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ZOOLOGICAL SCIENCE 19:1377-1388 (2002) @ 2002ZoologicalSocietyofJapan
MorphologicalComp arison among Drosophilallniandtts
Two New SibtingSpecies(DipterDaro:sophilidae)
M s t , T h a n g i m a Z a n n a t a n d M a s a n o r i J . T o d ak
lnstitu otieLovv femperatur eScience ,HokkaidO Ciniversi44
}<i t1a9 Nishi 8, Kita-k uS,apporo 060-0819 ,.Lapan
ABSTRACT-Morpholo'gliaciawlan differen carees investigat eudsing several culture strains ef three sibling spe-
cies collected frem and Guangdong in China and Pyinoolwi nand 't2ingon in Myanmar. Careful
examination of male terminalia reveals distinguishabl edifferenc eisn the paramere and the aedeagal basal
process among the three species, tn addition, a number of quantitati cvhearacters are compared, Kruskal-
Walli stests with Bonferron ciorrection, which are carried out separately for each sex, detec tsignificant dif-
ference sin 15 characters, of which two are male-specific, among the three species, Canonical discriminant
analysis using these characters reveals that the three species can be distinguish efdrom each other with
high confidence for both sexes, The results clearly show the presence of three good species, DrosophMa
(Sophopho rliah )iBock & Wheeler, 1972 and it stwo new siblings, The new species are describ eads
Drosophil a(Sqphopho rohan)ishii sp. nov. from Pyinoolwi nand Drosophil a(Sophopho rogau)mai sp, nov.
from Ytangon.Th emorphological differenti aamtonigon the three $ibling species does not coincide with the
degree of reproductive isolati o(nbase don a previou sstudy), The premating isolati epnatter n$uggests two
possibilit tiheast premati nigsolatio nhas been evolved or reinforced in sympatric population bsetween D.
ohnishii and a lin aind between a ohnishii and D, ogumai or that i thas evolved in a very restricted local
populatio onf a ohnishii, possjbl yby a few mutations.
Key words: discriminan atnalysis, kikkawaispecies-complex ,montium species-subgroup, Oriental
Regionq,uantitaticvhearacter
D. jambuti nthae ,D. kikkauai ,the D. nikananu and the D.
INTRODUCTION senata complex, The members of each species-complex
The Drosqphil a(Sophopho rmaon)tium species-sub- are hardl ydistinguisha biln emorphology and show various
group is the larges tin the D. metanogaster species-group, incipie nsttages in differentia tiofo ncharacters such as
comprising a tota lof 85 species from Asia and Afric a(Leme- genes, protein s,physiolog y,behavio rand morphology.
unier et al. ,1986; Toda, unpublished data> .This subgroup Those are very good materials for studying the speciation
include sa variety of species at differen tstages of speciation mechanisms,
process and provide svery u$eful materials for studies of A few geneti cinvestigat orhsave suggested the pres-
evolutionary genetic sT,he relationships among members of ence of two new species very closely related to Drosqphila
this subgroup have been investigat ebdy various methods; (Sophopho rfian )Biock & Wheeler, 1972. Drosophil afiniwas
hybridizat io{ncro sesxperiment) test s(Dav iedt aL, 1978; describe das a new species ot the montium subgroup on the
Kim et al., 1989), two dimensiona lelectrophoresis and con- basis of a strain (Texa 3s146,1 )established from a single
ventional starch gel electrophoresis (Ohni$ eht iaL, 1983a, female collected from Taiwan in 1968 (Boc kand Wheeler,
b; Ohnishi and Watanabe, 1984), chromosomal (metaphase 1972) ,When Tsacas and David (197 7)established the
karyotype a)nalysis (Baim aanid Chumchong, i980; Baimai kikkatataicompl etxh,ey include dD, iinii tnhi sspecies-com-
et al. ,1986) ,and eco-physiological comparison (Kimura, plex.
1987) .Through these studies several groups of very closely Ohnish iand Watanabe (198 4wa)s the firs tto recognize
related species have been recognized as specjes-com- the presence of a sibling species of D. Iin ibased on the
plexes :the D. auraria, the D. bakoue, the D, bocquea the results of two electrophoretic analyses, They analyzed pro-
' Corresponding author:TeFlA.X . ++8811--1111--770066--67B18472;. talelion zymdeif fvearrieant cioebny bty wos-tadricmhe nsgieol naelle cterloepchtorroepshiosr,es isusi ngan da
E-mail:[email protected] strain {MMY326 )originating from a single female col[ected
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Tablel. Isofemalestrainsinvestigated
SpeciesStrajns Source
D, fini Bowling Green 14028-0581.0 (Texa 3s146.1) Yun-Shui, Talwan, 1968
DHS401DHS501NKS9212 DinghushanG,uangdong,southern China,1992
dittoNankunshan,
Guangdong,southern China,1992
NKS9231 dittoPynoolwin,
D. ohnishii MMY307MMY326RGN3RGN206 central Myanmar, 1981
dittoYangon,
D, ogumai southern Myanmar, 19Sl
ditto
at Pyjnoolwj n,central Myanmar in 1981, along with $uch 1000C for several minute$. Drawings were made on the basis of
strains of other 28 species of the montium subgroup. They microscope photograp htsaken by a computer-interfaced digital
tpelnatcaetd iMvMeY nl3ay2m6ed a si taa sm eDmbreorsop ohfi tl halei nkFilkik kaTewh.ae inco,m pKliexm e ta nadL c Tah mee raA,Hlol ktkyapi desopeUcnimievnesrs iwetryMeus eduemp,osiHtoekd kian iSdyostUenmiavteirc sEinttySo,ampoploorgoy,,
O989 >examined the crossability between D. tin ajnd D. IinF Japa n (SEHU) .
lik e(MMY326 )a,nd revealed that the premating isolation
between them was almost compiete. Furthermore ,Kim etaL Terminology and quantitativ ceharacters
d(prl1ia9ac l9eD d3 NDeA)x. a fmiiinhne di1-8l i(rskepMeseMtcriYiecs3ti2 on6o fc)e lntozshyemese tm otn otp iDau,mt tMie irs ,nuM bsoogrfre oum pir,teocceh notanln-yd, aaannn dddi o rWZiehn adc anilfgccoue llsaa lt.nAeodd w teo T dtfoMaodcl rAa ol fbp (oi3nt11e h 9sq e9ux( 2ae1tsn)9,ot8 rit1t1 haO)e{ t coidh r veacemrhtoaaircrptnaheciortltseori g$tooi ofcn w areslmmreae altse u,emr reeaammsnieudrnne todoslnoegy
Oguma et al, {199 5e)xamined the reproductive isoiation character for female {Tabl 2e).
and courtship behavio rbetween nine isofemal estrains from
Tstaoickwa n no(.B o3w1l4i6 .Gn1rg=eDe.n sItiocnkO ,nmoa.i n{1a4nd028 -C0h5i8n1a, O=(TDeHxaS s3ol1d5, D sia btl ianF gain rassltpy,es cetisoe st,in d wmeea niinngttuelnsi vchea lreayxcatmeirnsed f otr hdriesec rstirmaiinnsa t(i nBtghGe 1t4hr0e2e8-
DHS401, DHS501, NKS9231; all from Guangdong Prov- 0581.0, MMY326 and RGN206), one from each species, for the
ince )and Myanmar (MMY326=D ,finNik eM,MY307 from above 42 quantitat cihavreacters, As some metrjc characters would
Pyinoolwi nR;GN3, RGN206 from Yangon). They suggested have been affected by the body size, i.e .culture conditions, corre-
the existence of at leas tthree genetical ldyistin csitblin gspe- lation with the thorax lengt h(ThL w)as examined for all charaeters
cies in them: D, tin i(BG14028-058 1DH.S031,5, DHS401, earxacteeplyt ffoor r tehaec hbo dseyx l. eSnegxthu ,ta hlde iwfinfger leennc gweats ha nedx atmhine ewding b wyi dutnhp,a isreepd-
DHS501 and NKS9231) distribute dfrom Taiwan to southern Ftest for each character and interspec diiififceren wcaes examined
China, D. fihi-li k{eMMY326 and MMY307} in central Myan- by Kruskal-Wall itsest separately for each sex, except for the four
mar and the other species (RGN3 and RGN206) in southern characters representing the body size. Bonierro ncoirrection (Rjce,
gMuyiasnhmeadr f. roHmo weeacvher o, thtehro smeo rpshpoelcoigeisca llcya,n baned h tahred lt wyod insewtin- 1tt9ie8ps9lt )esc owwmjaptsahr i sBaoponpsnl.ifeedr r toFono r ircc ooprtirhnegce tic ohnw,aitrha c 1pt0err osmba alsebesjlle ic tateneryddr o rbtsy Oc aKfuresmueadslk ebasy l w-meruWela-llis
sibling species have not been formall ydescribed yet, measured in each of the remaining sjx st rains, Combining such data
ln this study we compare the morphology precise lfyor of the nine strains ,we performe tdhe canonical discrimina natnaly-
both qualitati avned quantitati cvhearacters among the three sis ,using the computer software STATISTICA (StatS o2f00t0,), for
sibling species, using nine isofemai estrains of which eight the three species, separately for each sex.
are the same as in Oguma et aL ( 1995), and descri btehem,
two of them as new species.
RESULTS
MATERIALS AND METHODS Description
The followi tnhgree $pecies are very similar in general
S ofp et chTie emnnei nnmesa il esexoa fmaienndmeda lIe Os tfraeimna$l (e Tspaebci l1mee)n. sThe w esrterai nsex ahmianvede be efno r meaaicnh- mion raplhlo ltohgey .thr Eexe stpeecrineas lq uaarel iftiart si rtcevhfeaerrarcetde rtso icno mtmhoen lryedes csreiepn-
taine don a standard Dtosophil aculture medium for long periods tio nof D, hai but not repeated in the descripti oonf two new
since the establishment each from a single wild-caught female. species. Quantitati vceharacters that vary among the three
Extemal morphology was observed under a stereoscopic species are analyzed in the subsequent section.
microscope and metric characters were measured with an ocular
micrometer, The detaile dstructure ot head, male forele gand male Drosophila(Sophophottai}niBock & Wheeler, 1972
aan dco mfpoeumndale t elrimgihn atmliicra owsecroepe, obs eravfetder d eitn aac hdirnog ptlhee t oofr ggalnysc efrr ooulnm dtehre (Fig sI A., 2A-C)
body and cleaning them by warming in 1O% KOH solutjon at about DrDsophila (Sqphopho rLian )iBock & Wheeler, 1972: 59.
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Two New Siblin gosf Drosophil ahai 1379
Table 2.Detinitions of quantitati vchearacters examined
Both sexes
BL straight distanc efrom anterior edge ot pedjce lto tip of abdomen
ThL distanc efrom anterior notal margin to apex oi scutellum
WL distanc efrom humeral cross vein to wing apex
ww maximum wing width
dorsal number of dorsal branches of arista
ventral number of ventral branches ot arista
spcs number of supra$ervical setae
pocls number of post ocular setae
ocps number of oocipttal setae
ms number of medial cibarial sensilla
ps number of posterio cribaria lsensilla
FW/HW fronta wlidth / head width
ch/o maximum width of gena f maximum diameter of eye
prob proclinat oerbital seta f posteri roerclinate orbital seta jn length
rcofo anterior recliflate orbita lseta 1 posteri roerclinate orbital seta in length
vb subvibrissal seta / vibri$sa in length
dcl anterjor dorsocentra sleta 1 posterio drorsocentra sleta in length
sct] basal scutellar seta 1 apical scutellar seta in length
sterno anterior katepistern aselta / posteri koartepistern aselta in length
sterno2 mid katepistern aselta / posterio kratepi$tern aselta in length
distanc ebetween proclinat aend posterio rreclinate orbita lsetae 1 distance between inne rvertica land posterio rreclinate
orbito
otoita lsetae
dcpsctlpWWl/eWnLgct4 hcd4ivs5txaancoM ebetween ipsilate rdaolrsocent rseatlae 1 cross distanc ebetween anterior dorsocent rseatlae
distanc ebetween ipsilatera lscuteilar setae 1 cross distanc ebetween apical scutellar setae
maximum wing width rdistan cferom humera lcross vein to wing apex
second costal $ection between subcestal break and R2+3 f third costal section between R2+3 and R4+s
thir dcostal section between R2+3 and R4.s I Mi between r-m and dm-cu
Mi between dm-cu and wing margin fMr ibetwee nr-m and dm-cu
CuAi between dm-cu and wing margin 1 dm-cu between Mi and CuAi
thir dcostal $ection between R2.3 and R4.s / distan cbeetween djsta e1nds of R4+s and Mi
CuAi between dm-cu and wing margin /Mi between r-m and dm-cu
C3FMale lengt hof heavy setation in thir dcostal section / (leng tohf heavy setation in third costal section + lengt hof ligh tsetation in
thir doostal section>
scl number ot teeth in sex-comb of 1st tarsomere
sc2 number of teeth in sex-comb of 2nd tarsomere
epands number of latero-dors saeltae on epandrium
vlbs number of ventral lobe $etae on epandrium
surpg number of pegs on surstylus
sursp number of spines on surstylus
cers number of setae on cercus
cersp number of spines on cercus
pg2 number of pegs en secondary clasper
s2Female number of setae on secondary ciasper
ovm$ number of marginal ovisensilla on oviscapt
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'
t'
t.r'
nl x
Z7/
c
flA
vx
'" g
N
.!ILRC( >>ll
N
ww
d cl
x 4e
N
'N'
A sur B c
' ''
Fig. 1. Periphall iorcgans. A, Drosophil a(Sophopho rlian i)Bock & Wheeler, 1972; B, Drvsophla (Sophopho rohani)shii sp, nov,; C, Droso-
phila (Sophopho rogaum)ai sp. nov. epand=epandrium, v lb=ventr allobe of epandrium, sur=surstylus (prima crlaysper), cerc=cercus, 2nd
cl=secondary clasper. (Scale-lin mem)=,O.1
aed
t'veieik
aed b pr
#
,; `' pm
･ie1'1'
r't"'-'t'
,i
1. l
XN !j ilZ:
X "lt SN ･･,... ,
li･'
NX vt ....t.
tt/ t/t
I /･.･,,,.,..,,･,,..
t/･/'i/
hypd L ttl,i,,,/l/i,S
1i"'
"i /'v･,,,
l･,.
,
aed a
/1iii'"./,1
A
't/
di
'e
` 6'
c E F H
Fig, 2. Phalli corgans (A,D, Gwh:ole in ventral view; B,E,H: aedeagal basal processes in dorsa lview; C,F,1 :paramere) ,A-C, DiDsophiLa
(Sophopho rtain} Biock & Wheeler, 1972; D-F, DrosophiLa (Sophqphor ao)hnishii sp. nov.; G-l, Drosophil a{Sophcrph oogtumaai) sp. nov.
hypd=hypandrium, pm=paramere, aed=aedeagus, aed a=aedeagal apodeme, aed b pr=aedeag ablasal proces s(,Scale-lin mem>=.O.t
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Two New Sibling sof Drosophza alin i 1381
Diagnosis ,Paramere apically round and with 3-5 gle pale brown; ocelli orange. Anterior recfiinate orbital seta
minute sensilla, submedially with strong, triangul airn,ward much cleser to proclin asteeta than to posteri orreclinate
expansion of which posterio mrargin only slightl yconcave seta, Antennal pedice lusually yellowi sbhrown. Face and
(Fi g2.A,C} .Aedeagal basal process with fine ,irregula rser- gena yellowis hbrown. Carina convex, somewhat broader in
rations on slightly wider, apical margin (Fi g2,B). \ , Palpus and clypeus brownish yellow .Pa]pus with t
Description c7i and \ . Head: Eyes brigh tred, Postocel - prominent terminal and another subprominent, lateromedian
lar setae convergent, Supracervic asletae tapered, thin ,api- setae. Cibaria lposteri osretae long ,tapered, thin ,curved
cally slightly curved and sharp, Frons includi nogcellar trian - and apicali ysharp; medial and anterior $etae shorter than
Table 3. Mean±SD and range of 42 quantitat cihvareacters in three species
D. Iini O. ehnishii D. ogumai
Character Male Female Mele Female Male Female
Mean±SD Min, Max. {M Mean±SD Min. Max, (M Mean±SD Min .Max.(M Mean±SD Mim .Max, (N ) Mean±SD MIn. Max. (M Mean±SDMin.Max. {M
BLThLWLww2d.ooresea.lov7eon.1tB.rb9al5 lo2s..pe1c35 s1(p.o51c50 lv)se2o..ce2p77±sOOm,.s71p32stFO2W..ot0sH3 W2e.h39i oOpDTo)br1e.e9r4b±Ov.b1d0c l1s.c7t4 12s.t0e9rn(or$i tOe)2r,n2o22±Oo,d1e0it1o.d7c 4p2s.c09t lp(V1V0W)i2W.Lece4roc,4ov75O,x18a,19c6M C23.F1s7 cOlsOc) 2e2p.a5ntdisOv.i1b1s2s.u3r9pO2g..sS7u8 1r1s(,p61c70eO}r.sD7c.7e9r4sPpg2suovm$
o.so o.Bg (lo )D.g2±o.o2O,88 O.9 5{10)O.80±O.02 OJ7 O.82(ID }O.g3±D.02O,90 O.95 {10) ±O,021.o5,27s o.s4 oo} e,gl±o,o2 {10)1.82
1,45 1,6 4UO] 1,65±O.051.sa 1.71 {le)1.57±O.e3 1.51 1,M (10 }t.76±O.071.58 t.8 1(10) ±O.04O.17.143 1.58 {10 ) 1.73±O,05 {tO)O.83
O.69 D.78 (10 }O.79±O.02O,76 OS2 (10)O.72±O,e2 O,69 e.75 <ID )O,79±O,03e,76 O.85 (10) ±O.03O.66 O.76 {10 )O.8ri±O.02 {10)
4.0±O.O 4 4 (10 } 4.2±D.4 4 5 (10) 4.e±o.o 4 4 {10 )4,O±D.O 4 4 CIO) 4.0±O.O 4 4 CIO )4.2±O.4 4 5 {10)
3.0re,O14.e 3 3 (ID }2,9±O,3 2 3CtO) 3.e±o.o 3 3 {10 ) 3.0±O.O 3 3 CIO) 2.8±O.415. g 2 3 CID )3.0±O.O 3 3 {10)
±1.713.3 11 20 {50 ) 16.B±1.5 13 21 (50)14.S±O.9 12 17 C20 )15.2±2.0 10 oo (20) ±1.e14, 414 20 (2e )IB.O±1.4 15 22 {20)
±1.2ri1. 711 15 {10 ) 15.0±1.4 13 le (10)14,1±10.7 13 15 (10 )14.B±t.2 12 16 (10) ±1.315. 911 17 (10 )14.4±1.B ti 17 {10)
±t,6 S 15 {50 )13.9±2.0 le 21 (50)12.2±1.7 9 16 (20 )14.7±2.4 10 20 (20) ±l.9 12 21 (20 )16,1±1.7 13 25 {20)
7.3±O.820 . 06E. OO,94 5(10 )B.O±O.6 7 9(10) 7,9tO,7 B 9 (10 ) 8,2±l),4 s g"e} 7.5±D.921 . 56 9 (10 ] 7S±O.9 6 9 (10}
12 26 (5e )20.9±1.7 14 26 (50)192±1.6 16 24 (20 )19.2±2,1 12 24 (20} ±2,eO. 4I6St D.2052 o.(e2e 0)22,9±ri.6 lgO .4256 O.C0270O>.8O2,O4.B45O.78esgO.73O.47O.24O.51O.40ri.OsO.441.591.4t2.512.312.64O,92e.52
±O.02O.0o9.41 o,4e (so )o,4s±o,olO.42 O.47 (5e)o.47±o.ol o.4s o.4g (20 )O.47±l).elO.45 O.49 {20} OA4 O.48 (2 0)O.47±O.Ol (20)O.09
±O,OO1.0O0.07 O.09 (10 )O.09±O.Ol o.es o.11 uo) o.eg±e.o lo,o se.lo(10 )O.08tiO,ooO,08 O,09 {10) ±o.Dl1o,.1o07 o.lo "e} e.osto.oo (10)1.15
±O.05O.4o3.gl 1.12 oo) o.e7m.os O.74 1.03 <10}1.et±O.07 O.88 1.14 oo) e.B6±o.o7 O.7B 1.oo {10) ±O.D8O1..4o9o 1.25 (10 )O,94±D.10 (10)O.65
±O,D6O.OT.634 O.53 (so )e.46rc.06e.37 D.57 (50}o,4s±o.e4 o.3 eo.sri(20 )O.48ee.03 O.42 O.55 {20} ±O.03e.Oe.442 O.53 (20 }O,55±O,06 (20)e.gB
±O.tOo,o.s6Bo o.g6 cte }o.7s±o.ogO.67 O.93 {10}O,78±O,e6 O.67 O.B6(10 )O.76±O,13O,63 e.95 (10> ±o."o,6O1.69 1.00 <10 )O.6Stl).06 (to}O.63
±o.e4Oo..847sr co..ooss Oo.o5Z) roe..6coatOo..o2s4O,49 O.67 {tO)O.57±O.06 e.44 O.67oo} D.se±o,o3O.53 O,63 (10> ±o,e4o.Oa,256 O,68 (10 )O.6osl).Ol (10}O.99
O,76 1.08 (10 )O.91±O.04O.S2 O.96 OO} o.s4±o.og o.7o 1,oo (tO }D.85±O.07O.77 1.0 1{iO) ±o.osO.O5.974 O.90 {le )O.83ve.e7 (10>O.67
O,44 D.59 (10 )O.57±O.04e.49 O.65 (10}O.55±O,04 O,50 O,63 (10 )O.58±O,04O.49 O.63 {10) ±O,05oo..2s63 o.7o clo )o.61ve.es CIO)O.3S
±O.04O.6O9.15 O,29 (10 )O.29±O.07O.IB O.4 1(10}O.27±O.03 O.2e O.32 (10 )O.30±O.08 e.24 o.so {lo) ±o.esoD..6124 tOo,.3e0 soC.I4O6 )O.31±O.05 <10)O,66
±O,"e.62O.56 O.95 C50 )O,59±O.06O.48 O,67 {50}O,7or,O. 0O6.64 O.79 {20 )O.64±O.04 O.ss O.69 C20) D.5 1O.e6 (20 )O,61±O.04 {20)O.51
±o.o41.0O0.35 O.50 (10 )O,41±O,02O.39 O,43 {10)O.47±O,03 O,40 O,50{lo )o.4Mo.e2 O.39 O.46 (10) ±o,es1.O1.332 O.52 (10 )O.46±O.04 {le)1.37
±O.05O.4D7.86 1,16 C50 )1.05±O.07O.94 1.IB C50)1.0eeO.14 D,63 1,43(2o )1,os±o.e7O.90 1,13 (20) ±O.07O.O4.799 1.20 (20 )1.21±D.08 C20)O.49
±O.021O.,s43t LO.49 (10 )O.4B±O,02O.45 O.51 (10)O.46±O.Ol O,44 O.48oo) o.4sdi.e3O,42 O.54 (10) ±O.Ol1.O6,s4c4O .O1.251,0 75"D] O,47±O,02 (10)1,92
±o.es1.1a.26 71.93 CIO )1.95±O.06t.B3 2.03 (le)1,71±O,07 t.62 1.82 (10 )1,95±O,151.56 2.15 (10) 1.53 1.95 (ID ]1.B2±D.10 (10)1.73
±O.182.17.44 12.D7 C5e }1.46ro.061.36 1.57 (50)2.le±O.19 1.88 2.59C20 )1.45±O,081,32 1.58 (20) ±O,142,1S,457 2,D2 (2D }t.6ltO.09 (20)3.07
±O.232.822,44 3.15 (50 }2.41±D.07 2.43 2.54 (50)2.62±O.27 2.34 3.33(2D }2.55±O.172.27 2,7e (20) ±O,193.21,758 3,14 {2D }2,78±O,16 (20)3.13
±O,392.19.39 63.35 (50 }2.87±O,212.35 3.82 (50)2,59±O,24 2,35 3.09 (2 0)2.54±O.272.09 2.96 (20) ±O.3922..6789 3,91 {20 )2.69±D25 (20)3.18
±O.29os2g.58 3.57 (50 )2.79±O.162Al 3.07 C50}2.91±O,15 2,74 3.25 (20 )2.86±O,162.58 3.19 (20) ±O.ri72.14.81 13,OB t20 )2.SUO.tS (20)1,19
±e.loO.O5.385 1.17 {50 )ri.4±7OA2 O.B7 1.28 (50}e.95mo.1 2O.S2 1.25 {20 )O.9D±D.09D.74 1.02 <20) ±O.09O1.,5O8 l1,2 8C20 )1.03±O.OB (20}O,63
±O,0321O..947 O.56 {50 )O.53±O,03O.47 O,59 (50)O,52±O,04 O,46 O.60(20 )O.51±O.030.45 O.56 <20) ±O.02tO9..595 O.6 1(20 )O.5eee.04 {20}
±1.616. 716 tB {10) 20,±31,3 19 24 (10)C20)CIO)(20)(10)CIO)(±110.)B1(51. 60r)i (71203} <(1100))
±1,O 13 20 (50) 15.3±1.4 12 18 ±1.0 13 17 (2e)
5.6±O,5 5 6 (10) 6.1±O.7 5 8 6.9±O.710. 1 6 8 (le)
92±1.0 7 12 C50) 9,1±1.0 8 12 ±1.1 7 t2 C20)
3.5±O.5 3 4 (tO} 3.8±O.4 3 4 3.7±O.5 3 4 (10)
7.or:O,813. I M.65 8 (le) e.3±O.6 8 le B.5±O.715. t 8 ID OO}
12 14 (10) 13,1±O.9 12 15 ±IJ 12 t7 (10)
3,OtO.O 3 3 {10) 3.eto.o 3 3 3,e±o.o 3 3(10)
2.0±O.O 2 2 {10) 2.osO.O 2 2 2.0±O.O 2 2(10)
7.2±O.7 6 S CIO) 7,atO. 6 6 8 7.2iO.6 6 9 {10)14.8
t3.7±t.4 11 15 {10) 13.8±t.2 12 16 CIO) ±O.6 14 16 (10)
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TThheZeo olZoogiocallogical SSoociceityety ooff JJaapapnan
1382 M,T. Zannatand M. J. Toda
posterjo sretae. apjcal seta on tibia of fore -and midlegs. Longitudin aslex-
Thorax :Scutum and thoracic pleur abrownish ye-ow. combs presen talong entire lengths of lst and 2nd tarso-
Acrostichal setulae in 6 rows in fron tof anterior dorsocentral meres ot 8 foreleg.
setae, 4 rows betwee ndorsocentr asletae. Basal scutellar Wing transparent, slightly yellowis hV,eins brownish
setae paralle lap;ical setae cruciate at right angle. yellow .Basal medial-cubital (bm-c ucr)o$$vein absent,
Legs: Preapica ldorsa lseta presen ton tibi aof all legs; Abdomen: Second to 6th tergit eysellow ,each with very
distjn captical black band, except for 8 6th black dorsally.
Twiatbht et h4e .t hoQruaax nlteintgatt hii vn cethharea cttherrsee s isbhloiwnign gspe ciseisgnifican tcorrelations p u beMsacleen tt ,weirtmhi tnrailai naEg:pual naedrxprainusimo npa lceo vberriongwn , bbarsoe aodf ,snuort-
Species:Stra i nl D. tini D. ohnishii D. ogurnai stylus; ventral lobe apically round. Cercus separated from
BG14028-0581,O MMY326(PvalueR)GN206(Pvalue) epandrium, not pubescent ,triangul alro:wer ,oval part sepa-
(Pvalue) rated, differentia taesd secondary ciasper. Hypandrium
Male pubescent on caudolateral lobes and caudal margin, caudo-
medially with strong protrusi oanpically bearin ga pair of
spcs O.0460O.0292O.OO0 9tnsO.0421 ns stout paramedian setae as long as paramere, Paramere
ps ns large ,longe rthan wide, Aedeagus slender, apically finely
FWIHW ns nsO.0249O.0353 hirsut eb,asall ywith a pair of lobat eprocesses as long as
vb ns ns aedeagu$, fused to apodeme; apodeme rod-like, as long as
sctl nsO.0482 ns aedeagu$,
M nsO.0471 ns
chloFemale ns ns TKarbulsek a6l.- WaQlula itnsetsitt )aamtoingv echa trhace ttehrsre e wsiitbhli ngsi gsnpiefciiceasnt differenc e(by
Character Male{Pvalue) Female(Pvalue)
ocps O.0388 nsO.O076O.0261 nsnsnsnsns
prorb ns
spcsocpspsFWIHWchlOprorbr crOo.bOd0c8ls3c<tOl.$OteOrOnIotodeitos cOtl.pOWWOA0N3Lc'4c4v5xacMC3Fsclsc2epandsvlbssurspcers
dcp nsO,0432O.0251
O.OOOIt
sterno2 ns
<O.OOOIt <O.OOOI+
ovms ns
' Signific aafntter Bonferron icorrection. O.OOIOt ns
ns O.O064
Table 5. Quantitati vceharacters with significant sexual difference O.O042 ns
(b yunpaired Fte$t )in the three sibling species O.OO03' O,OO02t
SpecieslStra i n: D. Iini D. ohnishii D. ogumai O.0458 ns
BG14028-0581.0 MMY326(PvalueR)GN206(Pvalue) ns O,O156
(Pvalue} O.OI04<O.OOOIi O,0482<O.OOOI.
spcsocpspsmsFWtHWprOor,b0r2c8o5rbOs,tOeOrInoOs'terno2orOb.iOt0o3dc0pOc.4Oc0ac256xCO3.FO027O,0378O.O186O.O047O,OIOOO.OO15i
O,O098 <O.OOOI'
ns ns ns O.O121
nsO.O095O.OOIO' ns O.O085 O.O093
nsO.OO02tO.0490 O.O026 O.OO02+
O,O065 O,OO03t
nsO.0287 ns O.OO06t O,O066<O.OOOIt
ns nsO.0313 <O,OOOIt
nsO.OO07'nsO.O021O.0438 ns<O.OOOIt O,OOIIt <O.OOOI'
ns O.O020' O.OO03'
O.0265O,OO05t O,nsOO0.042130O.O122 O.O147<O,OOOI'
ns O.O041<O.OOOIt
ns nsO.0222
nsO.O190 fis O.O021
ns ns O.O167
' Signific aafntter Bonferro cnoirrection. 'SignificantafterBonterronicorrectjon.
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Two New Sibling sof Drosophil aha i 1383
Eipecimen sexamined. Ten 8 and 1O 9 each from the Comparison of quantitati vceharacters
followi ngisofemal estrains :BG14028-0581.0, DHS401, The mean ±SD and the range are shown to r42 quanti-
DHS501, NKS9212 and NKS9231, tativ echaracters, separately for each sex and fo reach spe-
Djstributio nC.hina (Taiwa Gnu,angdong). cies, in Table 3. In interspeci fciomcparison for each sex,
there is no character with nonoverlapping ranges between
Drosophit (aSophophor oahn)ishii sp. nov. species, which can be used as the specific diagnosis,
(Fig sIB., 2D-F) Using the data fo rthe three strains ,BG14028-0581.0
Drosophla (Sophopho riai)ni-l ikOeh,nish i& Watanabe, (D .LinD ,MMY326 (D .ohnishiO and RGN206 (D ,ogumaD,
1984: 802; Kim et al, ,1989: 178; 1993: 992; Oguma et al,, which were measured tor all the 42 quantitat cihvareacters,
1995: 312. correlatiens with the body size (ThL t:horax length) were
Diagnosis ,Paramere apically round and with 3-5 examined for 38 characters. Seven characters for male and
minute sensilla, submedially with moderate, inward expan- fiv echaracters for female showed significant correlations
sion of which posteri moarrgin distinc tclonycave (Fi g2D.,F}. with ThL in any of the three species (Tab l4e), However,
Aedeagal basal proces swith fine ,irregul asrerrations on
narrower, apical margin than in D. tin i(Fi g2.E),
Holotype 8 from MMY326, 5
Paraopes. Nine 8 and 10 9 from MMY326; 10 {7i and Male
tO \ fromMMY307. 4
DistTtbutioMny.anmar (Pyinoolwin),
3 A
Relationship Tsh.i$ species is very closely related to D. A A
tin rie:ciprocal crosses between these two species can pro- 2nytft o AAA AA b
duce Ft hybrids o,f whjch female sare ferti lbeut maies ster- A
il e(Oguma et aL, 1995) .Morphologically, these two species tt.! cooeootsAAOAooo oAoAo
can hardl ybe distinguish efdrom each other even by the oA aouDa
cshiasr. aAct ecrosmbi noaf tmiaolne toef rsmoimenal qiwuhaiachnti atraet ig vcihevaeran citne trshe dciaang nboe- .o2,8 oooooo ooe eooDooooo oSo o oDp DDDDD
used, but not absolutely, to discrimina ttehese two species ¢ uDD
(seebelow). .2
o
EtymologJ LPatronym, in honor of Dr. S. Ohnishi who oeo
found thi$species first. -s
-4
DrosophiLa (Sophophor oagu)mai sp. nov. .4･3.2.1 O 1
(Fig sIC., 2G-1> Oanonical variable 123g5
Diagnosis .Paramere apically narrow and with 2-3
minute sensilla, submedially with moderate, inward expan- 5
sion of which posterio mrargin distinc tcloyncave (Fi g2G.,1). Female
Aedeagal basal process irregul aonr apical margin but with- 4
out fine serrations (Fig 2.H).
Holot)pe 8 from RGN206, 3
o
Rarat}tpe sN.lne cn and 1O 5 1from RGN206; 1O 84 and oooo D
10 9 fromRGN3. 2cu!ft o D
eoooo8oeegooo o ooooDo
DisttthutiOMny,anmar (Yangon).
ixlg,?28
Relationshi pTsh.is species is also closely related to the D
foregoin gtwo species: crosses with them produc eFi fertile oo o%ooBe o
coOooeooefiooAaboAV
females but sterile males, but crosses between ohnishii
female sand qgumai males produce no Fi hybrid s(Oguma ., -
et aL, 1995), Morphological ltyh,is species can be distin- o OA AAAAAAA
guished from the other two species by the diagnosti cchar- -2
acters, but only for males. Combination sof some quantitative Ab
-3 AAA
characters can be used, not only for males but also for
females ,to discrimina ttehi sspecies from the others (see -4
below), .4-3-2.1 O 1
Etymology Patronym, in honor of Dr, Y, Oguma who Canenica vlariable 12s45
detecte tdhe presence of postmatjn gisolati obnetween this Fig. 3. Scatterpl oott csanonicai scores for 90 specimens: 50 ot D.
species and the other two species for the fir sttime, Iin i(circ alned) 20 each of D. ohnishii (trian ganlde )D. ogumai
(squar ei)n each sex.
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only FW/HW in maie of BGI4028-0581,O held a significant orbital seta) was significantly differe nbtetween male and
correlation after Bonterron icorrection. Since no character female in all the three species. Theretore t,he followi nagnal-
was thus regarded a$ being affected constantly by the body yses were carried out separately for each $ex.
size, i.e .culture conditions, all the characters except for Kruskal-Wall itsest sdetecte dsigniticant interspecific
those representing the body size were subjected to the fel- differen cien 23 characters for male and 17 characters for
[owin ganalyses. female (Tab l6e) .Of 26 characters that differ esdignificantly
Significa snetxual difference was detecte dby unpaired among the species in either sex, 15 characters showed still
Ftest for 16 characters in any ot the thre especies (Tab l5e), signiticant interspeci fdiicfference after Bonferroni correc-
Among a tota lof 27 such cases were seven cases stil lsig- tion .These 15 characters were used in the canonical dis-
nifjcant after Bonferron ciorrection; especially, prorb (relativecriminant analysis.
lengt hof the proclina toerbital seta to the posteri orreclinate Fig. 3 shows plot sof canonical scores for measured
Table 7. Factor structure coefficients (correlat iboentwseen the characters and the discriminan ftunciions a)nd classification function sferthe
three sibling species, resulting from canonical discriminan atnalysis in which classification swere based on equal a prior pirobabili tfyor each
specles
Factorstructurecoefficients Classificationfunctions
Functio n1 Function2 D. Iinip=.33333D. ohnishii D. ogumai
Aprioriprobability: M.33333 p=.33333
Male
spcs O.266227 n.ooo47s 2.393 2.475 2.311
ocps O.550960 -O.104288 O.B87 1.226 2.466
ps O.174051 -O.149233 -1.7291282.87 0-1.6651332.12 4 -1.6551326.998
FWIHW O.140366 O.429669
rcorb O.203348 O.O16609 104.630 119.611 135.53S
orbito -O.137896 O.094232 69.357 67.346 61.627
sctlp 0243104 O.207933 25251 33.425 29.349
4c O.153670 -O.162753 29.403 30.957 41.292
4v O.111083 -O.296968 51.253 46,618 46.572
5x O.208429 -O.374703 O.255 n.272 3.056
ac -O.096502 O.O17271 -7.206 -7.671 -13.344
M O.195389 -O.358703 34.881 35.268 30.307
C3F O.311355 n.356125 182.338 164.375 197.085
-O.188396 -O.397294
sc2 17.412 16,226 16.264
vlbs O.168478 -O.13e057 7.261 6.767-677.275 7.661
Constant -679.703 -721.889
Female
spcs O.153029 O,435593 -1.106 -2.088 -1.520
ocps O.363089 O,076757 -O.834 n,318 O,040
ps O.161460 O.249951 -2.734 -2.890 -2,540
FWIHW O.297272 -O,326475 2527,289 2634,603 2582.174
rcorb O.313515 O.039127 54.713 68.337 80.395
orbito O.077671 -O.241863 124,O13 139,055 129.650
sctip O,499406 O.258703 158,399 163,628 IB3.733
4c O.317850 O.229277 -31.2S2 -26,295 -4.801
4v O,151690 O.263008 123.466 119,749 114.503
5x -O.117790 O.238431 50.220 46.883 48,991
ac O.041"7 -O,077814 95.241 98.971 95,038
M O.021024 O.261844 -157.073 -151.888 -160.064
C3F O.295669 O,391708 338.958 313277 343.224
Constant -1012.122 -1061.614 -1104.296
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Two New Siblin gofsDrosophil alin i 1385
Table 8. Summary of posterio rciassifications, Rows: observed $ex, and the 15 characters were used in male and 13 of
classifications c,olumns: predicte dclassifications them, excluding two male-specific characters, in female for
Correct(%)D.tini D,ohnishii D,ogumai the analyses. At a glance of the figur et,he tir sctanonical
Male functio ncan be regarded as discriminat iDn.g ogumai from
D. Iini 88.0 442147 616t23 o21820 tDh.e oohtnhiesrhi isp fecrioesm ,t haen dot htehrse ,s eicon nbdot hf suenxecs,ti Toahsne d ifsicrr sfitumnicntaitoinng
D. ohnishii 80.0 means large ocps (th neumber of occipital setae) and C3F
D. ogumai 90.0 (th erelative length of heavy setation in the third costal sec-
TotalFemale 86,7 tion )in both sexes, and larg esctlp <th edistanc ebetween
ipsilate rsacultella rsetae / the cross distanc ebetween api-
D. fini 90.0leo.4o5o247 520126 oo1717 csaelct sicount eltloa rM is etbaeet)w, e4ecn (r-tmh raeenldat idvem -lceun),g t rhocto rtbh e( tthhi err edcloasttiavle
D. ohnishii lengt hof the anterior reclinate orbitai seta to the posterior
D. ogumai 85.0 reclinate orbital seta> and FW/HW (th reelative width of frons
Total 91.1 to head width) in female (Tab l7e) .The second function
seems to be reverse in the directi obnetween male and
specimens, resu}ting from the canonical discriminant analy- female plots m,eaning larg eFW/HW, small sc2 (th enumber
ses, separately for each sex. A tota lof 90 individuals ,10 of teet hin sex-comb of forele g2nd tarsomere), 5x (th reel-
each from the nine strains (fi vsterains ef D. Iin iand two ative lengt hof CuAi between dm-cu and wing margin to dm-
each of D. ohnishii and D, ogumaD, were measured in each cu between Mi and CuAD, M (th reelativ elengt hof CuAi
Table9,Degrees of premating and postmatin gisolati obnased on the data of Oguma et al .(1995)
FemaleSpecies Male
D. Iini D. ogumai D. ohnishii
StrainTWN-DHS315DHS401DHS501NKS9231RGN3RGN206MMY307 MMY326
A) Copulati orante calculated from the data of pair-matin egxperiment
D.lini
rwN O.1O.4o.oO.1- O. 1 O,3 O.4O.2O.3O.2O.2O.5O,1OO.,32O,3o,o02o.oO.2O.1o.o
DHS401 - O,2 O,2
DHS501 - O,4 3,O
NKS923t - O.2 O.O
D. ogumai
RGN3 O.3O.1 - 02 O.2 o.oO.3 O,3O.4 O.3O.4 o.oo,o
RGN206 - O.3 O.2
D. ohnishii
MMY307 o.o - o.o o.o o.o o.o o.o O.4
B) Strengt hof postmati nisgolation
D.tini
rwNDHS315DHS401'DoH.So5o0.1oNoK.So9o.2oo3.1o o,oo.o o.oo.oo.o o.oo.oo.oo.oO.5O.5O.5OO..55OO..55O.5O.5OO..55O.5O.5OO,.55OO,.55O.5O.5O.5
o.oo.oo.o
o.oo.o
o.o
D. ogurnai
RGN3RGN206 O.5O.5 O.5O.5 O.5O.5 O.5O.5 O.5O,5 o.o O,5O,5 O.5O.5
o.o
D. ohnishii
MMY307MMY326 O.5O.5 O.5O.5 O,5O.5 O.5O.5 O,5O,5 1.01,O O,51,O o.o
o.o
tTWNIBG14028-0581,O
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between dm-cu and wing margin to Mi between r-m and cally good species reproductively isolate fdrom each other
dm-cu), C3F and 4v (th reelative length of Mi between dm- almost completeiy (Ki met al. ,1989; Oguma et aL, 1995). Of
cu and wing margin to Mi between r-m and dm-cu) in male, tho$e three species, two were de$cribe das new species,
but larg eC3F and small FW/HW in temale {Tab j7e) .Clas- i,e ,D. ohnishii and D. ogumai, and the remaining known
sificatio nfunction fsor the three species, based on equal a species, D. Iin iw,as redescribed in the Iigh tof detailed
prior pirobabili ftoyr each species, are shown in Table 7, and examination of the male terminal iaand many quantitative
the results ot posteriorclassifica tofi otnhes 90 cases (spec- characters, However, they are very similar in morphology,
imens) in each sex are shown in Table 8. The percentage hardl ydistinguisha bflreom each other even by a few char-
of correct classifications was rather high in both sexes, acters designate das the diagnosi se,specially between D.
86.7% in male and 91.1% in female. Thus, the present finian dD. ohnishii, implyin gthat they are stil lin the process
results suggest that the 15 (f omrale) or 13 (fo tremale) of species differentiati one,ven though havin g reached
quantitati vchearacters in combination are effective to di$- almost to the level of good species. This is supported atso
criminate the three sibling $pecies, even their females ,from by parti acrlossability and fertil iotf yFi hybrid sbetween
each other with considerable confidence. them (Ki met aL, 1989; Oguma et al, ,1995),
l tis conceived, but implicit lthya,t character differentia-
DISCUSSION tio ndoes not always proceed in parall ealmong different
biologic aplroperti eisn the process of speciation .Here ,mor-
ln thi$ study, we examined morphological differences phologic adlifferentia taimoonng the three sibling species is
among three sibling species that had proved to be biologi- compared with the degrees of premating and postmating
Morphology Reproductiveisolation
Male Premating
OH5rO Ll
OH O,902 Ll
O.433
t.ooo
OG
Female Postmating
OH 7.0 Ll OH O,500 Ll
11.8 .500
OG
OG
Fig. 4. Relationshi psamong the three sibling species (LlD:. Iin i,OH: D. ohnishii, OG: D. ogumaO in morphology (measur ebdy squared
Mahalanobis distance) p,remating isolati o{nisola tiindoenxl ,see text )and postmatin gisojati o{nse etexO, the las ttwo based on data by
Oguma et aL ( 1995).
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