2lo.

I 8 9 ' I EF A S C T C UL E

DI PTERAGENERAINSECTORI.JM
D IRIGES PAR

P .V Y T S M A N

DIPTERAFAM.TANYDERI D.IE
By CH.rnrnsP. ALEXANDER
W'TH

I- PLAIN

PLATE

t927

En ventechezLouls Desmnr-VeRTENEUTL,
60-62,rue T'Kint, BRuxeuues
lmprirneur-liditeur,
Prospectusgratis et franco sur demande(Belgique)
Directionscientifique.' P. Wvrsmlx, Quatre-Bras,TnRvueRnr.t

DIPTERA
FAM. TANYDERIDADIPTERA
FAM. TANYDERID,E
By Cxanres P. ALEXANDER
WITH

I

PLAIN

PLATE

were for many years placed in the older
HE Flies included in the family Tanyderidea group to which they are now known to be rather
and better-knownPtychopterida,
distantly related. The first member of the family to be described was the long-lostNcmoloQusMacquart (r838) which is now placed in the subfamily Bruchomyiine.
The second genus to be defined was the fossil MacrochilcLoew (r85r), followed in quick successionby
ProtollosaOsten-Sacken(1859) andTorydcrzs Philippi (1865). The first recognition that members of
the family should be elevated to a higher rank came in r88o when Osten-Sacken(Verh. Zool.-bot. Ges.
using the latest genus to be
Wien, Vol. 29, p. 5q-5zz [t8Zg]) proposed the term Tanyderina,
proposed rather than one of the earlier ones. The group was accorded full family rank by the writerin r9r9 (Cornell Univ. Agr. Expt. Sta. Ithaca Mem. Nr. 25, p. 8S3). Recent studiesby Crampton andare most closely allied to the Psychodide, theEdwards have shown that the Tanyderide
discussedat this time forming a close connection between the two grouPs. The
Bruchomyiina
relationshipswith the Ptychopteride are likewise evident.
The members of the family constitute a small, palreogenic aggregation of species that occul inall the major regions of the World rvith the exception of the Oriental. Becauseof the excessiverarityof almost all of the known species, it is impossible to predict where or when new forrns will be
discoveredand it is quite possible that eventually they will be taken in the Oriental Region. Flies ofthis family have been taken in numbers only in New Zealand (Tarydcrus)and in Argentina (Bruchonyia)'
Gcologicel dittnlbution. - As mentioned above, the second genus of the family to be
characterized was the remarkable fossii MacrochilcspectrumLoew, described from tsaltic Amber(Oligocene). In r9ro, Handlirsch (Contrib. Canad. Palaont. Vol. z, Pt. 3, p. rzz-ri'3, fig' 3o)tcrtiaria from beds that were
described a new genus and speciesof Insect under the name Etoptycho2taa
supposedto be of Oligoceneage. The writer cannot believe that this Insect is a true Tanyderid,since it differs notably in the following features: The long fusion of veins R. and rRu,this fusion being
longer than the free branchesbeyond the fork; cell rsi M, is very long; two distinct anal veins' Two

DIPTERAs p e c i e so f . N e t n o l a e z r s f r o m B a l t i c A m b e r h a v e b e e n d e s c r i b e d b y M e u n i e r ( r 9 o 5 ) a n d E d w a r d s ( r 9 z l ) 'known and it is interesting to note that
The above are the only fossil records of the family so far madethey all pertain to the Oligocene.
discovered with
The immature stages of members of this family have never been
in detail and figured by the writercertainty. The supposed larva of.Protoplasafitchii was discussed
butits identity has never(cornell univ. Agr. Expt. Sta. Ithaca MeDr. Nr. 38, p. 769-772, Pl.r3 [r9zo])
its combination of characters; the body isbeen confirmed by rearing. The larva is very remarkable in
The terminal two segments of theencephalous; integument smooth and shiny white in color.
that is distinctiy fi've lobed at the tip'
abdomen are produced into a stout, non-retractile bleathing-tube
branched tracheal gills' The head is
At the base of this tube are borne two very large, pinnately
These very curious larve4)'
provided with numerous, scattered, multisetose punctures (Plate, Fig'
were found in saturated decaying maple logs.
are likewise but little known. Bruch (inlitt') discussesthe
The habits of adult Tanyderide
habits of Bruchomyraas follows :earth and in crevices of rocks'
n The flies always occur in damp places, being found on damp
thepieces of wood, roots, moss and other substances in these situations were examined critically but
and the larvr could not be located inearly stages could not be discovered. No plant-mines were foundof rest, the adult flies sit rather
the mud and wet earth along the margins of streams. ln a position
nrale rests on the female' their bodies
high-legged, the wings held obliquely divergent. In copula, thetake flight readilv when disturbed'
not being held in opposition. The flies are rather waly andwatch such a disturbed fly in flight and see
However this flight is of brief duration and one can readily
it alight again. I
osten-Sacken' near
Thomas R. Harris nade the following observationson Tonydcrusforcit'atus
a I found them hanging on small
the base of Mount Ruapehu, in the Nortl-r Island of New Zealand'
there having been lreavy rains and thebushes near the edge of the creek. The weather was rough,on one nightthey werefairl)' numerous'
creek was in flood. I got them only on two or three nightsbut
or two to the bushes, mostly low dorvn' I
They were sluggish and easily caught while hanging by a leg
were a few individu als of' Protoflasa
The only specimens of the family ever seen alive by the writer
the Sacandaga River' New York' in r9o9'
ftchii sweptfrom rank vegetation along
Biology.

-is the most generalized of the- The family TanyderideGhenectens of the Adult Flles.
the other families of crane-flies by the
recent families of Tipuioidean flies. They are readily told fromsector forking dichotomously' The twopossession of five radial veins, the four branches of thevery considerably from one another in theirsubfamilies considered by the writer as belonging here differ
are small. verv hairy flies that bear a considerable
general appearance. The Bruchomyiin€
to the typical members of the family
resemblance to Tipulid flies of the genus Molophilus or
have the eyes prominetrt, naked and markedly approximated
The Bruchomyiine
Psychodida.
of either 16 or 3o segments' Maxillaryon the vertex. The antenne are eiongate, setaceous,composed
than the preceding segments' Pronotum
palpi with the terminal segment elongate and more slendergibbous'
Laterocervical plate small' Nlesonotal prescutumsmall and inconspicuous, collar-like.
Tibie
Diptera'inotherbetter developed thanprojecting slightly over the head. Metanotum muchthat simulate spurs' Wings with the anal arrgle
without spurs, though otten with large powerful seta
Male hypopygium with the tergite twisted
lacking, the veins in this region being greatly reduced.
abdominal sternites. ovipositor with small
through rgo degreesso it lies on a plane with the remaining
fleshy valves that are retracted within the preceding segment'

FAM. TANYDERID,€
The Tanyderine,on the other hand, are almost invariably large, subglabrous flies.

The eyes
The mouthparts are generally short but in some cases greatly produced.
with a variable number of segments, ranging from r5 to at least z5 in the known species.

are densely short-hairy.
AntennePronotumsptrrred.

more massive.

Laterocervicalplate large to very large. Metanotum reduced. Tibia
Wings with a more or less conspicuous anal angle; rz present, closing the long cell rsl Mzi asingie well developed anal vein.

Supernumerary crossveins occur in various cells of the wings in many

species. iVale hypopygium not twisted, of primitive structure, there being a single dististyle.

Adeagusdividing into two or three elongate, slender prongs, each of which is penetrated by an opening, theenlire structure suggesting the condition found in the Tipuiid subfamily Cylindrotomina.
Ovipositor.
with fleshy valves.
Phylogeny.

-

The Tanyderid flies are presumably the most generalized of the living Diptera.

It appears probable that both the Psychodide

and Tanyderide
may have been derived from anfrom certain of the Bruchomyiine forms, as Bruchotttjia or
NcmolaQtts. Since no members of the famih' are known from geological formations that antedate theancestor that was not greatly different

Oligocene, we have no hint as to their ancestry front this source.

It is interesting to note that theoldest fossil Tanyderids known are very close to existing genera.

TABLE

OF SUBFAMILIES

Small flies with the body and wings densely hairy; eyes naked;
pronotum collar-like, inconspicuousI laterocervical plate small I metanotum large; tibie rvithout spurs; anal angle of wings lacking, the analand cubital veins correspondingly crowded and reduced; rz lacking,c e l l . r s lM , b e i n g o p e n ( P l a t e , F i g . a , 2 ) .

. r. Subfam. BRUcHoMyIIN,ts.

Larger flies with body and wings subglabrous or sparsely hairy;eyes hairy; pronotum more massive; laterocervical plate moderately
elongate (Mauoclrilc, Protollasa)to very elon gate (T anydarus);metanotum
reduced; tibie

spurred; wings witha conspicuous anal angle;

a r p r e s e r r t ,c l o s i n g c e l l r s l M , ( P l a t e , F i g . 6 - 1 3 ,l6)z. Subfam. TANYDERINTE.

l . S u B F A MB. R U C H O M Y I I N . €Bnuchomyiin*

Alexander, Ann. Amer. Ent. Soc. Vol. r3, p. 4o3 (r9zo).

Ncmopalpin-

E d w a r d s , A n n . N { a g . N a t . H i s t . ( 9 ) , V o l . 7 , p . 4 3 g$ g z r ) .

Phlebotomina

Tonnoir, Ann. Soc. Ent. Belg. Vol. 62, p. n7 $9zz).
Crampton, Ent. News, Vol,37, p. 33.39, 65-7o(tgz6).

Br.uchomyiln-

Remtnks. - The subfamily Bruchornyiine
w a s e r e c t e d i n t g z o t o r e c e i v e t h e n e w g e n u sa n d
species, pruchomyiaargentina. The follovvingyear, Edwards noted the relationship of the genusto thelong-lost and misunderstood NemopaQusXllacquart and called attention to the fossil genus Paloosycorax
Meunier which is now known to be a synonym of.NcnofaQus. For this gtoup, Edwards proposed theterm.N e m o p alp i n a. basing it on the earliest described genus. In rgzzr 'f o nn o i r placed the two genera
above discussed, together with Phlebotorzas
Rondani,Phlebotomine.
famiiy Psychodida,

In t9e6, Crampton

in the family Psychodidre

under the subfamilyr e c o g n i z e dt h e s u b f a m i l y B r u c h o m y i i n e ,which assignment now seems to be more nearly correct.

p l a c i n gi t i n t h e

Because of its many

DIPTERA
annectant characters, however, its seems desirable to discuss the group in the present connection. The
peculiarities
venationof thewingsisverysimiiartotheconditionfoundintheTanyderina,theonly
being such as are produced b_vthe loss of the anal angle and the consequent reduction and atrophy of
the anal and cubital veins. The distribrrtion and arrangement of the veins in the medial andradial
fields is quite the same in the two groups.
-

Fig. 3) having the general appearance of a Mololhilus(Tipu lidre) or a typical member of the Psychodida; body and wings covered with long, dense hairs.
Eyes naked. Antenne setaceous, with 16 (Ncmofalpus) or 3o (Bruchonyia) segments, the terminal
Ghenectens.

segment very small.

Small flies (Plate,Pronotum small and inconspicuous.

Mesonotum gibbous.

Metanotum large.

elongate; tibia without spurs; tarsal claws small. Wings with Sct present or
atrophied; R and M forking far back near the wing-base, the forks of all the veins thus being very deep ;crossvein zr lacking; anal vein short or subatrophied ; anal angle of the wings lacking, this region fringed
Legs with the coxr

Male hypopygium twisted to r8o degrees so the tergite lies on a level withthe sternites of the remainder of the abdomen; dististyles small and partly concealed (Bruchomyio) orwith unusually long sete.

larger and exposed (NcnoPaQus).

OF GENERATABLE

r. Antcnna with 3o scgmcnts. Distal sccliot of ucin Ca clongate,rearlyr. Genus BxucnouYll
as long as thc basalscclion(Plate, Fig. | )recurttcd'
Antemrc with 16 scgmcnts. Distol secliott of acin Cu short,

Alexander.

z. Genus Nnuoper-pus \{acquart.

to the margin (Plate, Fig. 2) .

l . G e r u u sB R U C H O M Y I AA t e x a r u o e R
Bnuchomyia Alexander, Ann. Ent. Soc. Amer. Vol. 13, p. 4o3 (tgzo).
Ghanactens. -Body (Plate, Fig.3)

very hairy, the head. thoraxandabdomenprovided with

Rostrum but stightly produced; palpi very large and conspicuous. very hairy,
apparently four-segmented. Antennr 3o-segmented; flagellar segments z8 in number, the last segment
long, dense, erect hairs.

minute; scapal segments subequal in size, small, subglobular, the second with a circlet of short,verticillate hairs; flagellum setaceous, the first segrnent about as iong as the following two taken together;flagellar segments 2 to 20 subequalrelongatecylindrical; remaining segmentsgladuallyreduced in length,
Eyes large, naked;
the last very small, button-like, the penultimate and antepenultimate oval.
ommatidia moderately large; eyes approximated. separated on the vertex only by a narrow strip that isfrom one-fourth to one-half the diameter of the basal segment of the scape. Legs comparatively stout;
tarsal segments gradually shortened, from the firstto the fifth; tibia provided with long, conspicuous seta, the sclerite terminating in one or two that are
more slender and simulate tibial spurs; claws small, subappressed to the end of the tarsus. the edges
coxr long and slender; femora shorter than tibie;

roughened, the rather long apex smooth.

Wings with no anal angle; veins and margins with abundant

long sete that are especially elongate and conspicuous in the region of the anal angle ; at the union ofr.m with Ru is a triangular chitinized area that is provided with conspicuous black hairs. Venation : Sc,
lacking, Sc, ending in .R before the fork of R2 ..,.3; Rs originating close to wing-base, slightly proximadof the tork of.M; Rs with four branch es; M forking far back near wing-base, each fork with two branches;
zr lacking; Cz simple, the distal section elongate; rz-ca conspicuous, without macrotrichie; a single,
semi-atrophied anal vein.

Male hypopygiun-rsnrall, inconspicuous, densely protected by seta; basistyles

FAM. TANYDERID.E

5

stout, each with a densebrush of stout black seta on mesal face near apexl the single dististyle is small,bifid.

Ninth tergite (the apparent sternite) produced into a flattened lobe, the apex bJfid.
Type tpaciest, Bruchomyiaargcntina Alexander (Plate, Fig. l, 3).

Geog naphlcal dirtnibution.
r . B r . a r g e n l i n aA l e x a n d e r , A n n . E n t . S o c . A m e r . V o i . 1 3 , p . 4 o 5 , p l . 3 2[r9zo] (Plate, Fig. |, 3).

Argentina.

2 . G E N U s N E M O P A L P U SM n c g u A R r
Nemopalpus

Macquart, Dipt. Exot. Vol. r, Pt. r, p.85

(1838)1 Becker, I\{itt. Zool. I\{us. Berlin,

Vol.4, p.Zr-72 (r9o8); Tonnoir, Ann. Soc. Ent. Belg. Voi.6z, p. rz5, plate; Key to known
species (rgzz).
Pal-oryconar

Meunier, Miscell. Ent. Vol. r3, p. 5o (r9o5).

Chanactens.
appressed.

-

Body densely hairy, the sete on head and thorax erect, on the abdomen

Head relatively small, mouthparts not prominent;

palpi elongate, 4-segmented, theterminal segment much ionger and more slender than the preceding segments. Antenne r6-segmented,
approximately as long as the body; basal segrlent of scape short-cylindrical I second segment cyathiform ;flagellar segments elongate, densely provided with conspicuous verticils.
Eyes large, naked, strongly
approximated on vertex.
seta.

Thorax gibbous, the prescutum with longitudinal rows of conspicuous erectLegs of moderate length, hairy; tibial spurs lacking.

Wings with the anal angle lacking ; Sc,

generally present, though sometimes atrophied; base of R hypertrophied; Rs originating near basal
quarter of wing; cell.R, shorter than (N. pililcs) to longer than (N. faaus, N. zclondia) its petiole; r-zr
far beyond the fork of.M ra2(N. p;li{x) orlying at orproximad of this fork m lacking; m-culong; distal
section of Cu much shorter than the basal section and attaining the wing-nrargin
one-fourth the length of the *'ing; anal vein reduced.

suddenly at about

Male hypopygium twisted upon itself throughrSoo so the ninth tergite lies on a plane with the remaining sternites of the abdomen; a single
conspicuous dististyle that tends to be bifurcate.

Ovipositor with fleshy valves that are retracted in theeighth abdominal segment.

Typc specier : N. flaausMacquart.
Geognaphical dirtnl butlon.
r. N.flaaus Macquart, in Webb & Bertholet, Hist. Nat. lles Canaries,Ent.
D i p t . p . r o e ,p l . 4 , f i g . a ( t 8 3 8 ) ; D i p t . E x o t . V o l . t , P t . r , p . 8 2 ,p l . r z . f i g . r , t a , r b ( r 8 3 8 ) ; B e c k e r ,M i t . Z o o l . M u s . B e r l i n ,V o l . 4 ,p.7r-72, pl. z, fig. z8 (r9o8).
z. N.molophilinusEdwards, Ann. Mag. Nat. Hist. (9), Vol. 7, p. +37, fig.
[r9z r] (P alaosycoroz).
3 . N.lilipcs Tonnoir, Ann. Soc. Ent. Belg. Yol.6z, p. r3o, fig. r-8 (rgzz).
+ ' N. tutiario Meunier, Miscell. Ent. Vol, 13, p. 5o [r9o5] (Patroosycorar).
J . N. zcland,ia
Alexander,Insec. Inscit. Menst.Vol. 9, p. r58 (r9zl); Tonnoir,
Ann. Soc. Ent. Belg., Yol. 62, p. r34, fig. g-tt lrgzz) (written- Plate, Fig. 2.
zcalandicus).

Canary Islands.

BalticAmber.
Paraguay.
Baltic Amber.
New Zealand.

DIPTERA

2 . S u e r r n r .T A N Y D E R I N , €Tanvdcnf na osten-Sacken,Verh . zool.-bot. Ges. wien, yol. zg,p. 5r7, fig. vz (rggo).
Tanydcninr

Alexander,Ann. Ent. Soc. Amer. Vol. 13, p. +o2 (r9zo).

Tanvdenld-

Crampton, Ent. News, Yol. 37,p. 33-39, 65-7o(t926); Bull. Brooklyn Ent. S-oc.yol. zt,
p. r-r4 Qgz6).

Ghlnectcnr. - Size usually large, the smallest speciesbeing members of the genus protollasa(P. bechcri,P. uand,*zcci).Rostrum moderately to excessivelyproduced, the palpi conspicuous. Antenne
with from r5 to z5 segments. Eyes with short, erectsete between the ommatidia.

Pronotum massive.
Laterocervical plates elongate (typical Tanydcras, Mischodcrus and. Radinodcrus),shorter and wider(Proloflasa), or greatly reduced in size (Mauoohilc, Pbiaguqomyina). Metathoracic spiracle close to baseof halter. Mesothoracicmeron fused with mesepimeron. Tibial spurs present. Wings usually witha well-developed anal angle; five radial, four medial, one cubital and one developed anal vein presentI
rz present.closing the elongatecell l"r Mr; supernumerarycrossveinsincertainof theradial cells of thewings (Tanydarus; subgenera Tarytd,crus,Mischod,erus,
Notlcoilcr*s and Neodcru.r)or in cell M, (protoflasa,s. s.). Male hypopygium with the dististyle simple, cylindrical (TaryrdarasrPirittguqomltina)
or more orless bifid (Macrochilc,Protoflasa);adeagus trifid, in Macrochile, more bifid.

TABLE OF RECENT GENERA
r. Rostrum alorgale, excceilingthc combincdhcad and lhorax, thc
ntouthfarts bornc at lhc cxtrcmc apcx; wings immacalotc;
dististyk of nole futlolltgiun sdmflc,aeryrclongate(Pfate,Fig. 16-18) .
I. Genus PfnrucueyoMyrNAAlexander.
.
Rostrun shortcr than thc hcad; wingslicturcd ; d,ististytcof nale
hypolygium shortz. Ccll R, rclatively short, R2+s forhirry bcyotd mid,lengthof thc
dislal scctiortof aein Rr; lronotum oid lotcroccnticolflates
rclotiacly short, not forming o consficuoasncch; disthtylc of
malclrypofugiumnore or lessbifd (Plate, Fig. | | - | 4) . z. Genus Pnoropr,rsr Osten-Sacken.
Ccll R, rclatiacly dec!, R2atforhitg just bqtoxd the bascof(Radinodcrus)lo just beforcmidlcngthof, thc distal sactionof
Rr; lronotum and,Iatcrocaraicalplates (ancft in Nothodcrus)
long and,consficuous,
fortning o conslicuottsnech;dististylcof
malchypolyghtmsimplc,
eylindricol(Plate,Fig. 6-lO, l9). 3. Genus Texyoanus Philippi.

I . G e r u u sP E R I N G U E Y O M Y I NAAl e x a r u o e n
P5nfnEueyomyine Alexander,Ann. S. Afr. Mus. Vol. r8, p. z3z(r9zr).
Ghanectens. - Rostrum elongate,exceed.ing
the combined head and thorax (Ptete, Flg. 1Zl,rather stout, cylindrical, with the base enlarged, the surface with numerous, subappressed.sete, themouthparts borne at the apex; maxillary palpi slender,
4-segmented,the three basal segmentssubequalin length, the terminal segmentabout a third longer tharr the penultimate; labial lobesfleshy. transverse.

FAM. TANYDERID,€
Antenna apparently with only 16 segments, moderately elongated, setaceous; second scapal segment
swollen, subglobular; three basal segments of the flagellum stout, the remaining segments graduallyelongated, slender, provided with long verticils that are about equal in length to the segments that bear
them; in addition to the verticils, the segmentspossessa rather abundant, suberectpubescence. Headnarrowed behind. Eyes large, broadly contiguous beneath (Plate, Fig. 16), narrowly separated aboveby the vertexl ommatidia small, with short erect sete between them.
conspicuous, the posterior pronotum

very narrow.

Anterior pronotum large andLaterocervical plates very small.

Legs with themargin of the posterior coxa swollen anteriorly and provided with a row of black sete; tibie with shortWings broad (Plate, Fig. | 5), with fi,ve radial, four medial and a single anal vein attaining

spurs.

the margin ; n-cu disttnct; anal angle of wing moderately prominentmacrotrichie.

Veins with abundant long delicate

IVIalehypopygium with the basistyles very slender, greatly elongated, gradually narrowed

to the tips, each (Plate, FiB. l8) bearing a single, very elongate, cylindrical dististylethatis provided
with a series of about 3o slender spines and numerous erect sete along its inner margin, at the apex withtwo elongate bristles.
Type specict: Pirittgucyonyinabarnardi Alexander (Plate, Fig. | 6- | 8).

Geog naphicrl d lrtnibutlon.
t. P.barnordr Alexander, Ann. S. Afr. Mus. Vol. r8, p. 233, fig. (r9zr).
(Plate, Fig. | 5- | 8.)

South Africa.

2 . G e r u u s P R O T O P L A S AO s r e N . S A c K E N
Pnotoplasa Osten-Sacken,Proc. Acad. Nat. Sc. Philad. p, z5r (1859).
Pnotoplasta

Osten-Sacken, Bull. U. S. Geol. Surv. Vol. 3, p. zo7 GSIZ).

f d l o p l a s t a O s t e n - S a c k e n ,C a t . D i p t . N . A m e r . ( e d . z ) , p . 3 6 , z z z ( 1 8 7 8 ) .
Members of the genus Proloflosa are con$,ned to the Northern

Hemisphere, three species beingfrom the Nearctic Region while one is Palaarctic.
Ghanactcns. -Rostrum shorter than the head. the palpi long and conspicuous. Antenne

r6-segmented; second scapal segment enlarged, cyathiform; flagellar segments elongate-oval, withverticils that exceed the segments in iength.

Eyes short-hairy.

shorter than the head excluding the mouthparts.
Mesonotum moderately gibbous.

Pronotum lelatively short and stout,

Laterocervical plates relatively short and wide.

Legs with short, rather sparseseta; tibiat spurs conspicuous. Wings(Plate, Fig. I l- | 3) usually with a prominent anal angle; in uanduzcafthe wing is long and narrow,showing the firstsigns of atrophy.

Venation as in the subfamily; cell R2 short, shorter than its petiole.

the base o{ the cell lying beyond midlength of the distal section of vein R,; cell tst Mr long, gently
widened distally, the cells beyond it comparativelv short; a supernumerary crossvein in ceil M, in the
typical subgenus, lacking in Protaryrd,crus. NIale hypopygium with the dististyle deeply bifid in membersof the subgenus Protanydcrus(Plate, Fig. l4a,

l4b),

in Protoflasa,s.S., simple but strongly bent near

midlength andhere provided withadistinct shoulderon outerface (Plate, Fig. l4c).
Typc speciet z Protollasafitchii Osten-Sacken (Plate, Flg.
TABLE

! I , l4c).

OF'SUBGENERA

in acll M, of aing (Plate,
A suPcnwmcrarycrossacin

Fig. | | ).

in any ccllsof wing (Plate, Fig. I 2, | 3).
No suferuutncrarl/crossvcins

Subgen. Pnoropr.ese Osten-Sacken.
Subgen. PnortHvnBnus Handlirsch.

DIPTERA
Gcognephical dlrtnibutlon.
Subgenus P notoplasa Osten-Sacken.
r. P. (P.)fitchii Osten-Sacken,Proc. Acad. Nat. Sc. Philad. p. z5z [1859]. Eastern North America.
( P l a t e ,F i g . I I , l 4 c . )SubgenusPnotanydcnur Handlirsch, Ann. Naturhist. Hofmus, Wien,
Vol. 23, p. 267 (rgog).
z. P. (P.) bcchri Riedel, Zool. lahrb. Abt. f. Syst. Vol. 43, p. 365, fig. A-D Turkestan.
(Plate, Fig. l4a.)
Ir9zo] (Tonydcrus).
3. P. (P.) aaniluzcei
Alexander, Ent. News Philad. Vol. 29, p. 285 (r9r8). Western North America.
(Plate, Fig. | 2. | 4b.)
4.P.(P.)rit'ioOsten-Sacken,Butl. U. S. Geol. Surv. Vol. 3, p. zo8 lrSZZl Western North America.
(Protoflasto);
type of subgenus.(Plate, Fig. 13.)

3. Genus TANYDERUS Pxlrlppt
Trnydenus

Philippi, Yerh. ZooI -bot. Ges. Wien, Vol. 15, p.78o (1865).
-

Remanks.

The genus Taryrdcrus was erected by Philippi

in 1865 forthe Chilian T.ficta^s.

Additions to the genus have been made, more especialty in the past decade, so that fifteen species are now
referable to it.

Handlirsch (tSog) was the fi.rst to attempt a subdivision of the genus and proposed two
new generic terms which are herein considered as being of subgeneric value. Of these subgenera,
Mischodcrus Handlirsch,

includes five New Zealand species (aunul'ifcnts,forcilatas, marginatus,ncpluttus

and wrifes); Rad,inoddrusHandlirsch,

includes four species from the Papuan Subregion (mhobilis,
ocul,alus,ornalissimus and solomonls); two species (occidcntatis and terra-rcg,inc) from Australia; andone(gloriosus)from Southern Chile; while Tanydcrus,s. s., includes only the genotype, lictus, of Chile.
In order to accommodate the two remaining species, australicnsisand.latagonirus, it is necessary to propose
two additional subgeneric groups which are termed respectively Nothodcnrs and Ncodcrus (r).
It should be noted that members of the genus are confi.ned to the Southern Hemisphere.
Ghanactens.

-

Rostrum moderately elongate, about equal in length to the remainder of thehead, or, in Nolhod,a,rus,
longer than the head; maxillary palpi apparently S-segmented, there beinga
small basal segment; in the subgenera Ncod,crus(Plate, Fig. l9) and Nothodcrus the maxille are
elongate, stylet-like, and the remaining elements of the mouthparts are likewise well developed. Eyes
short'hairy.
Antenna with from 15 to at least z5 segments. the flagellar segments varying fromshort-cylindrical to elongate-cylindrical in thevarious species; the number of segments varies greatly inthe differentsubgenera;Ta*yderas hasatleaste5 segments; Radinodcrushas r8 (gtoriosus),zr(mirabilis)
or 22 segments (oculatus,otnatissimus);Neodcrushas 17 segments- Mischoderushas 16 segments t Nolhodcnts
has the minimum

known nurnber of 15 segments.

Pronotum large. Laterocervical plates usuallylong and slender, much longer than the head excluding the mouthparts; in Nothod,crasshorter and.
stouter. Mesonotum large, moderately hairy.
Legs with conspicuous, erect seta; tibial spurs distinct.

(r) NothodeFu.'
nov. subgen. Characters as in Tanyilcruss, s., but .5c2fuses with .R for a distance and thea
breaks away, appearing as a distinct oblique vein that is provided with macrotrichia.
Anteana with r5 segments,- Tanlthr*s autralicnsis Alexander (Plete, Flg, g).
Typr of rubgcnur.
Ncodcrul,
nov. subgen. Characters as in Tanyderus, s. s., but cells R3 aad R5 of the wings a supertrur1erarycrosgvein. Atrtenna with r7 seg.ments.
Typo of lubgcnur,- Tarlihrus latagonios Alexander (Platc, Flg. 8, l9).

FAM. TANYDERID,€or more or less developed, the wings
lvings (Plate, Fig. 6- | O) with the anal angle acute (fictus)
in this latter subgenus' moreover'always large and. broad; Sc, usually present, lacking inNolhodetu.s;
provided with macrotrichie; cell R, isthe tip of vein Scu is preserved as a conspicuous oblique element
so, the base of the cell being only a shortalways deep, in the species of Radinod,crrsbeing unusuallyof R243 lies before midlength of the
distance beyond the end of vein Sc,; in other groups the fork
in four of the f've subgenera, in cell Rn
distal section of vein ftr; supernumerary crossveins developed
R, and R5 irr Neotlcrus; Analin Nothodcrtts and.Tonyd.crus; incells rR, and -Rn in Mischodcrus',in cellswith a single dististyle, thisvein well developed. Abdomen moderately elongate. Male hypopygium
cylindrical, of moderate length only, and unarmed except for seta; edeagus
lacking. Ovipositor with fleshy valves.

three pronged; gonapophyses

y'dzs Philippi.
TyFe rpecier z Torytdcrus
TABLE

OF SUBGENERA

r. No su,pcrnurncldr! ctossacinsin any ccllsof wing; Rz + sforhing shorlly
Subgen. ReorNooenus Flandlirsch.
bcyonduin Sc, (Plate, Fig. 6)
2.
in radial cells of wing; ocin Rra" longcr 'Sulenrumcrary crossocins

3.

z. A sufcrnunclary crossoein.in ccll Ro onllt
Sulcnnntcrary arosstcinsin tuo radial cclls'.t.

3. Antanna with tS scgncttls. Anal anglc of wing moduately dcttel'oPed;
Subgen' Nornoopnus' nov' subgen'(Plate, Fig. 9)
til of aein Scrprcserttcd.
Antcnno with at leastz5 scgmcnls. Anal anglc of wing *cutc; ti! ofSubgen. Tenvpenus Philippi.
acitt sc, atrophicd(Plate, Fie. 7)
in' cach of cellsR" and R.n' Arttcnna tuith
4. Sttfentu.merary crossocins( Plate, Fig. I O).
t6 scgmenls
h eachof ccllsR, and Rr.
Suldrnumerary crossacitts(Plate, Fig. 8)
r7 scgmcttts

Subgen. MtscnoosnusHandlirsch.

Antcnna with

Subgen.Nnoonnus,nov. subgen.

Type speciet z Taryrduusliclus Philippi.
G e o E n a p h i c a ld l s t n i b u t i o n .
Subgenus Radlnodenus Handlirsch, Ann. Naturhist. Hofmus. Wien,
V o l . 2 3 ,p . 2 6 + ( t S o S ) .
Alexander, Arkiv f6r Zoologi,Vol. 13, P'5, fig' (r9zo)'
I . f . (R.) gloriosus

Chile.

(Plate, Fig 6.)f.(R.) mirabilisdeMeijere,Nova Guinea,Vol' 13, P'5I, fig' I (r9r5); Now Guinea.
Tijdschr. v. Ent. Vol. 58, P. ro4, fig. I (r9I5)'
Alexander,lnsec. Inscit. Menst'Vol' r3, p'32(1925)' Western Australia.
3 . f. (R.) occitlcntalis
P' t43(tgzz)' New Guinea.
+.f.in.i oailahsRiedel,Ann. Mus. Nat. Hungar, Vol r8, lndie,vol.
t7, Amboina, Obi.
Nederl.
Tijdschr.
5 . r. (n.l. P . 8ornatissimus
0 | I 8 5 8 ] ( CDoleschall,l l i n d ' r o t oNatuurk.
ma);osten.SackEn,Berl.Ent.Zeitschr.
Vol. 3r, p. 22g(r8S7);tyPe of subgenus'
Alexander, Insec. Inscit. Menst' Vol' rz, p' r43 (tSz+)' Solomon Islands.
6 . I.(rR.) solornoiis
Queensland.
Alexander,ibidem, Voi. rz, p' r4r ('S'i'
7 . T. (R.) terra-rcgina
SubgenusNothodenus' nov. subgen.
Alexander,Rec. S. Austral.Mus.Vol' 2, p' 226(tgzz)'
8 . f . (N.) australicttsis(Plate, Fig. 9.)

Tasmania.

DIPTERA
SubgenusTanydcnuc Philippi.
9. T. (f.) fichc Philippi, Verh. Zool.-bot. Ges. Wien, Vol. 15, p. 78r,
p l . z g , f . 5 7 ( r 8 6 5 ) ; t y p e o f s u b g e n u s .( P l a t e , F i g . 7 . )Chile.

Subgenus Mlrchodonur Handlirsch, Ann. Naturhist. Hofmus. Wien,
V o l . 2 3 . p . 2 6 +( t g o g ) .
ro. T. (M.) annuliferusHutton, Trans. N. Zeal. Inst. Vol. 32, p.48, pl. 4, New Zealand.
fig. zr a-e(rgoo)r Edwards, ibidem, Vol. 54, p. 57r, pl. 27, f. 7 Q9z3).
rr. T. (M.)forci/atns Osten-Sacken,Verh. Zool.-bot. Ges. Wien, Yol. 29, New Zealand.
p. 5zo, f. r-z (r88o); Edwards, Trans. N, Zeal. Inst. Vol. 54,
p . 5 7 r , p l . z ? , f . . 4 , 5 Q 9 z 3 ) ; t y p e o f s u b g e n u s(.P l a t e , F i g . 6 , l O . )n. T. (M,)narginatxs Edwards, Ann. Mag. Nat. Hist. (9), Vol. rr, p. 625 New Zealand.
(r9z3).
t3. T, (M.) ncltunusEdwards, Trans. N. Zeal. Inst. Vol. 54, p. z7z, pl. zj, Nerv Zealand.
f. 6; pl. 32, f.. rz3 (1923); (doubtfully distinct fromJorcipatas).
r a . T , ( M . ) a a r i l c sE d w a r d s , i b i d e m , V o l . 5 4 , p , 2 Z 2 t p l . 2 7 , f . 8 , 9 ( 1 9 2 3 ) .
New Zealand.
Subgenus Ncodcnur, nov, subgen.
15. T. (N.) fatagonicusAlexander, Proc. U. S. Nat. Mus. Vol. 44, p. 332, Patagonia.
f. r-3 (r9r3).(Ptate, Fig. B, 19.)

FOSSIL TANYDERIN,€

I . G E N U SM A C R O C H I L EL o e w
M e c n o a h l l c L o e w , L i n n . E n t . S t e t t i n ,V o l . 5 , p . + o z ( 1 8 5 r ) .
ldloplasta Meunier, Mon. Tipulidre et Dixida de I'Ambre Baltique, Ann. Sc. Nat. Zool. Paris (9),Vol. 4. p. 39o(r9o6).
Mrcnochile

Crampton, Ent. News, Vol. 37, p, 33 (19z6); Bull. Brooklyn Ent. Soc. Vol. zr, p. rQsz6).
Rsmanks. -This interesting genus was proposed by Loew for the oniy known species,
M. sfcctrum. Osten-Sacken and Meunier believed that the fly was identical with the recent genus
Protollasa Osten-Sacken, but this is certainly not the case. Osten-Sacken placed the name in thesynonymy of. Protoplasaunder the belief that Macrochilewas not available because of the earlier use
of.Mauocltila Stephens and MacrochiloHiibner. A recent study of all the available specimens of this
irrterestingfly hasshown that it is curiously annectant between the subfamily Bruchomyiina
and thetypical Tanyderine.

The detailed papers by Crampton on the affinities and morphology of thegenus, cited above,should be consulted.
Chanactenr. - Mouthparts elongated.somewhatshorter in the femalethan in the male; rostrum
about one-half longer than the head, the very long labial palpi nearly twice as long as the rostrum;
maxillary paipi long and slender, 5-segmented, the basal segment short, only one-third the length of
the second; third segment neariy one-half longer and more slender than the second; fourth segment
about one-half the third; fifth segment long and slender, a little longer than the second and about onehalf lorrger than the fourth.
Antenne rg.segmented, the flagellum long, setaceous,the segments cylindrical, with short,scattered sete and sparse subbasal verticils that do not exceed the segmentsin length; flagellar segments
decreasing very gradually in length and diarneter to the last, which is a trifle longer than the penultimate.

FAM. TANYDERIDEII

If bent backward, the antenna would extend to about opposite midlength of the third abdomirral
segment. Head relativoly small. Eyes large, rounded, with small ommatidia, betweenwhich project
short, erect seta. Anterior vertex reduced, the males being holoptic, the femalesdichoptic.
Pronotum and laterocervical plates much reduced, inconspicuous. Halteres very short andstout. Legs with the coxa elongate; irochanters small I femora relatively short and stout, clothed withshort, suberect,black sete; tibia slender, the spurs long and conspicuous; tarsi with the segments
gradually decreasingin length and very slightly in thickness,each segmenton ventral face terminating
in a small spine; tarsal claws small, smooth, erect. Wings with,Sc relatively short, Sc, extendingto
about opposite one-third the length of R2as, Sc, a little longer than Scr; Rs a little longer than .Rs.L3,
gently arcuatedat origin; cell R, variable in length, in most casesa little more than twice its petiole;a short spur on r-tn closeto its union with Rn..,.u,jutting into cell R; M in direct alignment with Mr+2;
cell rsl Mo long and narrow, the basal section of. My12 from one-third to one-half Rr+si n-c* atthe
fork of Ms+*i Ca, well-indicated, extending almost to the wing-margin; a single well-preservedanalvein. Anal angle of wing well-developedbut not so squarelyso as in Protoflaso. Wings without a dark
pattern.
Male hypopygium with a single dististyle, this cylindrical, narrowed to the obtuse apex. on themesal face at basewith a slendercurved arm, the apex of which is chitinized and obliquely flattened,
directed toward a small rounded knob lying in its axil, the two lobes separatedfrom one another by asrnall circular notch. Mesal face of dististyle at apex with long erect seta. ,lEdeagusapparently bifid.
Ovipositor short and blunt, fleshy, consisting of a stouter basicercusand a more slenderdisticercus,
the latter tufted with short seta.
Type specier = Macrocbile
spcctrum
Loew.
Geologicrl dlstnl bution.
r . M . s l c c t r u t n L o e wL, i n n . E n t . S t e t t i n , V o l . 5 ,p . 4 o 2 , p l . z , f . z 4 , z 5 ( r 8 5 r ) ; B a l t i c A m b e r .
Crampton, Bull. BrooHyn Ent. Soc. Vol. 2r, p. r, pl. i-z (1926).

DIPTERATNDEX
argentiaa, Alexander (Bnchomjia),

Pegcr

5

annuliferus, Hutton (Ta4dnts)

roaustraliensis, Alerander (Taayderts)

BRU CHOMyll

Z
s
4
3

NE, Alexaader.

f,tchii. Osten-Sacken (p rotolrtasa)
flav us, Macqu art (N r rru
latlus)forcipafus, Osten-Sacken (Tany&rus)
.

6
5
ro

gloriosus, dexander (Toayilcrusl
tdloplertr

9(genus), Osten-Sacten
TtIo

Mecr.oohllo(genus), Loew

maf,g'inatus,Edwards (Ta4,6sas1.
mirabilis, de Meijere (Tanytuns)
Mkohodrr.ur(subgeu.), Handlirsch
molophiliaus, Edwards (Nemopaeu)
.

9
IO
J

Edwards(genus), Macq uart(subgea,), nov. subgeo.

Ncodrnur
rfeptunus, Edwards (Tanyiltms),
Nothodcrur(subgen.), nov. subgen

occidentalis,Alerander (Taryikrus\

3

)
IO
IO
9

9

9
.

Peborycor"er(genus), Meuaier
patagonicus, Alexan d.er
(Taayhr*s)
P5nlngucyomylna (genus),
Alerander
PHLEBOTOilIN€,
Tonnoir
pictus, philippi (Taryilemsj.
pilipes, Tonnoir (Nntofae*s)
Pnctanydenut (genus),
Handtirsch.
Pnotoplaaa (genus). Osten_Sacken. .
Pnctoplarte (genus), Osten_Sacken
Rrdlnodcnlr

9
5
IO

6
3
IO

5
8
7'8
7

(subgen.), Elandlirsch

solomonis, Alexatder (T,tryikrtts).
spectrum, Loew (M acrochilc).

IO
IO

NEMOpALpINE,
Ncmopalpur.
ornatissimus,Dolescball(Taagitrts)

9barnardi, A lerander (p cringuyomy
i na)beckeri, Riedel (p rotoflasa)
Bnuohomyla (genus),Alexauder.

Prgeroculatus, Riedel (Taryilerus)

TANYDERTDE, Alexander
TA NYDE R I NA, Osten_sacken
TANYDERI NE, Alexander.
Trnydcnur (genus), philippi.
terra.reg-ine, Alexander (Taa/hr*s),
tertiaria, Meunier (N cntolalpts\.
vanduzeei.Alexander (protoplasa).
varipes, Edwards lTanyiktts)
vipio, Osten-Sackea (p rotolla sc)
zelandie,Alexander(Namofalltrs)

9

9
II

6
6
6
8
9

8
IO

8

)

r3

FAM. TANYDERID,E
EXPLANATTON OF PLATE
Fig.

r. Venatio n of,BruchomyioarganlinaAlexander.
z. Venation of NcnofaQus zelandie Alexander.
3. Lateral aspect of adult Bruchonyia afgtiltina,Alexander.
4. Supposed larva of. Protollasa (Protoitlosa)ftchii Osten-Sacken.
5. Male hypopygiutn of Tanytlcrus(Misclnduus\ forcilatus OstenSacken; slightly flattened on slid6.
EXPLANATION OF ABBREVIATIONS. D : basistyle;d :dististyle.

ll:adeagus;

The terminology of the parts of the male hypopygium areinterpreted according to Crampton (Trans. Amer. Ent. Soc.
Vol. 48, p. zo7-zzS, ry23).
6. Venation of. Tonydcrus(Radinoclcrus\gloriosusAlexander.
7. Venation of Tanydcrus(Tanyderus)lictusPhilippi (after Philippi).
8. Venation of.Tanydcrus(Naodcrus)t'atagonicas
Alexander.
Alexander.
australaazsrs
9. Venati on of.T anyderus(N othoderus)ro. Wing of. Tanyderus(Mischoderus)
forcilatrs Osten-Sacken.
r r. Venation of Protoplasa(Protollosa)ftchii Osten-Sacken.
EXPLANATION OF ABBREVTATIONS. - Sc: Subcosta:
Scr : being the antelior or outer branch; ft : Radius, frr, ftu,
fte, Rr and Ru being the five branches,the last four being branchesof the Radial Sector (Rs); M : Nledia, with its four branches,M y M r , M r a n d M n ; C u : C u b i t u s ;A : A n a I v e i n .
The venation is here interpreted according to the Tillyard
modification of the Comstock-Needham System.
rz. Wing of Protollasa (Protanydcrus)aanduzczrAlexander.
t 3. Wing of,Protoploso(Protatryilarus)aipio Osten-Sacken.
r4. Dististyles of male hypopygia of species of.Protot'lasa.
A. Protoplasa(Prctanyderus)bechcriRiedel.
B. Protoplasa(Protarydcrus) uonduzeciAlexander .
C. Protoplasa(Protofl,asa)
ftchii Osten-Sacken.
r5. Wing of.Piringucyomyinabarnardi Alexander.
16. Head of Piriupteyomyinabanrurdi; ventral aspect.
I7. Head of.Piringuayomyinabanrcrdi; lateral aspect.
r8. Male hypopygium of.Plringacyomyinabarnardi; lateral aspect.
19. Head of Tanyderus(Ncodcrus)latagonicus Alexander.
EXPLANATION
OF ABBREVIATIONS.
lary palpus; lp : labial palpus.

Amherst, Mass. U. S. A. June r, 19z6.

-

mr!:

maxil-

GENERA INSECTORUMDIPTERAAru d?ongnht

ag@dw-Aaryeddrc"Yartory)pr,f

fag;dswgtLTebew

pdrqtou;MAla,vaattunn,va&bra-Tovdau

Iogftlaaa

zeland:afo.vr@EAS.tlW-4le.vaatie

2W%pHrEZId^mdret,$

P?pbpaqse

hronyT.'aa ?anAweq:

t\rE ti a s.,wadbtu

Alzz,wirg.

-L{:{i,bi
_6
"d
't,L
!.{
-dsfiz&E or ryryopyg
ie ", #.*

?r,o4a4@

vFu,Ag.,wtg.
-PAq:ngn4tortJ;lbu

Pam:nouqrongrbrL

bumatdt;

Ate

b@d,

AZce, t ed.,,fagdawarav;hbrarrAla,yaatbtu

whgt.
tu?Va%@6horptuaftttat
Iagrrtaqa&goriaw.lle.

O.S,wtatbtu

O,AsuWlm

rud.

ntn:ngqfoDgrb@

b@ae:

Alea_ZwdlatuqL.
a&Pe,

FAI.T. TANIYDERID.E.