A new species of Thraulus from Nanjing (eastern China) with single first gill (Insecta: Ephemeroptera: Leptophlebiidae)

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ZOOLOGICAL SCIENCE 231 641-645 {2006) @ 2006 Zoological Societ yof Japan
A New Sp Thraulusfrom Nanjing(EasteCrhnina)
of

SingleFirstGill(lnsecEtpah:emeroptera:
withecies
Leptophlebiidae)

Peng Li ,DongLiu andChang-Fa Zhou'
The Key Labotatory of Jiangsu Bioresourc eand BiotechnoiogJt; of Lii eSciences,
IVanjin Ngormal Universit yN,arv'in 2g1O09Z P.CRo. iCiheignea
thraulus temoreitu ssp. nov., a new species tound in eastern China ,is describe dtrom imagos, sub-
imagos, nymphs, and eggs. The nymphs are unique in the genus as well as in the Thraulus group
because of their single lanceolate gill s1. The imagos can be differentia tterdom those ot related
species by the tollowing combination of characters: slightly robust hindwing with a bluntt ryounded
costal projectio tnhat is $ttuated over halt the distanc efrom wing ba$e to apex, darkl ypigmented
basal portio nof hindwing ,and subapical dark band on hind temur, Subimagos resemble the ima-
gos in color patter nE.ggs have eight pair sof long threads on the polar portio nand slight ridges
on the surface. The reduced gill s1 in combination with entirely fringed gill s2-7 represent a new
evolutionary lineag eand ecological type in the thraulus group, but provide limite dinformation
regarding phylogeneti crelationships,
Key words:Ephemeroptera,Leptophlebiidae,Thraulus, new species, China
INTRODUCTION STUDY AREA AND METHODS
The genus thrauiu ssensu lat owas established by Nymphs and subimagos were collected in early June in small
Eaton {1881 a)nd redefined by Peters and Edmunds ( 1970). pools of a stream, The width of the stream varies from 1-2 m, but on
This genu$ and ciosely related species were reviewed by average, i twas only about 1 m wide and tes sthan 30 cm deep at the
Gweaiidrgtdhahititnni t osaLnpne,edc ipes tsePvoeep rthawelelrr ee s(obt i1hire9e rdtta 9aoi nse3 iepwdne)hcoc,i le isupn d r seTooe7fpv ieotrnshae iu gsdl teug hsnesee ennTrushaus r.Ahta austtvthreaui tcsg ttbroieo ,muelepnn, ttadlhlieeemmpc oeot cssutee rif dm restcaetoesnl iudLitlr enel cwHtd,taheiseoan pv,alo ybe Toossuhlist eLs tt , ,ah1pnmad,aonn5o yt-l1h 2s ef.m lsrl0musoeb,m as i tvrnwa eahnidtdsc eih,aa n mthd ehatt edeh s reow a.am t neTreeorhwt et s e minsvape n lelo[cotdic oheirsste ty opa n toeow fstao hsfl els w toweciarow semlo-ef
reported from the eastern hemisphere (Gose ,1980; Jacob, oi collect[on, The $tream is i na well-protected forest rpyark ,with
1988i Soman, 1991; Kang and Yang, 1994). bamboo and large trees in the riparian zone on both side$i therefore,
In the original definit ioof nthe Thiaulus group ,the imag- the canopy is virtually closed, and Mtle sunshine can reach the
ina lforewing venation and nymphal fimbria tgeMs are dis- stream direct lWya,ter pH was about 5,56.0; temperatur ewas
tinc tdiagnosti ccharacter$ of these seven genera in the fam- around 16.30C; and air temperature was about 20-240C at the time
idglie yvn,eelrrn sa t heheavivo slegu tnrioootnu apyr ,yebt obttereh enin mddaes gs,acolrst ihobauengdd h ( nPytmhephets ne yamrn pdshssh oEwd m ofus nofmdeosu,r oCPtaa ceronaljihesac tnsiupoenrc,ui es Msy,aoy utiH lZahiboe urscoo lpalhnedclte edbBir ioardaoe smsc iithj h,i2en 0esn0as3mies, aY nsodtur e saomamned i nGbcauleitu, idd1 e 9sdop9ne5e-,
cies.
1970; Peters and Tsui, 1972; Grant and Peters 1,993} ,How- The nymphs were collected by hand screen, and subimagos
ever, all known nymphs of the group uniformly have seven were coHected by mosqujto net near the same stream. Adults and
pair sof gill sea,ch consisting of dorsa land ventral lobes, some subimagos were obtained by rearing the nymphal stage in the
ln 2004, we collected several nymphs that have gil l1s laborator Tyh,e water used in rearing was obtained from the same
single and gil l2s-7 marginally fringe din a park stream of stream. The nymphs usuaHy hid under leaves and siones when
Nanjing city, Jiangsu Province, Easter nChina ,This kind of they were put int othe collecting pan or rearing barre Tlh.e gil lwsere
leptophleb iniymdph obviously represents an undescribed observed to move very rapid[y,
taxon of the T7itautu sgroup .In 2005, we obtained additional TAXONOMICDESCRIPTION
nymphs, more than thirt ysubimagos, and adults, and
deposite tdhem in our collection. Basect upon the characters thraulu(s fFeimogr1sa-.t3u0s )sp. nov.
g7iT7vrean ubleuls.ow ,we estab[ish and describe a new species of Ma t e ri a l ex a m in ed . Holo typ e8 China ,Jiangs uProv-
,
ince ,Nanjin g,Zij inMountai n{118"51'23 13"2Ee,03'461"N,
' Correspondin gauthor.PFEha-oxm naei:l: +/+88z66-h-22o55-u-88c335h599a887n328g38;f;[email protected] aP1leOtn .gg8 96L I m, iD)m,o ansggt orLesa I,m4U a 8nnd8ea rC 3h aL\in n\gg --sgFu ub aZiHtmaeOgmUop.sl, eP, a31r0-a Vtllya-pr2ve a0s1e0: ,s5s aL,8mee 'gg.'
doi:10,21081zsj.23.641 data as for holotype ;other materials: 4 larva e1,O-Vl-2004,
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642 P. Li et al.
bothwith setae, a small tuftof apical setae on thirdseg-
/
%,X ment; anterior margin of maxilla with dense thick spines and
zak fbienl eoswe ttahee,s ea,n aa dIdaitrigoe natlh ircowke ropf e3c0t ipneac ttipenraotje esectatei o onnp reisnennetr
lyifii 11) corner; outer margin of maxilla and cardo with setae <Fi 5g)..
Hypopharyn xwith a projecti aonnd a row of setae on lateral
arms of superlingua (Fi g6), ,Labium igloss awith thick plate-
lik esetae on ventrai suriace, fewer setae on dorsa lsurface;
paraglosswaith denserbutfewersetae on dorsaisurtace;

fir ssetgment of labi aplalp with thjck setae on margins, sec-
ond segment with setae on outer margin, apical segment
with thick and fine setae on margjn and surface, those near
apex of apical segrnent gathere dlnto $mall tufts s;egment 1
lengt hca, 1 .2 times segment 2 lengt hs,egment 3 Iengt hO.9
times segment 2 lengt h(Fi g7)..
Pronotum brown, washed with paie irregula rmarkings
on dorsai surtace. Mesonotum brown, median area pale.
Legs pale but for efemora gray and hind femora with con-
spicuous brown subapical band (Fi g10.) .Femora with thick
setae, those on outer margin longer than others; additionaj
setae on outer margin, Tibia ehave fin eand thick setae on
innermargin, and vety sparse anctthinsetae on outer mar-

gin ,but hind tibia ehave more and denser thick setae on
surface, Tars iof all leg swith several thic ksetae on inner
margin and sparse setae on outer margins (Fig 8s, .9, 1O).
Claws of all legs similar, with a row of denticles ,middle den-
ticle sslightly large trhan basa land apical ones {Fi g1,1).
Abdomen brown, with ind[stin cptaire dsubmedian stripe
along body axis, Terga 3-9 wjth posterolater aplrojectjons,
those of terga 8-9 large r(Fi g1),. Gill sgray ,on segment$
1 1-7; gill s1 single, slender, lanceola t(eFi g1.2) ;gil l2s-7
similar i nshape, with dorsa land ventral lamella ee;ach
lamella elongated-ovate, with entire margin$ fimbrlate ;tra-
cheae of gillcslear (Figs13.-14). Caudaltilamentwsith

spines and setae at articulations.
Fig .1. Thraulu siemoratu ssp, nov.: habitus ot male nymph, 6. 0 -M7a.l0e ,m m,ln teethramnionla; l fbiolday mle ennt12g,t Oh-71.30.-08 ,mOm ,mm, ce rcfio rteOw.Oi-ng
11.0 mm, forele g5.0-6.0 mm. Body reddish brown ,com-
other informati soanme as fo rtypes, Al ltypes deposite din pound eyes pale orange, upper porti oonval in dorsa lview;
the lnstitu toef Genetic Resources, College of Lif eSciences, maior axis nearly perpendicul atro longitudi nbaoldy axis;
Nanjin gNormal Universi tPy., R. China .Two nymphs col- distance between compound eyes les sthan half width of
Iected in 2004 have been deposited ]n Entomology, Florida eye. Latera locelli located closely to eyes but not contiguous
A & M Universi tUyS,A. (Fig 1s5., 16) ,Thorax reddish dark brown .Forewings b:ase
Etymology. The specific epithet is derive dfrom the with ciear, dark transverse band; vein Rs forked at O.2 dis-
Lati nadjective temotatu s{femu rin) r,eference to the color tance from base of vein to margin; vein MA forke dat O.5 dis-
patter nof the hindlegs, tance from base of vein to wing margin; vein MP forke dat
Mature nymph. In ethanol; body iengt h6.0-7. 0mm, O.3 distanc ferom base of vein to margin, for ksymmetrical,
antennae 3.0-4.0 rnm, terminai filamen t12.0-13.0 mm, 2 intercalari epsresen tbetween CuA and CuP {Fi g,17).
cerci ".O-12.0 mm, Head capsule with area between oce"i Hindwings much smaller than forewings ,with conspicuous
washed with dark brown ;area between median oceHus and color marking$ at ba$e of wings; costal projecti obnluntly
fron spale; median lin eof vertex pale; margins of clypeus rounded, apex of project isiotunated at ca, O.6 to O.7 dis-
parall e(lFi g1).. Mouthpart sl:abrum with posterio rmargin tance from base to apex of wings; apex of wings blun tw,ith
slightly convex, irregul asprarse setae on both dorsal and apicai and posteri oarreas expanded siightly (Fig 1s8.-19).
ventral surfaces, additionally median anct apical row of hairs Legs: for etarsus sljghtly longe rthan tibia ,both of them
dorsall yaiong the anterior margin; width slight[y greate trhan much longer than femur ;for efemur and apex portio nof fore
clypeus; anteromedian emargination broad (Fig s1., 2). tibia ereddish brown, other part sof forelegs pale ,mid- and
Mandible swith sparse later aseltae, prosthec aof Ief tman- hindleg psale except hind femora with larg ebrown $ubapical
dible with two thicke rspines than others (Fig 3s, .4) .Maxilla: band {Fig s2.0-22>. Claws dissimila ro:ne blunt ,pad-ike;
$egment 1 of maxillary palp ilengt h1,2-1,4 times segment one apically hookect ;hooked claw narrowed abruptly at
2 length ,outer margin with a few setae; segment 3 length apex (Fi g26.),
O.8-O,9 times segment 2 length ,outer and inne rmargins of Each tergum of abdomen has one pai rof straight dark
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New Species of 77iraut ufsrom China 643
3
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2
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14
12 13
Figs. 2-14, Maiure nymph of 77irauJ utsemoratus sp. nov,2) iabrum; 3) lef tmandible, 4) mandible; 5} maxilLa ;6) hypopharynx ,7)
lablu mdorsa l(le fatnd) ventral (righ 8t)) f;oreleg ;9) midleg, 1O) h:ndleg; 11) forecla we,nlarged,nght12) gi l1l; t3) gil 2l, 14} g,1 71.
20
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29 t t l t tt t tt l t t.t t2/ 3/ /'-.,O /27
Figs. 15-29. Imago of 77]raul uisemoratu ssp. nov,, Figs ,15-27, male' 15) head, dorsa lview, 16} eye, later avllew; 17) torewing ;1B) hlnd-
wing, 19} detai olf hindwing ,enlarged; 20) foreleg ,21) mLdleg; 22} h-ndleg; 23) genital ilaa,ter avliew, 24} genital evae,ntral view; 25) detai olf
penes, dorsa lview; 26) foreclaw, enlarged, 27) abdomen, dorsa lview). Figs ,28-29, female. 28) latera lwew of terminal segments ot abdomen,
29) sternum 9, ventral vtew,
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644 P. Liet aL
mm, midleg 2.8-4.0 mm, hindle g3,O-4,O mm. Antennal
lengt h1.2-1,5 mm and sockets outlined with ligh tbrown.
Eyes occupy most of head, upper porti oonrange gray ,basal
porti odanrk brown. Wings semi-hyaline, outer margins with
a row of lightsetae. Inother characters, male subimagos

resemble male imagos,

Female subimago, ln ethanol; body length 8.5-9, 5mm,
forewing 8.0-9. 0mm, hindwing O.8-1. 2mm, cerci 10.5-
11,5mm, and terminalfilament11.0-13.0mm. Foreleg4.5-

5.0 mm, midleg 3,5-4.5 mm, hindle g3,5-4.5 mm. Antennal
lengt hO.7-1. 0mm and $ockets outlined with ligh tbrown,
Eyes dark brown. In other characters, resembling female
imagos.
Egg, (Fi g30., dissecte fdrom femal ebody} ,Lengt hO,2
mm, breadt hO,1 mm, Egg ovoid, some drum-lik ee;xo-
chorion with tin yirregula rridges, poiar portions wjth eight
pairsof thread-likestructure on surface, lengthof thread

about a thir dof egg longitudi n[aelngt hn;o visible micropyle
on chorionic surface found.
Taxonomic comparisons, Nymphs of the new species
resemble those of congeners in having superlingua with lat-
eral projecti oonn it sarms and similar fimbria tgeil l2s-7,
However, the nymphs of T7iraul utserriora tspu.s nov, are
unique i nthe genus ,as well as in the 7Titaul ugsroup,
because oftheirsingle,slender, lanceolategill1s.Allprevi-

ously reported species in the group have gil l1s with double
lamella et,hough according to Peter sand Edmunds (1970),
the firs tgill sof T7]raut uarse highly variable. For example,
gil l1 may consist of either a single dorsal lanceolat eportion
with a ventrally fringedlamella(asinmost species inthe

genus ,such as Z torrentis t)w,o ovate lamella weith fringed
margins (a sin the firs tundescribed species of Peters and
Tsui (1972 o)r) t,wo lanceolat elameliae (su cahs in 71 bellus
and T,bishopD,Singlegill1s,as describedherein,are a

new variation. Furthermor et,he dark bandin gof the hind
femora may help to distingui tshhe new species.
The males are simi[a rto congeners with regard to their
separated compound eyes, bluntlyrounded costal projection

Fig .30,Egg of 7hrautus iernorat ussp.nov.SEMphotograph. of the hindwin ga,nd the patter onf the genital iHao.wever,
the shape of the hindwing of the new species is peculiar,
too, because of it scostal projectio nsituated over half the
stripes at middle dorsally ,those on terga 1-6 ciearer than distance from the wing base to the apex, the darkly pig-
others: tergum 1O with truncated apex (Fi g27.). Caudal fil- mented basa lportio na,nd the slightly expanded apical and
aments pale apicall ywith reddish basa larticulations. outer margins. Other species of 7T)taul uussually have the
Genitali a{Fig s2.3-25): Stylige rplat ewith slightly con- costai projecti olnocated at most half the distance from the
vex posteri omrargin, Forceps segment 1 narrowing gradu- base to the apex of the leading margin or have an acute
ally from base to apex; apical one thir dcurved mesally; seg- costal projecti oAni.thoug h71 bettu hsas a robust hindwing,
ment 1 length ca. 6.0 times segment 2 length ;segment 3 $everal species of 77itauiu shave dark basal portion sof the
lengt hca. O.8 times segment 2 length ;basal portio nof for- hindwin g(e, g7.1 ,lasciatusan Td. gopalanD,an d71 bishqpi
ceps pale ,most part sof segment1 dark brown ;apical seg- and T demoutin ihave a more apically situated costal pro-
ment of forceps small, Penes straight, narrow; inne rmargins jecti o7nh,raulu siemotatu ssp, nov, i sthe fir switth this com-
strajght, outer margins converging apically; short spjne-like binatio nof characters. The hindwin gof T iemoratu ssp.
setae aiong outer margin dorsally. nov. is similar to that of species of the genus Sulu Grant and
Female. in ethanol; body lengt h8.0-9,O mm, cerci Peter s,but the respective costal projectio anrse not alike:
10,O-11.0 mm, and termina lfilamenM2,O-13.0 mm, Body the former has bluntl yrounded projectio nwhse,reas the lat-
color patter rnesemble that of male, Posterio rmargin of ster- ter have acute or acutely rounded projectio nIsn .addition,
num 7 straight (Fi g2.8), sternum 9 extended posteriorly thei rmale genital diiaff ecornsiderably. The adults of Z iem-
{Fi g29,), oratus sp, nov, also have a dark subapical band on their
Male subimago. in ethanol; body length 6.7-8. 0mm, hind temora.
forewin g6.0-7.5 mm, hindwing O,6-1.0 mm, terminal fila- Koss and Edmunds (197 4r)eported that leptophiebiid
ment 11,5-13.5 mm, cerci 10.5-12, Omm. Forele g4,O-5.0 eggs have the greatest diversi tiy nthe order, with the [ack
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New Species ofThrauius fromChina 645
of pola rcaps and the presenc eof an entirely chorionic fun- criticism ,and improvement of the manuscript. We expre$s special
nelform micropyle being considered the only features appreciation to Mrs, Janic ePeter s(Flori Ad a& M Universit UyS,A)
sbaehngeadgersn eYd fad oonrbcgfyu i m(vae le1nsl tp9kee9cndio4 e wsw)nb.hyo , sEM pgapegzcrsizoe iosvnf.ii oduaWren iddc tnl heewiGanra istSpnheEocr Mia(esu 1Jpu9h soa9 r,te e0oga gvg)nresdray pK hhssaai mnvo-gef fTU(FohnoNr iiuhasvn weendorrra jkest niiicN ot nwoou UaynrgrsoSa,fAmg e)afCmleih nfnitUoannr nail cv(iien Gaogr rful$s iaultipsynht p toi CNtyir ocshc,tsoseirtd.rnu eadc3yt )0 ib,5 foy7tno s0otr ,2hhM0ei 0r as N.n adh Lteau ilntkodpon e a iM3J]n r0aNs, 3ccaKa0otne0bnu-i0urnY3sga u 7l( S)W cPAaeiuNngeGdgrsn d,cauee
ila rto those of 71 tatuu sKang and Yang with regard to their fund from Nanjin gNormal Universi t(yNo 2.005104XGQ2B70),
shape and thread length and location .However, eggs of the
former species have fewer distinc tridges on surface than REFERENCES
those of the latterspecies.DISCUSSION Eaton AE O881 }An announcement of new genera of the Ephemer-
idae ,Entomol Mon Mag 17/ 191-197
Gose K (198 0T)he mayflies of Japan. (7 )Key to the familie sg,en-
Although the single gil losf the new species provid ea era, and species. Aquabiology 8: 2" -215 (i Jnapanese)
good diagnost ichcaracter, they should be used with caution Grant PM, Peter sWL (1993 )Description ot fou rnew genera ot
tisih nnep gdhleyolugobiglele nlpgelitlailt c soerfe I.mcnaoynfLsleiteprstucotp iohorlinge.ib naiRliliydeak ed,{ePr1eit9ve7erd3s fs)uragnogdems taed Epdrmiu nmtdihstative TrreoTdpietrseacrraui:lp uts LLogenrpo tuoofpp hSmliayemfboliiteihsdr aaAfeutr:laoul maos nptdhlh ee Cbiehiainsuatees)r a,nT nrohaepnmhsi stApe hm{beerEireap Ehnanetdom-e-
(197 0p)r ovide dsom e interpreta t oifo tnhes e cha racters in Ja c obmolU So(c11199:88 71)731i-r1a6u8lutshrakersp. n, aus Bulgarien(lnsecta,
order to discus sthe phylogen yof the genera .According to E phe meropt era /Leptop hlebiidae). Re iche nbachi a26 : 1-3
thei rhypothesis ,the gi lwlith single lamella may be inter- Kang SC, Yang CT (199 4L)eptophlebiid aoef Taiwan (Epheme-
preted as plesiomorph iacn,d the gi lwlith doubl elameHae roptera). J Taiwan Mus 47/ 57-82
may be interpre teasd apomorphic. On the other hand, Koss RW, Edmunds GF Jr (197 4E)phemeroptera eggs and their
degenerati oonf gil l1s is a common phenomenon in may- contribution to phylogenet isctudies ot the order. Zool J Linn
flie sV,estigi aorl reduced gill s1 have been found in several Soc 55: 267-349
families,includingEphemeridae, Potamanthidae,Ephemer- Mazzin iM, Ga[no E (1990 O)ogenesis and involvemen tof chorionic
ellidae,C aenidae ,and Neoephe meridae, structures in Ephemeroptera taxonomy. In "Mayflies and Stone-
gtru or ue Ppoefs t igeni rtl shi aesn1 dg. e PnEeudt sme ur7 naTd nisdia Tu(stuu1 iss 9en(7s u10r9e)c l7oga n2tilzo)a ebdtaes rese dpt ahornrae tete dhsep e scttirheusec-m P e t efAcralcts iaiWeodLsn e:,L m ioiEtf ecd mtHPhujuesnb tdelosaris itsGeehrs Fnear n J d$Drho, (reBdm1rrieo9$cl7pho th0gR,e y)e rp"vEe pi dL9 se5ib-poy1 t n0oo4Ifp Ct hhClea egmbepinbieed rla(ile E c,cpKlalhsusewiefmire--
into four ecological types based upon abdominal gi lsltruc- roptera). Pacifi lcnsect s12: 157-240
tur e{simH daoruble gil l1s-7, gil l1 sdissimil tao rothers, ven- Peter sWL, Tsui PTP (1972 N)ew species of 77iraul ufsrom As[a
tra llamella reduced, and dorsa] lamella reduced). These (Leptophlebi Eipdhaeemeroptera )O.rienta llnsect s6/ 1-17
authors pointe dout that gi lslhape is correlated with water Riek EF (197 3C)lassificat ioto nthe Ephemeroptera ,ln "Proceed-
movement and aquatic oxygen concentration, and sug- ings of the Firs tlnternationa lConferenc eon Ephemeroptera"
gOtebysvptieeo,a duntsdhla ytt ht,tehhee r enldewuacse dttshpregeciei eltslyt pseraesnp rddeseewnretslilv-ed edfvaer lonoemp wetdh e ecfgoiliorgls ilctoa2nsle-,7 S o m aEtnodp hbAlyKe bW{iLi1d P9aee9 t/ Ae1Atr )asnl,e woJpGh slPpeeebcitieiesnrase o, >Ltfe r7iohdmr eNani u,Bllrgui is(l r lpEiS,psop ,1uh6t0eh -m1l7ne8drioa J.p BtLoeempr--a:
sugge st a new lineag eada pt ed toquietwater, silty sub- bay Nat His tSoc 88: 99-1e2

strate, and probabl ylow aquatic oxygen concentration. {Recei vOectdober 30, 2005 1 Accepted March 21 , 2006)
ACKNOWLEDGMENTS
We thank two reviewers for their valuable advice, constructive
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