vfl

THE DIPTEROUSFAMILY TANYDIIRIDIIIN JAPAN (INSECTA)C H A R L E SP . A L E X A N D E R
Pro/bssorofErltomolog!,]|.,IassaclmsettsStateCollege'
Antherst, Mass', U'S'A'
TIVO FIGURES(ReceivedNov' rE' I93r)

Kyirshir' Japan' andsobosan'oncollcctecl
Tipulide
Among somek i n d l y s e n t t o r n e f o r d e t e r m i n a t i o n b y m y f i - ai e nbeautiful
d ' . l t : l n tnew
: : t ^ lsPecles::::ofspecintens
tltreeEsaki, there were ipclude6
no mentber of thisSirrceTanyderida'of two-wingecl fly of the. family
i' the Japa'ese Empire'
take'
been
ever
had
archaic farniry of Diptera
b'e'hti (Riedel)'i+o"*.aous
species,othersingleaand, in fact, o'ly
a someadvisable to Prepare
seems
it
continent,on the entire *lurasianof Japaneseeutobenefit
theforrvhat detailecjaccount of the family
BibliograPhy that
the
to
refer
mologists. All clates in parenthesis
conciudesthe Paper'.

I.

GENERAL ACCOUNT OF

THE FAMII,Y

I.ANYDERID.IE

Diptera''fanyclericlaincludes the tnost primitive of all livingThe
in the family\l'ere
grouP
this
of
mernbers
theIJi:"dFor manly years
Philippi' r865)' still
rggo,'r886;
r869,
Tipulidne(osten Saclcen,r859,
19I3 ; F)nder(Alexander'
Dipteralower
tf,.
later, various stuclentsof
fliesrg"r) placed these
Rieclei,
to
Ig
;
r9",,
leirr, rgr2; Hancilirsch,
a subfanilY' the Tanyderinaasgenerallyin the fanrily PtycSopterida,
to full family
elevated the .. *loun
19)(i9Alexand.r
or Tanyderinr.
in conrnron rvith
littleha'erank. T5e members of the' Tanycleri6afamily for theirclistinctarequirethe Ptychopteridr and certainly
adult flies' the menlthe
ofaPPearance
reception. Fronr tlte general
273

274

CHARLES P. ALEXANDER

superfamilybers of this grouP have been held to belong to thethat theTipuloidea but recent studies by Crampton (tgz) have shorvu
and areadult flies afe very sirnilar in structure to the Psychodida
discovery
better placed in the superfamily Psychodoidea. The recent
a, l93o b)of the immature stages (Alexarrcler,I93o a; Crampton, I93o
has shown that ti:e group is highly isorated from either the Psychodidaon the one hand or the Ptychopterida on the other' The.Bruchomyiina'
forrnerly placed rvith the Tanyderida, are now placed as a subfamilyof Psychodidae(Alexander, rg2o c, tg27 b, tgz8; Crampton' t92si
Torrnoir, tgzz).
plrylogcny.. 'fhe first insect that can definitely be referred to the(Alexander,
Tanyderida is the Baltic amber ll(acrochile specyutn Loew
Tertiary
rg27 b, I93I ; Crarnpton, ry26; Loerv, r85r)' The lorverin
Etoptychoptra (t{andlirch, lgto) shows certaitt points of resemblance
ha's twothe nature of the medial and radial. fields of the rving but
no very
anal veins. Likervise the Mesozoic Eoptychopterida shorv
the familyclose relationship to the Tanydericla. The discovery ofbackPermotipulida (Tillyarcl, rg2g) has carriecl the order Diptera
shorvsto the Palaozoic. The insect in question, Permotipula Tillyard,
a highly modified radial field of the wing'grouP
IVforllzologl.' The general rnorphology of members of this(ry26i) and in aof flies has been discussed in cletail by Cranrpton
nrauuscript that is ready for the press by a sttrclelt of Dr' Crampton'
figuredMiss Inez Williams. The rving-venationhas been describerlandb,inany times (Alexander, I9I3, r9r9, rg2o b, tg2r' r g 2 7 a , r g 2 7
t924i
t g z } b , r g 2 g a , t g z g b , I 9 3 o a , l 9 3 o b , I 9 3 I ; Conrstock,Loew, l85l ;Crampton, tgz6; Edrvards,tg4 a i Handlirsch, ryq;
MacGillivray,rg23; cle lHeijere,I9I5 a' I9I5 b; OstenSacken, t869;

Philippi, 1865; Riedel, rg2o).
of
Imrnature Stages.' T[e very curious early stages of a member
and havethis family, .kotoflasa lftcltii, fiave recently been discovered(t93o a'been discusseclin detail by Alexander (193o a) and Crarnpton
solllel93o b). The larva eartier refierred to t[is sallle genus rvithqu"rti* (Alexander, rg2o a) has thus proverl to belong to sotne othergroup of flies, still unknoll'n.
memThe early stages of Protoplasa, and presumably of all otherTANYDERIDE

IN JAPAI.I

275

bers of the family, occrlr in wet sandy soil at margins of major streams,
ipdicating an aquatic or nearly aquatic habit. The rvhitish larva has
the borJy eucephalousand terete, rvith lateral spiracles on the prothorax
and eighth abclominal segment. No creeping rvelts on body. Caudal
encl ol,bo<ly with six long filarnents, of rvhich one pair are borne near
the caucralends of two elongate anal pseudopods,the latter terminating
in a circjet of short outer crotchets and a central group of long slendercrotchets" trotr simple anal gills. Length, total, 17 - t8 nrnr.; filaments
alone, 4.S- 5 mm. ; greatest diameter of body, I.5 nlmThe pupa has an appearance very much. resenrbling a Hexatontine
Tipulid. Head surmountedby a high bispinous crest. Pronctal breathing-horns small, smooth, equal in size. I-eg-sheathslying side-by-side,
the fore pair shortest, hind pair longest. Venation showing clearlyon wing-pads. Abdominal segments rvith a tuberculate armature,
chiefly near posteriormargins of segments. Length, 8.5 - 9 mtn-; rvidth,
clextro-sinistral,I.5 - t.6 mrn.; depth, dorso-venttal, t.6- t.8 mrn.
Distribation.' Eleven genera rvith twenty-four species are notv
known. These genera have the following general distribution, thefigure in parenthesisfollowing the generic nanle indicating the present
nurnber of species.
AraucoclentsAlexander (l).-Neotropical (Chile) (Alexander, tgzg b).
EutanyderusAlexander ( I ).-Australasian (Australia) (Alexander, I 928 b).
ll[acroclzileLoew (t).-Lower Oligocene (Baltic Amber) (Loerv, l85l).
Il'IischoderasHandlirsch ( S).-Au stralasian (New Zealand) (Handlirsch,r909).
NeoderusAlexander (r).-Neotropical (Patagonia) (Alexander, 19I3).
tYolhoderusAlexander (r).-Australasian (Australia) (Alexander, rgzz).
PiringueyonryinaAlexander (r).-Ethiopian (Cape Colony) (Alexander,ryzt).
ITotanydcnts Handlirsch (+).-Holarctic (Handlirsch, r9o9).
ProtoplasaOsten Saclcen(r).-Eastern Nearctic (Osten Sacken, r8Sg).
Iladindcrus Handlirsch (Z).-Australasian (Papua, Australia) (HandIirsch, t9o9).
TanydentsPhilippi (r).-Neotropical (Chile) (Philippi, r865)Iiron_rthe above digest, it rvill be seenthat the'Australasian Regionis richest in nrembers of this palaogenic group, with four genera and276

CHARLESP. .TLEXANDER

fourteen species (Alexander, l9z8 b, I93o a ; Iidrvards, rg23 a; deMeijere, rgrs a, r9r5 b;. Riedel, rgzt). The Neotropical Ilegion(Alexander, rgzg b) has three genera and three species, all in the
Chilean subregion. The Ethiopian Region has only the very distinct
Piringacyomyina bcwnardiAlexander. (l9zl).
In the eastern Nearctic
is found only the much discussedProtoplasafitchii Osten Sacken(Osten
Sacken,1859,r8@; Alexander, r9r9, rg2o a, t9z7 b, I93o a; Crampton, rgzg, rg3o a, t93o b). The sole remaining recent genus is
Protanydenn, u'hose four known speciesare as follorvs:
Piotanydenrs becheri (Riedel).-Turkestan (Riedel, rgzo).
-Japan (Kyushe).
P. esahii sp. n.
-UnitedP. uanduzeei(Alexander)
States (California) (Alexander,r 9r 8 ) .
-United States (British Columbia, south
P. dfio (Osten Sacken).
to Cilifornia, east to Colorado (Osten
Sacken; r82il.
From tlre above data it will be seen that the, occurrenceof amenrber of this fanrily in Japan was entirely to be expected. The
various genera occurring in the Northern Hemisplrcre (llfacruhile, Pro
tanyderus,Protoflasa) are all more specializecltltan are the Antipodalgroups in the lobing or bifurcation of the dististyle of the nrale hypopygium. It nright be expected that the group has attainedits present
distribution in the southernhemisphereby migrations across the former
Antarctic continent, thencenorthward along the Andean chaitr of SouthAmerica to western North America, sending one branclt into theeastern Nearctic (llotoy'Iasa) and another (Protanydenu) northward andwestrvard acros.sthe Bering land-connectionto Eurasia.
II.

DESCRIPIION

O}- .PRO'fANYDERUS $AKII

SP; N.

Protanydens esabii sp. ll.
General coloration of thorax light brown, the prescutum with a darker stripe on either side of nredian line ; legs yellow, the tips offentora narrowly but conspicuouslydark brown; wings whitish subhyaline, with a heavy brown pattern, arranged chiefly as three slightly
obligue crossbands,interconnectedat ceftain points.

TAI\TYDERID.E IN JAPT\N277

fiIale.-Length, about 7 mm.; wing' 7 mm'
Fenalc.-Length, about 7 - 7.2 mtn.; wing, 7'5 8 mm'
Itostrum chestnut brown, tlte elongate nrouthpartsblack' AntennapalelGsegmented, the scape and pedicel dark brorvn, the flagellum
onlybrown; flagellar segments oval to long-oval, rvith verticils thatslightly exceed the- segments in length ; terminal segment reduced'
gray, theapfro*i-ately one-third the length of the penultirnate. Headposterior vertex indistilctly marked with brownish, appearingas longiwith
tudinal areas on either side of the median line. Anterior vertex
a small tubercle immediately caudaclof each antennal base.
MesoCervical scleritesdark brown, tlte pronotum paler brown'
on .either
notum pale brown, the prascutal regio' with a darker strilleyellow, thesicle of the mediarr line. pleura light brown. Halteresknobs dark brown. Legs with the coxa light brown; trochanterstestaceou.s;fernora light yellow, the tips narrorvly but conspicuously
the .
dark brown; tibia yeilow, the basesvely narrorvly dark broln,segments
tips narrowly and weakly infuscated; tarsi yellow, the outera heavynrore infumed. Wings (textfig. l) whitish subfiyaline, with
but isbrorvn pattern that varies in its details in different individuals
at level of origin
arrange{ chiefly as three slightly oblique crossbancJs
tst IltI., the firstof Rs, along corcl, and at level of outer encl of cell
two connectedalong vein Al, the secondand third narrorvly connectcd
crucifornt
in cell 5b,; ,ving-nfex further variegateclby a more or lessvein R' to ]'1'
area, thc vertical arnl. crossing the wing from end ofbasally
the crossbarextending along cell R, to rving-nrargin,connecting

Fig. r-Wing of ?rotanydcnts esakii sP' t''-I'rotanydcnts csarii sp. n.
ri[. z_Dististyle of malc hypopygium of278

CHARIJS

P.. AIJSXANDER

with the outer band; additionaldark areasin cell C, at rving-margin
at ends of veins R, and IltI", smalldots and spots in interspaces,on Rs
M, and along R5; anterior prearcular region darkened; the trvo outercrossbanclsare paler in centersthan on the margins; veins brown,
light yellorv where traversing the ground-color. Anal angle of wing
very conspicuous,forrning an approximate right angle (slightly more
prominent tharr shorvn in figure). Venation : lt*, longer than thesinuous&; a spur from vein .R,jutting basadinto cell R; m angulate(as shown) to straight.
Abdomen brown, the sternites sonrervhatdarker brown, the caudal
margins of segmentsnarrowly pale. Male hypopygiunr rvith the dististyle (textfig. z) profoundly bifid, the nrore basal ann stouter, theapex narrorvly blackenedand acute; outer arm slender, blackenedon
distal third and here set with short stout setre.
I{ab. Japan (Ky&shir).
Holotype, 6, sobosan,Bungo,about4oo metersin altitude, July 16,.
rg3r (Esaki and Fujino).
Alloto/otype,g, July r8, r93r.
Paratopotype,g, July 19, r93r.
The type male and fenrale are preservedin the lintomological
Laboratory of the Kyirshfi Imperial University, Fukuoka; the paratype
fernalein the writer's collection.
I take very great pleasurein dedicating this striking crane-fly to
my friend and colleague,ProfessorTeiso Esaki, to whorn I' am very
greatly indebted for m.arr} favors in the past.
Of the four knorvn speciesof .l-lotanyclcrus,two P. becheriand P.
uanduseei,are srrrall species, showing a tendency to rving reduction,
and with the rving-patternpale and diffuse. The rernaining -species,P.
esahii and P. aifiio,are larger, broad-wingedflies, with the rving-patternclearfy defined and very handsome. P. esabii is well-distinguishedfrorl
the Nearctic P. aipio by the coloration of the bocly and the very
different banded pattern of the wings. All four speciesof Protarydertts
are before me at time of writing this article.
Anrong the Tipulida, species of tl:e farniliar genus Efi,hragnaand, in the Japanesefauna, of the Pediciine genus Heterangeus, havea somewhatsimilar banded wing-pattern and may be temporarily con-TANYDERIDzII tN JAPAN279

fusecl in collections u'ith members of the Tanyderidie. Thus, the Amencan Protoplasa ftchii has almost invariably been found in museum
collections to be placed in series of crane-flies of the fanriliar species,
Epiphrag ma fasciPennis(S"y). A glance at the venation, as the Possession of five branches of radius and the po.ssessionof a single anal vein,suffices to tell the present family fronr all other known Diptera excepting the srnaller,plain-tvingedPsychodida.

III.

BIBLIOGRAPHY OF THE TANYDEIIIDJ|E.

Alexanrler,C. P. r9r3 A revision of tlre South Anrerican Dipterous insects of the family
Ptychopteridr. Proc. U. S. Nat. Mus., 442 33r'335, 3 6gs.
rgrg New speciesof crane-fliesfrom California (Dip.). Ent. News, 2gi 285-286.
'l'hecrane-flies of New York. Part I. Distribution and tasonomy of the adultr9r9
flies. Cornell Univ. Agr. Ilxpt. Sta., Mern. z5 : 8B3r r fig'
'fhe craneflies of New York. Part II. Iliology and Phylogeny. C.ornell Univ.
rg2ogAgr. Expt. Sta., Mem. 58:. l6y7lz, pl- 13,
rgzob A new genus end specics of net-winged ruidge (Blepharocerid*) and an undescribed species of Tanyderidc (Diptera). Arkiv fdt knlogi, 13, N.,. 7: 5-7, tig.
rg2oc A new subfamily of Tanyderid flies (Diptera). Ann. Ent. .Soc.America, t3:
4o2'4c17,Pl. 32.
rg2r A new genus and speciesof Tanyderid (Phittgaeyoni*t bornonil in the 9uth
African Nluseum(Diptera). Ann. South African Mus., 18; e3r-234, fig.
rg22 Undescribed crane-flies (I'anyderidoe and Tipulidc) itr the South Atrstraliani\Iuseum. ltec- South Australian Mus', z: zz6'zz7'
tg24 Tw,r undescribeclspecies of ?'anydetrs fmtn tlre Australasian llegion. Insec.
Inscit. It'Ienst.r12 2 r4r'L43.
'fanyrgzs An undescril>e.dspecies of Tanydcnts from Western Atstralia (Diptera,
deridr). Insec. fnscit. Menst., t3t 32-34.
Diptera,rgz1x,The interpretation of the radial ficld of tlre wing in the Nenratocerous
Prnc. Linn. .Soc.New South Walesr5z:
rvith specialreferencet6 the Tipulidr.
44-45, r fiB'
tgzTb Tanyderidc.

Genera Inscctortrnr,Fasc. r&1, 9 figs'

rgz8aTlreAustralgsianspeciesoftlregenusA,cuo|a(,tts(Psychodide,Diptera).Proc.
Linn. Src. New South lVales, S3t 2gr'294, z frgs'
19:85 Tlre I'anyderickeof Australia (Diptera). Proc. I.inn. $rc. Nes' South Wales, t3:367374, figs. 4.
r9r9a A conrparisrrnof the systenrs of nomenclaturethat have been epplied to lheradial field of the wing in the I)iptera. IV. Internat. Congressl)ntomol', z:
7 @ - 7 0 . 7 , 3P I s .

CHARLES P. ALEXANDEII

28O

rgzgb Diptera of Patagoaia anct South Chile, Part I. Crane-flies. British Museum(Nat. Ilist.), pp. 227-229.
r93oa Observations on the Dipterous family Tanyderide. Proc. Linn. Soc. New South
Wales, 55 | 22r-23o, z Pls., r fig.
r93ob l'anyderus pictus Philippi (Fanr.Tanyderidc, Ord. Diptera). RevistaChinela deHist. Nat., 34: rro-rr3, r fig.
Crane-flies.ofthe lSaltic Amber (Diptera). Bcrnstein-Forschungen,tleft z: r'r35'
trgs. r-3.
Comstock, J. Ff. rg24 An introduction to entornnlogy, P. 796' fu.
Crampton, G. C. rg21 A phylogeneticstudy iof the thoracic sclerites of the non-Tipuloid
NematocerousDiptera. Ann. !lnt. Soc. America, 182 49-7415 Pls'
r93r

1935 The external anatomy of the primitive Tanyderid Dipteran Mocrcchilc sy'cct'ran
Loew, preservedin Beltic Amber. Bull. Brrxrklyn Ent. Soc., zr z r-r4, z pls.
rg2g A swarm of males of the rarc aad prirnitive cranefly Ptttoplasa flct;i observed
near Chandler in the Gasp6 Pcninsula (Diptera). Canadian llnt., 61 2 70'72.
r93oa Sorne anatomical detaits of the prpa of archaic Tanyderid Dipteron Ptotollasd: 83-98' 3 Pls.
ftchii, O. S. Proc. Ent. Soc. Washington, 3z
r93ob A comparisonof the more important structural details of the larvs of the archaic
Turyderid l)ipteron Ptotophsa ftchii, with otlrer llolometabolq from the stand'print of phylogeny. Bull. Brooklyn Ent. Soc., z5 z z3g-258r4 Pls'
Bdwards, F. W. rgz3a, A preliminary revision of the crane-fliesof New Zealand (Aniso'
p.rdide, l'anyderide, Tipulicla). Trans. New Zealand Inst., 542 27o'272, pl, zl,
6gs. 4-9, pl. 32, Frg. rz3,
r9z3b New species of crane-flies collected by Mr. (i. V. Hudson in New Zeiland.
Ann. Mag. Nat. Ilist., (g) rr : 625-626Enderlein, G. rgr2 Studien iiher die Tipuliden, Linnniiden, Cylindmtomiden und Ptycho'
pteriden. 7t>ol. Jthrb., Syst., 3z: 84'8J.
Ilandlirsc5, A. tpg Zur Phylogenieund Fliigelrnorphologieder Ptychopteriden(Dipteren).
Ann. k.-k. Naturhist. Ilofmus. Wien, z3: 263'272,Pl'rr, figs' r'r3'
rgro Canadian Fossil Insects. Contributions to Canadian Palamntologr 2z r22-t23,
r fig.
W.
rgoo The Tipulidc, or crane-flies,of New Zea,laod. Trans. New Zealand
Ifutton, F.
Inst., 3z s 48-49,pl. 4, frgs. zt'zzLrcw, I.I. r85r Beschreibungeiniger neuen Tipularia terricola. I.innaea entomologica,5:4o2-4o3,figs. 24-?5.
MacGillivray, A. D. rg23 External Insect-Anatomy, PP. 322'34, 6gs' 45'47'
Meijerc, de, J. C. r9r5a Diptera gesammett durch die 3t" Sud-Neu-GuiueaI'lxPedition.
Nova Guinee 13 3 5r'52t 69. l.
r9r5b Dipteni aus.Nord-Neu-Guinea.Tijd. vmr Entonrol., 58: ro4'ro6, fig. I.
Osten Sacken,C. R. 1859 New genera and species of North American Tipulidr, withslort palpi, with an attempt at a new ctassification of tlre tribe. Proc. Acad.
wat. Sdi. Plrilartelphia,1859: 197'256.
1869 il[rrnogrcphs of the Diptera of North Anrerica. Part IV. Smitlrsonian Misc.
Coll., zr9: 316-319,f:ry,7.

TANYDTIRIDIE IN JAPAN

z8t

r87T Westem Diptera. Bull. U. S. G€ol.'Survey.3: zo8.
l88o Die Tanyderina, einc mcrkwilrdige Gruppe der Tipuliden. Verh. zrnl.-bot. (les.
Wien, zg2 5r7-52zr6gs. t-2.
'
1886 Srudieson Tipulidc. Part z. Rcview of the published genenrof the Tiptlidc
brcvipdpi. Berliner Ent. Zeitschr., 3r: zz.8-41o.
.
Philippi, R. A. 1865 Aufzithlungdcr chilenischenDipteren. Verh. zml.$ot. Gcs. Wien'
15: 78o-782,pl. z% fig. 57.
Riedel, M. P. rgao Tanydents hccfreri.n.sp. aus Turkestan (Dipt.). Dic erstc rezcntet
pellarctishce Art der Farnitie Tanyderid.re. Zrnl. Jahrb, Syst., 43: 365-37o,+ figs.
rg2r NenratoceraPolyneura aus Nan-Guinea.usw. des Ungarisclrcn National.llrluscunrs
in Budrpdot. Anu. Mtrs. Nat. I{ungarici, 18: 143-144.
Tillyard, Il. J. 19z6 The insectsof Australia and Ncw 7*altnd, PP. 346-347,68. W gr'
rgzg Permian f)iptera fronr lYarner's Bay, N. S. lV. Nature, May 18, 1929,PP. I'4(scparatepagination).
(Dipt. Psyclndidc) et descriplion rl'une
Tonnoir, A. rg22 Notcs sur le genre Nemo1raQas
espdccnouvellc. Ann. Soc.-Ent. Iiclgique, 6z: 115'136,pl.