Expansion of the Distribution Range and Pollinators of Canavalia rosea (Fabaceae) in the Ryukyu Archipelago, Japan

2004·0.53 MB·English

by  Matsumura, Shun-Ichi

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ISSN 1346-7565 Acta Phytotax ,Geobot.55 (3) :207-212 <2004)
Expansion of the DistributionRange and Pollinatorosf Canavalia
(Fabaceaien)theRyukyu Archipelago,Japan
rosea
SHUN-ICHI MATSUMURAi*, YOICHI TATEISHI2, YOKOYAMA3and

MASAYUKI MAKI3JUN
iBiological institut eG,raduate Sehool ofStien lcbeh,oku U}iiversi Soelndai ,Miyagi 980-8578 , 2College of
.]mpanr
Education,U)iivensoifttyhReyukyusN,lshiharaO,kinawa 903-O129, SDivision ofEcotog&:Elv?olutionao,
,lap)an;
BioloEp lGraduate Sbhool ofLij S2criences 7,bhoku U)iivensi4 .S;endoi ,Miyagi 980-8578 ,JLLpan
New localiti oefs Canavalia rosea were fbund on Okinawa and in the Amami Islands of the Ryukyu
Archipelag oT.he locali toyn Amami-Oshima marks the northern-most limi tof the species in Asia and
extends it srange approximately 490 km northeastward, At it snewly discovere ldocation sC,. iosea is
pollinat ebdy femal elea fcutter bees and bears frui tW.e suspect that the presence ofthree closely relat-
ed species of dnnavali awith flower msorphologically similar to those of C rosea is probably one of the
main reasons fbr thc visitation by leaf cutter bees to C, rosea in the Ryukyu Archipelago T.he female
bee sexhibited two pattern osfbehavier durin gfiowe rvisitations ancl effbctively pollinat etdhe fiowers
ofC rosea only when collecting both polle nand nectar. Pol1inator sofa iosea have been reported as
being anthopholid bees elsewhere, but are leafcutter bccs in thc Ryukyu Archipe]ago,
Key words: Canavali atvsea, leafcutt ebrees ,new distributi poonl,linato rRsy,ukyu Archipclago
The Ryukyus are a subtropical archipelago ofcon- 1993, Hatusima & Amano 1994) ,In this paper we
tinental island ssituated between [faiwa nand the report new tocalit iofes C. rosea from the northern
Japanes emainland. The vegetation is basicallyRyukyus and observations on reproductions in the
evergreen forest sof the Sino-Japanese Floristic species in the newly discovere hdabitats.
Region (Takhta j19a8n6). Nutnerous plant swith
their center of distribut iion nthe tropics have rep- New distribut ireocnotds
resentative in the archipelago. Canavali arosea Between 1995 and 2003 we fbund CZinavat iroasea
(Sw. D)C, (Papiliono iFdaebaacee,ae) i,s a typical at seven new localiti eCsYbnagu Knuim,e, Ikei,
plant oftropical seashores (Saue 1r964) with fruit Okinawa, Okjneerabu ,and in the Amami-Oshima
adapted to dispersal by ocean currents (RidleyIslands i)n the Ryukyu Archipe]ago (fab l1 ,e Fig. 1),
1930) .Previousl yit was known in Japan from only These discoveri eesxtend the distributi orannge
the Yaeyama and the Miyako Island sin thc southcrn approximately 490 km northeastward, with the
Ryukyu Archipelag o(Walk e1r976, Shimabuku Amami-Oshima Islan dpopulation sat 28"16'N,
*present address ;Graduate School of Engineering and Science,Universityof the Ryukyus,Senbaru, Nishihara,
Okinawa903-0213,Japan
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TABLE 1. New localiti oefs Canavalia ro.s'ea and sites where drif tseeds were collected in the Ryukyu Archipelago. Stage efplants is
as fbllow sFL;. floweri nFgR;, timitin gS;L, seedlings; DS, drif iseeds. For number of new localiti essec, figur e1.
lstand Locality Stages &)ecitneni Observationyears
TakaraKikaiAmaOmhig-omori DSDSFL. S.M.DS.2464 200220021999L9992001t998200220021999$1999*200]l996-2003

lke.iiUken S.M. DSH2701

Village A,shiken-Ikegach i(1) FRDSDSDSDSYD.S FTL, ,& T. K. s.n.
'I',
Oshima Uken VMage, Funakoshi Y. DS-796
Kasari Town, Domori Beach S.M. DS-g87

Naze City, Chinaze Y. T & T, Y. DS- t559
Naze City, Arira S, M, DS-27S2
IJI(eOkinoerabIukejiEasteni S,M, DS-2566

coast ofCape Hanzaki (2) FRFL. s. M. ggoee1
Naikina Beach <3) FRDSFL, s,M. ggoeo2

UjijiBeach S.M. DS-g51

Okinawa KunigamiVillage,Hentona(4) FR, SL Y, T. & al. 43517
NakljinVillage,Nagahama DSDSDSFL, Y.T& T.Y.DS-797 1999199519961999*1999-2001**

lc-jimaScsokoIkciKumeMiyako Y.TDS-1561

'1'
Westeim beach Y. & al. DS-1562
Easter nbeach (5) FRFL, S. M, 990003
Shimajir i{6) FRDSDSDSp S. M, s.n,
Bora Peit, Cape Higashi-hcnna S. M, DS-800 19992001199919961996199619991996.
DittoWbeach,CapcIIigashi-henna S.M, DS-2042

FLp FR s. M, Dg-sot
lshigaki Cape Hirakubo DS, FL, FR S. M, DS-1571
AkaishilnodaYoshihura DSDSDSDS, S.M. DS-1563

S.M. DS-1568,1569

s.M. Ds-go2

TodorokRiiver FL.FR S.M. DS.803,1172 1997t

ShitaheHigawa DS, FL,FR S.M, DS-1564 -IS66 1994.1996'l'

Yonaguni Beach (7) FL,FR S. M. s,n. 1995*
1 Abbreviatien sofcollecter naines are as t'ollow sY:, T. = Y. Tateis hTi, ,K ' , T, Kurosawa, T. Y. = T. Yamashiro,
S. M, = S. Matsumura.
'1'hese
" population swcre absent in 1996 on Yonaguni Island ,andin 2001 on Okinoerabu and Ikei islands.
"" We observed fiuiting plants only in 1999,
'1'
Recerds are years c)fobservation ofdrift seeds, Although we observec{ flewerin gand fuiting plant sat a few localiti oens
Ishigak iIsland, years wcrc not recorded becau$e the plant wsere common.
129015'E being the northeinmost limi otf the species the Okinawa site were evidence of regencration.
in Asia .The populations on YOnaguni, Ikei ,and We did not find fi:ui itns the population on Kume
Okinoerabu islands have been lost since we dis- Island in either 2000 or 2001, but was sa"r frui tansd
cevered them. At ali newly discovere dsites C. a pollinato irn 1 999. W ℃ presume that the plant swe
iosea was a trailin gplan ton beache snext to the sea. fbund originated firo mseeds canied by oceanic cur-
The plant sbore firui tats all sites. We observed rents. We collected drif tseeds of C ivsea on the
fiui tisn the Hentona population on Okinawa Island coasts ofOkinoerabu in the Amami-Oshima islands
¢very year from 1996 to 2003 and in the Shimajiri and at other place s(Tab l1e, Fig, 1).
populatio onn Kume Is]and in 1999. Seedling sat
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t
Canavalia (Sw.D)C. 'l'
rosea
':"
o?
e locality
new
.t.-.-
A driftseeds

k
- Amami lslands1 Ammi-ohshima
200 km
28"N

' -t
p .J.J
-
.
ino au
Okin
" wa
t-p.t".-t
Ku

. e'
-
;... -.- , -..t.-."NNt
N
ds
2
'zi '
N
N-e..----- --`"'2-i4oN
.. ,,,!lilii$X,\af'
t -T-a'i-wan -- ------- -- --- ---- -- -- -------- d T -pt i-c -o-f- Cance -r--
"
'
"
1220E 126eE 1300E
'
FIG. i . New loca[it ioefs Canm,aiia rvsea and sitcs whcrc drif steeds were collected in the Ryukyu Archipeiag o.Broken aiid thin lines
show previou sand ncw northern ]imi tof$distribution, respectively. For number ofnew localiti esese, Table 1,
Reproduetion in the new localities three sites and by Chaticodoma monticola (Smith)
Although thcre were relatively few individual sat at one sitc (fab l2e, Fig. 2).
each site, ftuit wsere well-developed at alS of the In addition to Canavalia rosea. two other
sites we djscevere d.Because Canavali arosea is species of Ckenavalia ,C. Iineat a(Thunb D.C). and
selflincompatiblc (Gros s1993) and because the C. cathartica Thouars, are also distribute din the
stigma and anthers are hidden by the two kecl RyukyL iArchipelag oB,oth are closely relatcd to C.
petals ,the fiowe rhas to be pollinate dby a vector tvsea (Saue r1964), and are mainly pollinat ebdy
tha thas the ability te forc eopen the petal (sFacg &ri leafcutt ebrees and carpenter bees (Kat o2000, S.
van der Pij 11979). We observed that C. ivsea was Matsumura et al. unpubLished data) , Except for
pollinate dby femaies of the lcaf cutter bees the carpenter bees, thc pollinat oorfs C. Iineat aand
ILdegachi ioekinawana Yasumatsu & Hirashiin aat C. cathartica are the same as those of C. rosea,
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TABLE 2. Flower visitors of thnavalia rosea in the Ryukyii Archipela agnod un Lanyu Island f,aiwan.Numbers of flowe rvisitors arc
shown in parenthesis.
Flower visitors
Megachilidae (le acuftter bec)chi'71c5demaAnthophoridae
Island Mbgachiie bicogo rgroup other Megaehite Xl,loeopa Date
Amami-
Oshima NQobservation
Okinoerabu M okinawana Oct99
(1 ?)
Okinawa M. okinan'ana C. monticola 20.Jun.97
(3<g) (2\)
M okinc-vana C. monticola 27Jul,97
(1`.-4) (3<4)
IkcjKume No observation*
M, okinawana 1.Sep.99
a4)
Miyako it4/ yae.vantaensi,y il4. igni,y('opatu 24, 25 June 1999
(3<9, 1< gi) e9)
lshigaki ,Lt vaey'amae}tvis 17.Aug.99
(s<\, 3< db
Lanyu I. i'qficepb' 10.Sep.98
( 1g)
' Matsumura observed the holes on the standard, which the leafcutte brees mudc by chewing with the mandibles,
in 27 Aug.1999
The rnoxphological feature osf fiewers of the three upside down, They remain inverte dwhile bitin tghe
species of Canavali aare quit esimilar elsewhere basa lpart ofthe standard with their mandibles and
(Saue 1r964 ,Stirto n1977, Gottsberge r1988) ,as while grippin gthe wing petals .They then fbrce
they are in the Ryukyus. The presence ofthe three open the keel!win pgetal complex and actively col-
closely related species with similar fiower sis prob- lect pollen with their lcgs . The polle nadheres
ably one of thc main reasons fbr visitations to C. mainly to the scopae and legs and is late dreposited
rosea by leaf cutter bees in spitc their small popu- on the stigma of other fiowers (Fi g2.) .They there-
latio snizes. fbr eact as pollinator sonly when collecting both
pollen and nectar.
Behavior ofpollinators Male leafcutt ebrees do not effl]ct pollination
The female leaf cutter exhibit two pattern sofbehay- in spite oftheir fireque fnltower visits; they collect
ior when they visit flo",e rofs CZinavat iivasea. Tb only nectar and never come in contact with the
collect nectar, they alight on the standard petal and stigma. Other fiower visitors were also observed;
press their head to the keeVwing petal complex CZrratin aokinawana Matsumura & Uchida (Antho-
while sucking ncctaz In doing this ,they do not con- phoridae) ,Campsomerielta annulata Fabricius
tact the stigma, because their body is at a distance (Scolii danade )a ,fe" ispccies ofLepidoptera. We
from the keelfwing petal complex. When they col- doubt their effectiveness as pollinato rhso,wever,
lec tboth polle nand nectar the bees alight on the because Ce. okinawana appeared only to rob the
standard fir satnd then quickl yturn their bodies pollen and the other visitors never came in contact
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RG. 2. Fernale leaf cutter bee, Mlegaahile okinavvana, visitin fgluwer sof Cattavali raosea at Hentona, Oknawa Island B.ee pushes down
keelfwing petal complex with legs while bitin gstandard. Ivhntra lsurface of abdomeri ofbee covered by yellow polle inn contaet
with pisti l(whit aerrow). Legs ofbee in contact with anthers near abdernen. TXvo bite marks on basal part of standard (black
arrow). Scale bar = 1.5 cm.
with the stigma. Those flowe rvisitors are nectar the Ryukyu Archipelag (oYlima ent eal. 1999) ,and
andlor polle nrobbers, belong to the Ac tbicolo rgroup (Yasumat s&u
Other anthophorid bees have been reported Hirashima 1964).
previousl ays pollinato rofs CZxnavali arosea, for Pollinatio nby leaf cutter bees is rare in the
example, X),locqpa]ho (nOtlailviis eXlr o)rt,hogo- genus CZinavali a.Only Stirto n(197 7re)ported
naspis Moure, Centri sfuviat iFrliiesse ,Centris tha tC, virosa (Roxb W.i)ght & Arn. was pollinated
sp., and Acanthqpus patmatus (eliv iien Brr)azil by the lea fcutter bee, Megachil eeombusla Smith ,in
(Gottsbe retg ael.r 1988), X violacea L, in South South Aftica ,with the main pollinat boering a car-
Afric a(Sco Etltlio 1t891) andX latip e(sDrur aynd) pente rbee ,Jfylocqpafavoxtij Drea Greer .Mbgachile
X conjiisa Perez in Indonesia (Gros s1993). combusta was an opportunist and did not always
Additionally w,e observed CL rosea being pollinat- succeed in pollinatin tghe fiower sof C, virosa. Our
ed by X[ rtgliceps Fries eon Lanyu Islan dsoutheast of observations indica tteha tthe pollinato rosf C rosea
Tlaiwan (Tab l2)e. Cbnavalia rosea was mostly pol- have switched from being anthophorid bees to leaf
linate bdy ?Lfegachi yiaeayamaensis Yasumatsu & cutter bees in the Ryukyu Archipelago ,even though
Hirashima a,nd rarely by ?tt i[gniscopata Cockerell thre eendemic species oflfylocopa (Anthophoridae)
in the southem Ryukyu Islands ([I} L1b)l .eBoth occur there (Yaman eet aL 1999) .We do not know
M oldnawana and M yaayamensis are endemic to why carpenter bees do not visit the flower sof C.
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rosea. It may be that leaf cutter bees are relatively Society ofOlcinawa, Okinawa. (I nJapanese)
more predominant in the Ryukyu Archipelago and Kato, M. 2000. Anthophirous insect community and
their abundance may reduce flower visitation by plant-inse citnteractio onns Arnamj Islands in the
carpenter bees through resource competition. Ryukyu Archipelag oJ,apan .Contr .Biol. Lal) .Kyoto
Univ.29]157-252.

Ridley, H. N. 1930. The Dispersa olfPlants throughou tthe
We are gratefu t1o two anonymous reviewers, Drs, H. Wbrld.L.Reeve& Co,,Ltd,,Kent.

Ohashi,T.Nemoto,M. Kato,J.OkazalciA,,Naiki,M,
Sauer J, .1964. Revisio nof Cbncivali Bar.itton i1a6: 106-
Hotta, T. Kajita, T, Kurosawa, Y/ Iwokawa and )rkr sN,. 181,

Nikaidoh for providing usefu1 advice, and Dr. C. H. Scot tEllio tG,. E 1891. Notcs on the fenilisati oonf
Stirto fnor sending us reprints of his papers .We also Sout hAfrican and Madagascar fiowerin gplants.
thankDr.T.Yamashiro,Mses,K.Kurima,YLMukon M.
Ann. Bot. 5: 333-405,
Kamizato and E. Tfamaki for their help in the field and in Shirnabuku, K. 1993, Distributio nof leguminous plant isn
thelaboratory.
the Ryulcyus. lsland Studies in Okinawa 11: 1-13. (in
Japanese)

References
Stinon ,C. H. 1977. The pollinati oofn Cdnavalia virosa
by xylocopid and megachilid bees .Bothali a12: 225-
Faegri, K. & L. van der Pij1 .1979, The Principle osf 227.
Pellination Ecology, 3rd editien. Pergarno nPress, kkhtajan A. 1986. Florist iRcegions of the World,
Oxford. Universit oyf Califbrni Paress ,Los Angeles,
Gettsberge rG,,, J. M, E Camargo & I, Silberbauer- Walker ,E, H. 1976. Flora of Okinawa and the Southern
Gottsberge r1.988. A bee-pollinat etrdopica lcom- Ryukyu Islands .Smithsenian Institution Press,
munity: the beach dune vegetation ofIlha de S5o Washington, D,C.
Lu{s, Maranhao, Brazil B.ot. Jahrb ,Syst ,109i 469- Yamane, S., S. Ikudome & M. Terayama. 1999.
500, Identificat iGeunide te the Aculeat aof the Nansei
Gross C,. L. 1993 .1[he rqproductive ecology of Cbnavalia Islands ,Japan .Hol(kaid oUhiversit Pyress ,Sapporo.
rosea (Fabace aoen ) Anak Krakatau, Indonesia. Yhsumatsu, K & Y Hirashima. 1964. Red-haire ldeafcut-
Austra lJ.. Bot. 41: 591-599. ter bees of the group of A4bgachile bicolor from
Hatusima ,S. & T. Amano. 1994, Flora ofthe Ryukyus, Japan ,the RyukyiJs and Formosa. Kontyu 32: 175-
South of Amami Islan d2.nd edition. The Bielogical 187.
Received Jiin ela 2004 ;accepted Septemher 28, 2004
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