. ro), p. 5[i6[r75g] (Tilula).

Nigeria.
Europe.
india.
Cameroun.
South Africa.
Europe.

P. JuscibcsGmelin, Syst. Nat. p. zStgfxTgz) (Tifula).

7 , P . d a i m i oA l e x a n d e r , I n s e c . I n s c i t . I \ { e n s t .V o l . 9 , p . 8 o ( r 9 z r ) .
8 . P . d i s l i n c l aB r u n e t t i R e c . I n d i a n M u s . V o l . 6 , p . z 3 z ( r 9 r r ) ; F a u n a B r i t .
I n d i a , D i p t . N e m a t o c e r a ,p . z l r , p l . 5 , f . r ( r y n ) .
P.lformosarsls
Alexander, Insec. Inscit. N{enst. Vol. re, p. +g Ugz+).
9.
ro. P. hancllirschiCzizek, Zeitschr. des Mrihrischen Landesmus., vol. 17,
p. t4 [1924] (Liriofc).
rr. P. jafonico.A.lexander',The Canad. Ent. Vol. 45, p. r98, pl. 3, f. ?,p\. 4,
f. rr-r6 (r9r3).
rz. P. lacustrisMeigen, Syst. Beschr. Zweifl, Ins. Vol. 6, p. zgr ( t 8 3 o ) .
t 3 . P . / a z l sO s t e n - S a c k e n ,B u l l . U . S . G e o i . S u r v . V o l . 3 , p . z o 6 (r8ZZ).
1 4 . P . l o n g i c a u d aT o n n o i r , A n n . S o c . E n t . B e l g . \ r o l . 5 9 , p . r z r[r9r9](Paraptycholtcra).
r 5 . P . t n i n o rA l e x a n d e r , E n t . N e w s , P h i i a d . V o l . 3 r , p . 3 ( r 9 z o ) .
t6. P. minula Tonnoir, Ann. Soc. Ent. Belgique, Vol. 59, p. rzo ( t g t g ) .

Japan.
India.
F'ormosa.
Central Europe.
Japan.
Europe.
'Western

North America.

Europe.
Western North America.
Europe.

FANI. PTYCHOPTERID,€
Europe.
1 7 . P . p a l u d o s aM e i g e n , K l a s s . V o l . r , p . U z , p l . 3 , f , r 3 ( r 8 o 4 ) .
17a P. lalud.osucocrulcaStrobl, (]lasnik Zem. Mus. Bosni i Hercegov. Sara- Europe.
jevo, Vol. ro, p. 6oz (1898).
Europe.
p.
Proc.
Acad.
Philad.
(t859).
rufocincta
Nat.
Eastern North America.
r8. P.
Osten-Sacken.
Sc.
z5z
Europe.
t g . P . s c u t e l l o r i sM e i g e n , S y s t . B e s c h r . Z w e i f l . . I n s . V o l . r . p . z o 6 ( r 8 r 8 ) .
z o . P . s u b s c u t a l l a rAi sl e x a n d e r , I n s e c . I n s c i t . M e n s t . V o l . 9 , p . 8 r ( r 9 z r ) .
Japan.
P. scutellarisMatsumura (nec Meigen), Thousand Ins. Japan, Add. Vol, z,
P. +73 17916).
zt. P. surcouf,S€guy, Encycl. Ent. Dipt. Vol. z, p. zzftgz1)(Porot'tltchoftcra).
zz. P. tibialis Brunetti, Rec. Indian Mus. Vol. 6, p.233 (r9rr); Fauna Brit.
India, Dipt. Nematocera. p. z8z, pl.6, f. r-4 (r9rz).
P. atritersisBrunetti. Rec. Indian Mus. Vol. 0. p. 234 (r9rr); Fauna Brit.
Iudia, Dipt. Nematocera,p. 283, pl. 6, f. 5-6 (tgtz).

Species of Uncertain

Algeria.
India.

Synonymy:r. P. fasciataScopoli,Ent. CarnioL.p.3zr [1763](Tifula),

Europe.

? P. p.,ludasd I\{eigen.

z. P. netallica Walker, List Dipt. Brit. Mus. Pt. r, p. 8o (1848).

Western North America.

? P. lcnis Osten-Sacken.

Europe.

3. P. nigra liabricius, Syst. Antl. p. zr (r8o5).
? P. sculellaris Meigen.

4. P. ocrcaltrSchrank. Enurn. Ins. Austriaca, p. 427 [r78t] (Tit'ulo).
? P. albimanaFabricius.
5. P.lcctinala Macquart, Hist. Nat. I)ipt. Vol. r, p.77 (1831.
6. P. quadrifusciataSay, Long's Exped. to St. Peter's Rivers, Appendix,
p. 35s (r824).
7P. rufucincta
Osten-Sacken.

Europe.
Europe.
Eastern North America.

2 . S u e r a n rB
. ITTACOMORPHIN.€
Bittacomonphlne

Alexander. Cornell Univ. Agr. Expt. Sta. Mem. Nr. 38, p.779 Qgzo).

Remanks. -has
T h e r e m a r k a b l e a p p e a r a n c eo f t h e f l i e s o f t h e s u b f a m i l y B i t t a c o m o r p h i n agiven to them the ratherappropriate nameof rr phantom crane-flies r. They areallof delicate, ethereal
buiid, with semi-transparent.unmarked wings, and with long and conspicuous black and white banded
legs that form the most evident part of the insect as it drifts easily in the wind with all the legs radiating
outward like,the spokes of a wheel.
Ghanactens. -The essential characters of the subfamill' a1" the presence in the adult flies ofelongate, zo-segmented antenna and the complete fusion of veins Mrand, Mrof. the wings. Thelarveof the two included genera are very distinct in their general appearance . Bittaconorfia possessing a iong,rusty.red larva u'ith the body-integument relatively smooth and the anal end tapering gradually into a
Bittacomorfhclla, on the contrar.v*,has a short. stocky larva. blackin color, with the body-integument covered with long, horny projections and with the short, yellow
long, partly retractile breathing-tube.
breathing-tube entireh' retractile.
TABLE

OF

GENERA

Adults.
Wings with macrotrichie in the distal cnds oJ thc radial ond'mcdiot cclls: bositarsiof hgs not dilalcd.
Wings

r. Genus BtllecoltoRPHELLA, Alexander.

without macrotrichia in thc cclls; basilarsi of lcgs
cottsl i cuouslydilatcdz. Genus Brrtecouonpne,\A/estr,r'ood.

DIPTERA
Larve.
Colorution blach, thc breath.htg-tubelight yllow;

breathing_ttfic
cntirely rctraclile; body coueredwith acry lortg projcctiorts

which are cncascd,in a blach, horny substance;nautliblcs
uith. an innar conb of tceth
Coloratiott rusty-red; body ta/cring gradually to thc long, slendcr,

r. Genus Brrtrco^tronpHEr-t.A,Alexander.

t'artly rdtraclilc, btcathing-tubc; body coacrcdwith transaclsa lous o1fshortcr, stellatc tuberclcs; maudibles without
on inncr combof tccth

z. (]enus Brrracovonpne, lVestwood.
Pupe.

Right breathhryhorr small, dcgcncratc;abdominal,
tubdrclcs
wcoh,
tiplcd with scuual slrong scto.

Risht brcathing horn clongate,fil,iforn,r. Genus BrrrecovonpHELLA, Alexander.
longcr thon thc body;

abdominol tubcrclcs strong, el,ongatc,crowud. by a airclet
of -four or fac slines and, tillcd with q sct,ifcrous
falilla.

z. Genus Brrrecouonene, Westwood.

l . G e r u u s B I T T A C O M O R P H E L L AA u e x a n o e n
Bittacomonphelra

Alexander, proc. Acad. Nat. Sc. philad., p. 5+5 (r9r6).

Remankr. The adult flies of, B. jorcsi
Johnson, the best known of the pygmy phantom
crane-flies occur in cool, shaded woods, usually near small
trout streams or near springs. In somecases the flies lurk beneath low, dark bridges and. beneathculverts. The Iarve drvell in the richorganic mud in these shaded woods.
Ghanactens of Adults- - Fundamental characters generally as
in Ptycholteru bvt the generalappearance (Plate, Fig. l) very different. the coloration
being black, more or less pruinose, with therelati'ely long and slender legs conspicuously banded withblack and white.
Fronto-clypeus large. rnol'e or less arched dorsally. the labial palpi
conspicuous; maxillary palpi
iong and slender. Anter:na (Plate, Fig,. 7) zo-segmented,
in the males long and slender, in thefemales shorter; scapal segments small, especiaiiy the second;in the males. the flagellar segments areelongate-cylindrical; basal segment very long but partly divided
near midlength by a nearly cornplete
suture to form two subequal pseutlo-segments;flagellar segments
bevond the second.gradually elongatedto the seventh. thence gradually shortening to the end of theorgan; terminal segment very small; allflagellal segrnents clothed with a short but dense pale pubescence.
Antenna of female shorter than in
the maie.
Thoracic structure ver-y simiiar to that of.Ptychoptcra. Pronotum
much more reduced; laterocervical plates vely small- Legs relatively long and slender;
coxre moder.atelyelongate ; femora at bases
very narrow and breakiDg very easily from the trochanters;
basitarsi notdilatedbutallthe
legsprovided
witl-r long, conspicuous, suberect sete that are especialiy conspicuouson the tibia and tarsi; these seta
are black on the darl<ened portions of the legs, snowy-white on thetibial and tarsal annuli ; tibial spurs
very slender and deiicate; lrngues and empodia about asin Ptychoptcra. Wings approximatelv as in
Ptychol>tera,
the chief distinctions being in the unforked conditionof vein Mt+zl distal section of vein
Cu, verv strongly sinuous. Veins Rs, R
M
Cu
much
thicker and more conspicuous than are
z+t,r.and,
veins R r + s, Rr, 'R, or M t+ z. Conspicuous macrotrichie in the distal cells of the wing, more extensiveFAN,I. PTYCHOPTERID,€

9n B. joncsi (Johnson) and B. sackcnii(Rcider) where nearly the outer sixth of the wing is included. very
reduced in B. niplonazsls(Alexander), rvhere they are restricted to a few trichie in cells zo4 f t r , R s a n d . R r .
Abdomen long and slender, the general structure of the male hypopygium and the ovipositor'
approximately as in Plchoptcra.
Chanactens-

of Lanv-.

Body short, covered with very prominent projections u,hich are

longest or-rlateral and caudal parts of body (Plate,curved claws.

Fig. 6). Pseudopods prominent, with very lar.ge,
Breathing-tube short, entirely retractile within body. Head subquadrate, the forarnen

ventral in position; sete of hehd unbranched.
irilobed, cephalic margin untoothed.
Chantctcns
middle tarsi.

iVlandible with an inner comb of teeth.
Coloratiorr black; breathing-tube light yellow.

Mentum-of Pup-.

Right breathing horn very short, degenerate. F-ore tarsi overiying
Tubercles on abdomen moderately elongated, not crowned by a circlet of spines but

tipped with a long seta.
Type specles z Bittaconorfhclla jonasi Johnson.
Geog nephical distnibution.
r. B. joncsiJohnson, Psyche, Vol. rz, p. 75 [r9o5] (Bittoconorlho).
Eastern North America.
z . B , n i l f o n a z s r sA l e x a n d e r , I n s e c . I n s c i t . M e n s t . V o l . r z , p . 5 r ( 1 9 2 4 ) .
Japan.
3. B. sachcniiRdder, Wien. Ent, Zeit. Vol. 9, p. e3o Ir89o] (Bittacomwlha). Western North America.
Fossil Species.
4. B. mioccnicaCockerell, Bull. Amer. Mus. Nat. Hist. Vol. 28, p. z8o, f. z
Ir9 ro] (Bittacomorlha).

Miocene. Colorado.

2 . G e r u u sB I T T A C O M O R P H A
Wesrwooo
Blttrcomonphe

Westwood, London and Edinburgh Philos. Mag Vol. 6, p. z8r (1835).

Rcmenkr.

-

The curious Flies of the genus Bittacomorphaare very characteristic of alder swamps

and the wet margins of ponds.

The larve are usually abundant in decaying vegetable matter in the richorganic mud about ponds and in swamps.

In some cases the larve become scavengers and fill an

important function in sewage elimination.
Ghanactens-

of Adultr.

The characters are generally similar to those of. Bitlacomorphclla,differing in the following essential features: Basitarsi of all legs in both sexesconspicuously dilated intoa hollorv, fusiform structure (Plate, Fig. 9) that is covered with conspicuous seta.

Wing-membranewithout rnacrotrichia in the cells.
Chanactens-

of Lanv-.

retractile breathing-tube.

Form elongate, body gradually narrowed behind into a partly

Integument with transverse rows of tubercies.

segments one to three prominent, with large curved claws.

Pseudopods on abdominal

Head subpyriform, the dorsurn with rowsof consl-licuousblack spots that converge behincl.

tr{andible stout with a single powerful outer tooth.

Mentum biiobed, the anterior rnargin not toothed.

Coloration of body rusty red.

Ghanactens of Pup-.

-

Right pronotal breathing horn very elongate, the left very small andshort, subdegenerate. Fore tarsi o'i'eriying middle tarsi.

Tubercles on abdomen very long, located on

IO

DIPTERA

broad transversebands of chitin, each tubercle with a starof four or five spines surrounding the apex
which bears along seta.
Ttrpr rpcclct: Bittuonorlk

olaailx Fabricius.

Gcognrp hlcel dlrtr.l tufl on.
t. B. cladlcs Fabricius, Spec.Ins. Vol. e, p.
4o4[r7gr] (Ti!ila).
' z. B. occ'iilattc&s
Aldrich. psyche; vol.7, p. zot
1iA9Sy.

Eastern North America.
Western North America.

FOSSIL PTYCTIOPTERIDEThe only fossil Ptychopterida
that have been recorded are the pecholtcrc delata
Novak(oligocene, Krottensoe) and' Biltmototlhclla mioce*,iea(cockcrell) which has been discussedunder the
genus Bittrcormrfhcllo. According to Scudd,er, Novak's
species (Sitzber. Akad. Wiss. Wien, Vol.
76,
p' 88' pl' z,f' r(877)is certainly referableto the Ptychopterida
but is probably not a true pectnfrua.

FAM. PTYCHOPTERID,E
INDEX

africaaa, Alexander (P tyaho!tcra't
albimaaa, Fabricius (.PQchoflera).
annaudalei, Brunetti (P tjtoholtcra),

Pagcr
6

(geaus), Westwood

6

6
6

alritarsis, Brunetti lP$oholtcra).
BlttacomonphaPrgctlongicauda, Tonnoir (Ptycholtcrd)
metallica, Walker (P tltcho! tera)
miuor, A feran d,er(P tycholtcrd).

6

minuta, Toonoir (Ptlchoptcro)o98

miocenica, Cockerell (Bittacomorlhella).

(genus), Alexander

BITTAGO MORP H I NE. Alexander.

7

aigra, Fabricitrs \PtyehQtcrt) .
nipponensis, Alexander (Bittaeomoehcllal .

camerouneosis, Alexander (P $choptera\ .
capeosis. Alexauder (P leholtcra)
clavipes, Fabricius (Bittacomorlha1

6
IO

contaminata, Linneus (Pttcho|tt/al

6

Bltteocmcnpholla

4daimio, Alexander (P troho?h/ a).
deleta, Novak (Ptjlcholtaa)

6
IO

distincta. Brunetti (Pgcholtera).

6

occidentalis, Aldrich (Bithtconorlhd)
ocreata, Schrank (P tychopterc).

formosensis, Alexander (Ptlohoftera) .
fusci!cs, Grrelin (Ptltohollra).

6

7

paludosa cerulea, Strobl (Ptycholteta) .
(genus),Tonnoir .

PenaFiyohcptora

4

pectinata, Macquart (PQahoftra)

PTYCHOPTERIDE,
7
6

IO

paiudosa, Meigen (P tyaholtcr a)

PROPTYCHOPTERI DE .
Ptyohoptcna (genus), Meigen

EOFTYGHO PTERI DE .
fasciata, Scopoli (P g eho!t eru)

7
g6

(genus), Rondani.

Gtcncocnie

9r Io

3

.

Crempton
PTYCHOPTERIt

Handlirsch, Alexander,
314

llA, Ostea-Sacken.

PTYC H O PTER I NE,

Schiner.

3314

geinitzi, Handlirsch (Eolimnobia).

quadrifasciata, Say (P tlchof tcra)

handlirschi, Czizek (P tyoholttcra)

rufocincta, Osten-Sacken (Ptleholtcra) .

?

sackenii, Rdder (B itacomor4hellal.
sculcllais, Matsumura (Ptjtcholtera)

9japonica, A'ferander (Ptlchoftaral .

6jonesi, Johnson-(Bittucomor/hella)9lacustris, Meigen (Ptyohoptera),6leais, Osten-Sacken (Ptlehoptera) .
liasina, Handlirsch (P roltycholtcra)
Llnlope (genus), Meigen.

+scutellaris, Meigen (Ptlchoftera)

Llnloplda(genus), Griinberg .

3

6

a)
simplex, Geinitz (Eo!t11choltcr
subscutellaris, Alexander (P tychoftcra),

3

surcou6, Sigrty (Ptycholtcra)n4

tibialis, Brunetti (Ptyehoptora).

DIPTERAEXPLANATION OF PLATE
venation of the Ptychopterida.
System.)(Interpreted according to the comstock-Needham-Til\rard

Symbols. - A - Anal vein ; C : Costa
I Cu :Cubitus;/:fold; i : humeral crossvein;
M : Mediai ,rr-cu: medial-cubital crossvein_
R
Radius ; .Rs : Radial sector t_rn _radial_medialI
I
crossvein.
Fig'r ' Bittacomorrhciloj owsr

Johnson, lateral aspect of adult mare.
z. Prycholterorufocinctaosten-sacken, d.orsalaspectof larva.
3, PtltcholtcrarufociactoOsten_Sacken,lateral
aspect of pupa.
4. Ptltcholterajay'onicoAlexander, raterar aspect of femare ovipositor.
5' Ptyahoptera
jalonica Alexander, raterar aspect ofmale hypopygium.
6. BittacomorlhcllajarcsiJohnson, dorsal aspectof larva.
joncsfJohnson, antenna of male.
7. Bittacomorphclla
8. PQcholrerorqfocincta
osten-sacken, antenna of male.
9. Bittaaomorlhaol,adfesFabricius, posterior leg.
ro. Ptycholtcrajalmica Alexander, wing.
r t. Plcholtua lformoscnsl.s
Alexander, rving.
rz, Bittacomorlhcllasa+heaiiR6der, wing.
13. BittacomorphcllariploncnsisAlexand,er, wing.
14. Bittacomoehclla
jonev Johnson, wing.
r 5. Biltacomorfhcllaclaailcs Fabricius, wing.
A. Pecholhra albimanaFabricius, venation.
B. Bittocomorlhooccidcnfa&s
Alclrich, venation.

Amherst, Mass. (U. S. A.),

June r, 1926.

DIPTERAGENERAP,riupbvJq@@
Alei,
ovtpofltur

mlfoctrete
PNuDhre'O.S..Zuvtu"-Bit/amndvh&tuJbnsgt
JohN,,laru

ZlarmuphzllaTw:
Jolw.,{Mtam

FAM. PTYCHOPTERIDA.